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+Leonardo da Vinci (English: /ˌliːəˈnɑːrdoʊ də ˈvɪntʃi, ˌliːoʊˈ-, ˌleɪoʊˈ-/;[a] 14/15 April 1452[b] – 2 May 1519),[4] was an Italian polymath of the High Renaissance who is widely considered one of the greatest painters of all time (despite less than 25 of his paintings having survived).[c] The Mona Lisa is the most famous of Leonardo's works and the most famous portrait ever made.[5] The Last Supper is the most reproduced religious painting of all time[6] and his Vitruvian Man drawing is also regarded as a cultural icon.[7] He is also known for his his notebooks, in which he made drawings and notes on science and invention; these involve a variety of subjects including anatomy, cartography, painting, and paleontology. Leonardo's collective works compose a contribution to later generations of artists rivalled only by that of his contemporary Michelangelo.[8]
+Properly named Leonardo di ser Piero da Vinci,[d] Leonardo was born out of wedlock to a notary, Piero da Vinci, and a peasant woman, Caterina, in Vinci, in the region of Florence, Italy. Leonardo was educated in the studio of the renowned Italian painter Andrea del Verrocchio. Much of his earlier working life was spent in the service of Ludovico il Moro in Milan, and he later worked in Rome, Bologna and Venice. He spent his last three years in France, where he died in 1519.
+Although he had no formal academic training,[9] many historians and scholars regard Leonardo as the prime exemplar of the "Universal Genius" or "Renaissance Man", an individual of "unquenchable curiosity" and "feverishly inventive imagination."[6] He is widely considered one of the most diversely talented individuals ever to have lived.[10] According to art historian Helen Gardner, the scope and depth of his interests were without precedent in recorded history, and "his mind and personality seem to us superhuman, while the man himself mysterious and remote."[6] Scholars interpret his view of the world as being based in logic, though the empirical methods he used were unorthodox for his time.[11]
+Leonardo is revered for his technological ingenuity. He conceptualized flying machines, a type of armoured fighting vehicle, concentrated solar power, an adding machine,[12] and the double hull. Relatively few of his designs were constructed or even feasible during his lifetime, as the modern scientific approaches to metallurgy and engineering were only in their infancy during the Renaissance. Some of his smaller inventions, however, entered the world of manufacturing unheralded, such as an automated bobbin winder and a machine for testing the tensile strength of wire. He is also sometimes credited with the inventions of the parachute, helicopter, and tank.[13][14] He made substantial discoveries in anatomy, civil engineering, geology, optics, and hydrodynamics, but he did not publish his findings and they had little to no direct influence on subsequent science.[15]
+Leonardo was born on 14/15 April 1452[b] in the Tuscan hill town of Vinci, in the lower valley of the Arno river in the territory of the Medici-ruled Republic of Florence.[18] He was the out-of-wedlock son of Messer Piero Fruosino di Antonio da Vinci, a wealthy Florentine legal notary, and a peasant named Caterina,[e] identified as Caterina Buti del Vacca and more recently as Caterina di Meo Lippi by historian Martin Kemp. There have been many theories regarding Leonardo's mother's identity, including that she was a slave of foreign origin or an impoverished local youth.[17][20][21][22][f] Leonardo had no surname in the modern sense—da Vinci simply meaning "of Vinci"; his full birth name was Lionardo di ser Piero da Vinci,[2][25] meaning "Leonardo, (son) of ser Piero (from) Vinci."[18][d]
+Leonardo spent his first years in the hamlet of Anchiano in the home of his mother, and from at least 1457 lived in the household of his father, grandparents and uncle in the small town of Vinci.[26] His father had married a 16-year-old girl named Albiera Amadori, who loved Leonardo but died young[27] in 1465 without children. In 1468, when Leonardo was 16, his father married again to 20-year-old Francesca Lanfredini, who also died without children. Piero's legitimate heirs were born from his third wife Margherita di Guglielmo, who gave birth to six children, and his fourth and final wife, Lucrezia Cortigiani, who bore him another six heirs.[28][29]
+In all, Leonardo had 12 half-siblings, who were much younger than he was (the last was born when Leonardo was 40 years old) and with whom he had very little contact.[g]
+Leonardo received an informal education in Latin, geometry and mathematics. In later life, Leonardo recorded few distinct childhood incidents. One was of a kite coming to his cradle and opening his mouth with its tail; he regarded this as an omen of his writing on the subject.[31][32] The second occurred while he was exploring in the mountains: he discovered a cave and was both terrified that some great monster might lurk there and driven by curiosity to find out what was inside.[27] He also seems to have remembered some of his childhood observations of water, writing and crossing out the name of his hometown in one of his notebooks on the formation of rivers.[26]
+Leonardo's early life has been the subject of historical conjecture.[33] Vasari, the 16th-century biographer of Renaissance painters, tells a story of Leonardo as a very young man: A local peasant made himself a round shield and requested that Ser Piero have it painted for him. Leonardo, inspired by the story of Medusa, responded with a painting of a monster spitting fire that was so terrifying that his father bought a different shield to give to the peasant and sold Leonardo's to a Florentine art dealer for 100 ducats, who in turn sold it to the Duke of Milan.[34]
+In the mid-1460s, Leonardo's family moved to Florence, and around the age of 14,[30] he became a garzone (studio boy) in the workshop of Verrocchio, who was the leading Florentine painter and sculptor of his time.[35] Leonardo became an apprentice by the age of 17 and remained in training for seven years.[36][37] Other famous painters apprenticed in the workshop or associated with it include Ghirlandaio, Perugino, Botticelli, and Lorenzo di Credi.[27][38] Leonardo was exposed to both theoretical training and a wide range of technical skills,[39] including drafting, chemistry, metallurgy, metal working, plaster casting, leather working, mechanics, and wood-work, as well as the artistic skills of drawing, painting, sculpting, and modelling.[40][h]
+Much of the painting in Verrocchio's workshop was done by his employees. According to Vasari, Leonardo collaborated with Verrocchio on his The Baptism of Christ, painting the young angel holding Jesus' robe in a manner that was so far superior to his master's that Verrocchio put down his brush and never painted again, although this is believed to be an apocryphal story.[41] Close examination reveals areas of the work that have been painted or touched-up over the tempera, using the new technique of oil paint, including the landscape, the rocks seen through the brown mountain stream, and much of the figure of Jesus, bearing witness to the hand of Leonardo.[42] Leonardo may have been the model for two works by Verrocchio: the bronze statue of David in the Bargello, and the Archangel Raphael in Tobias and the Angel.[20]
+By 1472, at the age of 20, Leonardo qualified as a master in the Guild of Saint Luke, the guild of artists and doctors of medicine,[i] but even after his father set him up in his own workshop, his attachment to Verrocchio was such that he continued to collaborate and live with him.[27][43] Leonardo's earliest known dated work is a 1473 pen-and-ink drawing of the Arno valley,[38] which has been cited as the first "pure" landscape in the Occident.[j][44] According to Vasari, the young Leonardo was the first to suggest making the Arno river a navigable channel between Florence and Pisa.[45]
+In January 1478, Leonardo received an independent commission to paint an altarpiece for the Chapel of St. Bernard in the Palazzo Vecchio,[46] an indication of his independence from Verrocchio's studio. One anonymous writer claims that in 1480, Leonardo was living with the Medici and often worked in the garden of the Piazza San Marco, Florence, where a Neoplatonic academy of artists, poets and philosophers organized by the Medici met.[20] In March 1481, he received a commission from the monks of San Donato in Scopeto for The Adoration of the Magi.[47]
+Neither of these initial commissions were completed, being abandoned when Leonardo went to offer his services to Duke of Milan Ludovico Sforza. In 1482, he casted a silver stringed instrument[k] from a horse's skull and ram horns to bring to Sforza,[48] whom he wrote a letter describing the diverse things that he could achieve in the fields of engineering and weapon design, and mentioning that he could paint.[38][48][49]
+Leonardo worked in Milan from 1482 until 1499. He was commissioned to paint the Virgin of the Rocks for the Confraternity of the Immaculate Conception and The Last Supper for the monastery of Santa Maria delle Grazie.[51] In the spring of 1485, Leonardo travelled to Hungary on behalf of Sforza to meet king Matthias Corvinus, and was commissioned by him to paint a Madonna.[52]
+Leonardo was employed on many other projects for Sforza, including the preparation of floats and pageants for special occasions, a drawing and wooden model for a competition to design the cupola for Milan Cathedral (which he withdrew),[53] and a model for a huge equestrian monument to Ludovico's predecessor Francesco Sforza. This would have surpassed in size the only two large equestrian statues of the Renaissance, Donatello's Gattamelata in Padua and Verrocchio's Bartolomeo Colleoni in Venice, and became known as the Gran Cavallo.[38] Leonardo completed a model for the horse and made detailed plans for its casting,[38] but in November 1494, Ludovico gave the bronze to his brother-in-law to be used for a cannon to defend the city from Charles VIII.[38]
+With Ludovico Sforza overthrown at the dawn of the Second Italian War, Leonardo, with his assistant Salaì and friend, the mathematician Luca Pacioli, fled Milan for Venice.[54] There, he was employed as a military architect and engineer, devising methods to defend the city from naval attack.[27] On his return to Florence in 1500, he and his household were guests of the Servite monks at the monastery of Santissima Annunziata and were provided with a workshop where, according to Vasari, Leonardo created the cartoon of The Virgin and Child with St Anne and St John the Baptist, a work that won such admiration that "men and women, young and old" flocked to see it "as if they were attending a great festival."[55][l]
+In Cesena in 1502, Leonardo entered the service of Cesare Borgia, the son of Pope Alexander VI, acting as a military architect and engineer and travelling throughout Italy with his patron.[54] Leonardo created a map of Cesare Borgia's stronghold, a town plan of Imola in order to win his patronage. Maps were extremely rare at the time and it would have seemed like a new concept. Upon seeing it, Cesare hired Leonardo as his chief military engineer and architect. Later in the year, Leonardo produced another map for his patron, one of Chiana Valley, Tuscany, so as to give his patron a better overlay of the land and greater strategic position. He created this map in conjunction with his other project of constructing a dam from the sea to Florence, in order to allow a supply of water to sustain the canal during all seasons.
+Leonardo had left Borgia's service and returned to Florence by early 1503,[57] where he rejoined the Guild of Saint Luke on 18 October of that year. By this same month, Leonardo had begun working on a portrait of Lisa del Giocondo, the model for the Mona Lisa,[58][59] which he would continue working on until his twilight years. In January 1504, he was part of a committee formed to recommend where Michelangelo's statue of David should be placed.[60] He then spent two years in Florence designing and painting a mural of The Battle of Anghiari for the Signoria,[54] with Michelangelo designing its companion piece, The Battle of Cascina.[m]
+In 1506, Leonardo was summoned to Milan by Charles II d'Amboise, the acting French governor of the city. The Council of Florence wished Leonardo to return promptly to finish The Battle of Anghiari, but he was given leave at the behest of Louis XII, who considered commissioning the artist to make some portraits.[63] Leonardo may have commenced a project for an equestrian figure of d'Amboise;[64] a wax model survives and, if genuine, is the only extant example of Leonardo's sculpture. Leonardo was otherwise free to pursue his scientific interests.[63] Many of Leonardo's most prominent pupils either knew or worked with him in Milan,[27] including Bernardino Luini, Giovanni Antonio Boltraffio, and Marco d'Oggiono.
+In 1507, Leonardo was in Florence sorting out a dispute with his brothers over the estate of his father, who had died in 1504. By 1508, Leonardo was back in Milan, living in his own house in Porta Orientale in the parish of Santa Babila.[65]
+In 1512, Leonardo was working on plans for an equestrian monument for Gian Giacomo Trivulzio, but this was prevented by an invasion of a confederation of Swiss, Spanish and Venetian forces, which drove the French from Milan. Leonardo stayed in the city, spending several months in 1513 at the Medici's Vaprio d'Adda villa.[67] In March of that year, Lorenzo de' Medici's son Giovanni assumed the papacy (as Leo X); Leonardo went to Rome that September, where he was received by the pope's brother Giuliano.[67] From September 1513 to 1516, Leonardo spent much of his time living in the Belvedere Courtyard in the Apostolic Palace, where Michelangelo and Raphael were both active.[65] Leonardo was given an allowance of 33 ducats a month, and according to Vasari, decorated a lizard with scales dipped in quicksilver.[68] The pope gave him a painting commission of unknown subject matter, but cancelled it when the artist set about developing a new kind of varnish.[68][n] Leonardo became ill, in what may have been the first of multiple strokes leading to his death.[68] He practiced botany in the Gardens of Vatican City, and was commissioned to make plans for the pope's proposed draining of the Pontine Marshes.[69] He also dissected cadavers, making notes for a treatise on vocal cords;[70] these he gave to an official in hopes of regaining the pope's favor, but was unsuccessful.[68]
+In October 1515, King Francis I of France recaptured Milan.[47] Leonardo was present at the 19 December meeting of Francis I and Leo X, which took place in Bologna.[27][71][72] In 1516, Leonardo entered Francis' service, being given the use of the manor house Clos Lucé, near the king's residence at the royal Château d'Amboise. Being frequently visited by Francis, he drew plans for an immense castle town the king intended to erect at Romorantin, and made a mechanical lion, which during a pageant walked toward the king and—upon being struck by a wand—opened its chest to reveal a cluster of lilies.[73][74][o] Leonardo was accompanied during this time by his friend and apprentice, Francesco Melzi, and supported by a pension totalling 10,000 scudi.[65] At some point, Melzi drew a portrait of Leonardo; the only others known from his lifetime were a sketch by an unknown assistant on the back of one of Leonardo's studies (c. 1517)[76] and a drawing by Giovanni Ambrogio Figino depicting an elderly Leonardo with his right arm assuaged by cloth.[77][p][q] The latter, in addition to the record of an October 1517 visit by Louis d'Aragon,[r] confirms an account of Leonardo's right hand being paralytic at the age of 65,[80] which may indicate why he left works such as the Mona Lisa unfinished.[78][81][82] He continued to work at some capacity until eventually becoming ill and bedridden for several months.[80]
+Leonardo died at Clos Lucé on 2 May 1519 at the age of 67, possibly of a stroke.[83][82] Francis I had become a close friend. Vasari describes Leonardo as lamenting on his deathbed, full of repentance, that "he had offended against God and men by failing to practice his art as he should have done."[84] Vasari states that in his last days, Leonardo sent for a priest to make his confession and to receive the Holy Sacrament.[85] Vasari also records that the king held Leonardo's head in his arms as he died, although this story may be legend rather than fact.[s][t] In accordance with his will, sixty beggars carrying tapers followed Leonardo's casket.[87][u] Melzi was the principal heir and executor, receiving, as well as money, Leonardo's paintings, tools, library and personal effects. Leonardo also remembered his other long-time pupil and companion, Salaì, and his servant Baptista de Vilanis, who each received half of Leonardo's vineyards.[88] His brothers received land, and his serving woman received a fur-lined cloak. On 12 August 1519, Leonardo's remains were interred in the Collegiate Church of Saint Florentin at the Château d'Amboise.[89]
+Some 20 years after Leonardo's death, Francis was reported by the goldsmith and sculptor Benvenuto Cellini as saying: "There had never been another man born in the world who knew as much as Leonardo, not so much about painting, sculpture and architecture, as that he was a very great philosopher."[90]
+Florence at the time of Leonardo's youth was the centre of Christian Humanist thought and culture.[35] Leonardo commenced his apprenticeship with Verrocchio in 1466, the year that Verrocchio's master, the great sculptor Donatello, died.[v] The painter Uccello, whose early experiments with perspective were to influence the development of landscape painting, was a very old man. The painters Piero della Francesca and Filippo Lippi, sculptor Luca della Robbia, and architect and writer Leon Battista Alberti were in their sixties. The successful artists of the next generation were Leonardo's teacher Verrocchio, Antonio del Pollaiuolo, and the portrait sculptor Mino da Fiesole.[91][92][93][94]
+Leonardo's youth was spent in a Florence that was ornamented by the works of these artists and by Donatello's contemporaries, Masaccio, whose figurative frescoes were imbued with realism and emotion; and Ghiberti, whose Gates of Paradise, gleaming with gold leaf, displayed the art of combining complex figure compositions with detailed architectural backgrounds. Piero della Francesca had made a detailed study of perspective,[95] and was the first painter to make a scientific study of light. These studies and Alberti's treatise De pictura[96] were to have a profound effect on younger artists and in particular on Leonardo's own observations and artworks.[91][93][94]
+A prevalent tradition in Florence was the small altarpiece of the Virgin and Child. Many of these were created in tempera or glazed terracotta by the workshops of Filippo Lippi, Verrocchio and the prolific della Robbia family.[91] Leonardo's early Madonnas such as The Madonna with a carnation and the Benois Madonna followed this tradition while showing idiosyncratic departures, particularly in the latter in which the Virgin is set at an oblique angle to the picture space with the Christ Child at the opposite angle. This compositional theme was to emerge in Leonardo's later paintings such as The Virgin and Child with St. Anne.[27]
+Leonardo was a contemporary of Botticelli, Domenico Ghirlandaio and Perugino, who were all slightly older than he was.[92] He would have met them at the workshop of Verrocchio, with whom they had associations, and at the Academy of the Medici.[27] Botticelli was a particular favourite of the Medici family, and thus his success as a painter was assured. Ghirlandaio and Perugino were both prolific and ran large workshops. They competently delivered commissions to well-satisfied patrons who appreciated Ghirlandaio's ability to portray the wealthy citizens of Florence within large religious frescoes, and Perugino's ability to deliver a multitude of saints and angels of unfailing sweetness and innocence.[91]
+These three were among those commissioned to paint the walls of the Sistine Chapel, the work commencing with Perugino's employment in 1479. Leonardo was not part of this prestigious commission. His first significant commission, The Adoration of the Magi for the Monks of Scopeto, was never completed.[27]
+In 1476, during the time of Leonardo's association with Verrocchio's workshop, the Portinari Altarpiece by Hugo van der Goes arrived in Florence, bringing from Northern Europe new painterly techniques that were to profoundly affect Leonardo, Ghirlandaio, Perugino and others.[92] In 1479, the Sicilian painter Antonello da Messina, who worked exclusively in oils, travelled north on his way to Venice, where the leading painter Giovanni Bellini adopted the technique of oil painting, quickly making it the preferred method in Venice. Leonardo was also later to visit Venice.[92][94]
+Like the two contemporary architects Donato Bramante (who designed the Belvedere Courtyard) and Antonio da Sangallo the Elder, Leonardo experimented with designs for centrally planned churches, a number of which appear in his journals, as both plans and views, although none was ever realised.[92][97]
+Leonardo's political contemporaries were Lorenzo de' Medici (il Magnifico), who was three years older, and his younger brother Giuliano, who was slain in the Pazzi conspiracy in 1478. Leonardo was sent as an ambassador by the Medici court to Ludovico il Moro, who ruled Milan between 1479 and 1499.[92]
+With Alberti, Leonardo visited the home of the Medici and through them came to know the older Humanist philosophers of whom Marsiglio Ficino, proponent of Neo Platonism; Cristoforo Landino, writer of commentaries on Classical writings, and John Argyropoulos, teacher of Greek and translator of Aristotle were the foremost. Also associated with the Academy of the Medici was Leonardo's contemporary, the brilliant young poet and philosopher Pico della Mirandola.[92][94][87] Leonardo later wrote in the margin of a journal, "The Medici made me and the Medici destroyed me." While it was through the action of Lorenzo that Leonardo received his employment at the court of Milan, it is not known exactly what Leonardo meant by this cryptic comment.[27]
+Although usually named together as the three giants of the High Renaissance, Leonardo, Michelangelo and Raphael were not of the same generation. Leonardo was 23 when Michelangelo was born and 31 when Raphael was born.[92] Raphael died at the age of 37 in 1520, the year after Leonardo died, but Michelangelo went on creating for another 45 years.[93][94]
+Within Leonardo's lifetime, his extraordinary powers of invention, his "outstanding physical beauty," "infinite grace," "great strength and generosity," "regal spirit and tremendous breadth of mind," as described by Vasari,[98] as well as all other aspects of his life, attracted the curiosity of others. One such aspect was his love for animals, likely including vegetarianism and according to Vasari, a habit of purchasing caged birds and releasing them.[99][100]
+Leonardo had many friends who are now renowned either in their fields or for their historical significance. They included the mathematician Luca Pacioli,[101] with whom he collaborated on the book Divina proportione in the 1490s. Leonardo appears to have had no close relationships with women except for his friendship with Cecilia Gallerani and the two Este sisters, Beatrice and Isabella.[102] While on a journey that took him through Mantua, he drew a portrait of Isabella that appears to have been used to create a painted portrait, now lost.[27]
+Beyond friendship, Leonardo kept his private life secret. His sexuality has been the subject of satire, analysis, and speculation. This trend began in the mid-16th century and was revived in the 19th and 20th centuries, most notably by Sigmund Freud.[103] Leonardo's most intimate relationships were perhaps with his pupils Salaì and Melzi. Melzi, writing to inform Leonardo's brothers of his death, described Leonardo's feelings for his pupils as both loving and passionate. It has been claimed since the 16th century that these relationships were of a sexual or erotic nature. Court records of 1476, when he was aged twenty-four, show that Leonardo and three other young men were charged with sodomy in an incident involving a well-known male prostitute. The charges were dismissed for lack of evidence, and there is speculation that since one of the accused, Lionardo de Tornabuoni, was related to Lorenzo de' Medici, the family exerted its influence to secure the dismissal.[104] Since that date much has been written about his presumed homosexuality and its role in his art, particularly in the androgyny and eroticism manifested in Saint John the Baptist and Bacchus and more explicitly in a number of erotic drawings.[105]
+Gian Giacomo Caprotti da Oreno, nicknamed Salaì or Il Salaino ("The Little Unclean One," i.e., the devil), entered Leonardo's household in 1490. After only a year, Leonardo made a list of his misdemeanours, calling him "a thief, a liar, stubborn, and a glutton," after he had made off with money and valuables on at least five occasions and spent a fortune on clothes.[106] Nevertheless, Leonardo treated him with great indulgence, and he remained in Leonardo's household for the next thirty years.[107] Salaì executed a number of paintings under the name of Andrea Salaì, but although Vasari claims that Leonardo "taught him a great deal about painting,"[74] his work is generally considered to be of less artistic merit than others among Leonardo's pupils, such as Marco d'Oggiono and Boltraffio. In 1515, he painted a nude version of the Mona Lisa, known as Monna Vanna.[108] Salaì owned the Mona Lisa at the time of his death in 1524, and in his will it was assessed at 505 lire, an exceptionally high valuation for a small panel portrait.[109]
+In 1506, Leonardo took on another pupil, Count Francesco Melzi, the son of a Lombard aristocrat, who is considered to have been his favourite student. He travelled to France with Leonardo and remained with him until Leonardo's death.[27] Melzi inherited the artistic and scientific works, manuscripts, and collections of Leonardo and administered the estate.
+Despite the recent awareness and admiration of Leonardo as a scientist and inventor, for the better part of four hundred years his fame rested on his achievements as a painter. A handful of works that are either authenticated or attributed to him have been regarded as among the great masterpieces. These paintings are famous for a variety of qualities that have been much imitated by students and discussed at great length by connoisseurs and critics. By the 1490s Leonardo had already been described as a "Divine" painter.[110]
+Among the qualities that make Leonardo's work unique are his innovative techniques for laying on the paint; his detailed knowledge of anatomy, light, botany and geology; his interest in physiognomy and the way humans register emotion in expression and gesture; his innovative use of the human form in figurative composition; and his use of subtle gradation of tone. All these qualities come together in his most famous painted works, the Mona Lisa, the Last Supper, and the Virgin of the Rocks.[111]
+Leonardo first gained attention for his work on the Baptism of Christ, painted in conjunction with Verrocchio. Two other paintings appear to date from his time at Verrocchio's workshop, both of which are Annunciations. One is small, 59 centimetres (23 in) long and 14 centimetres (5.5 in) high. It is a "predella" to go at the base of a larger composition, a painting by Lorenzo di Credi from which it has become separated. The other is a much larger work, 217 centimetres (85 in) long.[112] In both Annunciations, Leonardo used a formal arrangement, like two well-known pictures by Fra Angelico of the same subject, of the Virgin Mary sitting or kneeling to the right of the picture, approached from the left by an angel in profile, with a rich flowing garment, raised wings and bearing a lily. Although previously attributed to Ghirlandaio, the larger work is now generally attributed to Leonardo.[113]
+In the smaller painting, Mary averts her eyes and folds her hands in a gesture that symbolised submission to God's will. Mary is not submissive, however, in the larger piece. The girl, interrupted in her reading by this unexpected messenger, puts a finger in her bible to mark the place and raises her hand in a formal gesture of greeting or surprise.[91] This calm young woman appears to accept her role as the Mother of God, not with resignation but with confidence. In this painting, the young Leonardo presents the humanist face of the Virgin Mary, recognising humanity's role in God's incarnation.
+In the 1480s, Leonardo received two very important commissions and commenced another work that was of ground-breaking importance in terms of composition. Two of the three were never finished, and the third took so long that it was subject to lengthy negotiations over completion and payment.
+One of these paintings was Saint Jerome in the Wilderness, which Bortolon associates with a difficult period of Leonardo's life, as evidenced in his diary: "I thought I was learning to live; I was only learning to die."[27] Although the painting is barely begun, the composition can be seen and is very unusual.[w] Jerome, as a penitent, occupies the middle of the picture, set on a slight diagonal and viewed somewhat from above. His kneeling form takes on a trapezoid shape, with one arm stretched to the outer edge of the painting and his gaze looking in the opposite direction. J. Wasserman points out the link between this painting and Leonardo's anatomical studies.[114] Across the foreground sprawls his symbol, a great lion whose body and tail make a double spiral across the base of the picture space. The other remarkable feature is the sketchy landscape of craggy rocks against which the figure is silhouetted.
+The daring display of figure composition, the landscape elements and personal drama also appear in the great unfinished masterpiece, the Adoration of the Magi, a commission from the Monks of San Donato a Scopeto. It is a complex composition, of about 250 x 250 centimetres. Leonardo did numerous drawings and preparatory studies, including a detailed one in linear perspective of the ruined classical architecture that forms part of the background. In 1482 Leonardo went to Milan at the behest of Lorenzo de' Medici in order to win favour with Ludovico il Moro, and the painting was abandoned.[20][113]
+The third important work of this period is the Virgin of the Rocks, commissioned in Milan for the Confraternity of the Immaculate Conception. The painting, to be done with the assistance of the de Predis brothers, was to fill a large complex altarpiece.[115] Leonardo chose to paint an apocryphal moment of the infancy of Christ when the infant John the Baptist, in protection of an angel, met the Holy Family on the road to Egypt. The painting demonstrates an eerie beauty as the graceful figures kneel in adoration around the infant Christ in a wild landscape of tumbling rock and whirling water.[116] While the painting is quite large, about 200×120 centimetres, it is not nearly as complex as the painting ordered by the monks of St Donato, having only four figures rather than about fifty and a rocky landscape rather than architectural details. The painting was eventually finished; in fact, two versions of the painting were finished: one remained at the chapel of the Confraternity, while Leonardo took the other to France. The Brothers did not get their painting, however, nor the de Predis their payment, until the next century.[38][54]
+Leonardo's most remarkable portrait of this period is the Lady with an Ermine, presumed to be  Cecilia Gallerani (c. 1483–1490), lover of Ludovico Sforza.[117][118] The painting is charcterised by the pose of the figure with the head turned at a very different angle to the torso, unusual at a date when many portraits were still rigidly in profile. The ermine plainly carries symbolic meaning, relating either to the sitter, or to Ludovico who belonged to the prestigious Order of the Ermine.[117]
+Leonardo's most famous painting of the 1490s is The Last Supper, commissioned for the refectory of the Convent of Santa Maria della Grazie in Milan. It represents the last meal shared by Jesus with his disciples before his capture and death, and shows the moment when Jesus has just said "one of you will betray me", and the consternation that this statement caused.[38]
+The novelist Matteo Bandello observed Leonardo at work and wrote that some days he would paint from dawn till dusk without stopping to eat and then not paint for three or four days at a time.[119] This was beyond the comprehension of the prior of the convent, who hounded him until Leonardo asked Ludovico to intervene. Vasari describes how Leonardo, troubled over his ability to adequately depict the faces of Christ and the traitor Judas, told the Duke that he might be obliged to use the prior as his model.[120]
+When finished, the painting was acclaimed as a masterpiece of design and characterization,[121] but it deteriorated rapidly, so that within a hundred years it was described by one viewer as "completely ruined."[122] Leonardo, instead of using the reliable technique of fresco, had used tempera over a ground that was mainly gesso, resulting in a surface subject to mould and to flaking.[123] Despite this, the painting remains one of the most reproduced works of art; countless copies have been made in various mediums.
+It is recorded that in 1492, Leonardo, with assistants painted the Sala delle Asse in the Sforza Castle in Milan,  with a trompe-l'œil depicting trees, with an intricate labyrinth of leaves and knots on the ceiling.[124]
+In 1505 Leonardo was commissioned to paint The Battle of Anghiari in the Salone dei Cinquecento (Hall of the Five Hundred) in the Palazzo Vecchio, Florence. Leonardo devised a dynamic composition depicting four men riding raging war horses engaged in a battle for possession of a standard, at the Battle of Anghiari in 1440.  Michelangelo was assigned the opposite wall to depict the Battle of Cascina. Leonardo's painting deteriorated rapidly and is now known from a copy by Rubens.[125]
+Among the works created by Leonardo in the 16th century is the small portrait known as the Mona Lisa or La Gioconda, the laughing one. In the present era, it is arguably the most famous painting in the world. Its fame rests, in particular, on the elusive smile on the woman's face, its mysterious quality perhaps due to the subtly shadowed corners of the mouth and eyes such that the exact nature of the smile cannot be determined. The shadowy quality for which the work is renowned came to be called "sfumato," or Leonardo's smoke. Vasari, who is generally thought to have known the painting only by repute, said that "the smile was so pleasing that it seemed divine rather than human; and those who saw it were amazed to find that it was as alive as the original."[126][x]
+Other characteristics of the painting are the unadorned dress, in which the eyes and hands have no competition from other details; the dramatic landscape background, in which the world seems to be in a state of flux; the subdued colouring; and the extremely smooth nature of the painterly technique, employing oils laid on much like tempera, and blended on the surface so that the brushstrokes are indistinguishable.[y] Vasari expressed the opinion that the manner of painting would make even "the most confident master...despair and lose heart."[129] The perfect state of preservation and the fact that there is no sign of repair or overpainting is rare in a panel painting of this date.[130]
+In the painting Virgin and Child with St. Anne, the composition again picks up the theme of figures in a landscape, which Wasserman describes as "breathtakingly beautiful"[131] and harkens back to the St Jerome picture with the figure set at an oblique angle. What makes this painting unusual is that there are two obliquely set figures superimposed. Mary is seated on the knee of her mother, St Anne. She leans forward to restrain the Christ Child as he plays roughly with a lamb, the sign of his own impending sacrifice.[38] This painting, which was copied many times, influenced Michelangelo, Raphael, and Andrea del Sarto,[132] and through them Pontormo and Correggio. The trends in composition were adopted in particular by the Venetian painters Tintoretto and Veronese.
+Leonardo was not a prolific painter, but he was a most prolific draftsman, keeping journals full of small sketches and detailed drawings recording all manner of things that took his attention. As well as the journals there exist many studies for paintings, some of which can be identified as preparatory to particular works such as The Adoration of the Magi, The Virgin of the Rocks and The Last Supper.[133]
+His earliest dated drawing is a Landscape of the Arno Valley, 1473, which shows the river, the mountains, Montelupo Castle and the farmlands beyond it in great detail.[27][133][z] According to art historian Ludwig Heydenreich, this is "The first true landscape in art."[134] Massimo Polidoro says that it was the first landscape "not to be the background of some religious scene or a portrait. It is the first [documented] time where a landscape was drawn just for the sake of it."[44]
+Among his famous drawings are the Vitruvian Man, a study of the proportions of the human body; the Head of an Angel, for The Virgin of the Rocks in the Louvre; a botanical study of Star of Bethlehem; and a large drawing (160×100 cm) in black chalk on coloured paper of The Virgin and Child with St. Anne and St. John the Baptist in the National Gallery, London.[133] This drawing employs the subtle sfumato technique of shading, in the manner of the Mona Lisa. It is thought that Leonardo never made a painting from it, the closest similarity being to The Virgin and Child with St. Anne in the Louvre.[135]
+Other drawings of interest include numerous studies generally referred to as "caricatures" because, although exaggerated, they appear to be based upon observation of live models. Vasari relates that if Leonardo saw a person with an interesting face he would follow them around all day observing them.[136] There are numerous studies of beautiful young men, often associated with Salaì, with the rare and much admired facial feature, the so-called "Grecian profile."[aa] These faces are often contrasted with that of a warrior.[133] Salaì is often depicted in fancy-dress costume. Leonardo is known to have designed sets for pageants with which these may be associated. Other, often meticulous, drawings show studies of drapery. A marked development in Leonardo's ability to draw drapery occurred in his early works. Another often-reproduced drawing is a macabre sketch that was done by Leonardo in Florence in 1479 showing the body of Bernardo Baroncelli, hanged in connection with the murder of Giuliano, brother of Lorenzo de' Medici, in the Pazzi conspiracy.[133] In his notes, Leonardo recorded the colours of the robes that Baroncelli was wearing when he died.
+Renaissance humanism recognised no mutually exclusive polarities between the sciences and the arts, and Leonardo's studies in science and engineering are sometimes considered as impressive and innovative as his artistic work.[38] These studies were recorded in 13,000 pages of notes and drawings, which fuse art and natural philosophy (the forerunner of modern science). They were made and maintained daily throughout Leonardo's life and travels, as he made continual observations of the world around him.[38]
+Leonardo's notes and drawings display an enormous range of interests and preoccupations, some as mundane as lists of groceries and people who owed him money and some as intriguing as designs for wings and shoes for walking on water. There are compositions for paintings, studies of details and drapery, studies of faces and emotions, of animals, babies, dissections, plant studies, rock formations, whirlpools, war machines, flying machines and architecture.[38]
+These notebooks—originally loose papers of different types and sizes, were largely entrusted to Leonardo's pupil and heir Francesco Melzi after the master's death.[137] These were to be published, a task of overwhelming difficulty because of its scope and Leonardo's idiosyncratic writing.[138] Some of Leonardo's drawings were copied by an anonymous Milanese artist for a planned treatise on art c. 1570.[139] After Melzi's death in 1570, the collection passed to his son, the lawyer Orazio, who initially took little interest in the journals.[137] In 1587, a Melzi household tutor named Lelio Gavardi took 13 of the manuscripts to Pisa; there, the architect Giovanni Magenta reproached Gavardi for having taken the manuscripts illicitly and returned them to Orazio. Having many more such works in his possession, Orazio gifted the volumes to Magenta. News spread of these lost works of Leonardo's, and Orazio retrieved seven of the 13 manuscripts, which he then gave to Pompeo Leoni for publication in two volumes; one of these was the Codex Atlanticus. The other six works had been distributed to a few others.[140] After Orazio's death, his heirs sold the rest of Leonardo's possessions, and thus began their dispersal.[141]
+Some works have found their way into major collections such as the Royal Library at Windsor Castle, the Louvre, the Biblioteca Nacional de España, the Victoria and Albert Museum, the Biblioteca Ambrosiana in Milan, which holds the 12-volume Codex Atlanticus, and the British Library in London, which has put a selection from the Codex Arundel (BL Arundel MS 263) online.[142] Works have also been at Holkham Hall, the Metropolitan Museum of Art, and in the private hands of John Nicholas Brown I and Robert Lehman.[137] The Codex Leicester is the only privately owned major scientific work of Leonardo; it is owned by Bill Gates and displayed once a year in different cities around the world.
+Most of Leonardo's writings are in mirror-image cursive.[143][44] Since Leonardo wrote with his left hand, it was probably easier for him to write from right to left.[144][ab] Leonardo used a variety of shorthand and symbols, and states in his notes that he intended to prepare them for publication.[143] In many cases a single topic is covered in detail in both words and pictures on a single sheet, together conveying information that would not be lost if the pages were published out of order.[147] Why they were not published during Leonardo's lifetime is unknown.[38]
+Leonardo's approach to science was observational: he tried to understand a phenomenon by describing and depicting it in utmost detail and did not emphasise experiments or theoretical explanation. Since he lacked formal education in Latin and mathematics, contemporary scholars mostly ignored Leonardo the scientist, although he did teach himself Latin. In the 1490s he studied mathematics under Luca Pacioli and prepared a series of drawings of regular solids in a skeletal form to be engraved as plates for Pacioli's book Divina proportione, published in 1509.[38] While living in Milan, he studied light from the summit of Monte Rosa.[63] Scientific writings in his notebook on fossils have been considered as influential on early palaeontology[148][149] and he has been called the father of ichnology.[150]
+The content of his journals suggest that he was planning a series of treatises on a variety of subjects. A coherent treatise on anatomy is said to have been observed during a visit by Cardinal Louis d'Aragon's secretary in 1517.[151] Aspects of his work on the studies of anatomy, light and the landscape were assembled for publication by Melzi and eventually published as A Treatise on Painting in France and Italy in 1651 and Germany in 1724,[152] with engravings based upon drawings by the Classical painter Nicolas Poussin.[153] According to Arasse, the treatise, which in France went into 62 editions in fifty years, caused Leonardo to be seen as "the precursor of French academic thought on art."[38]
+While Leonardo's experimentation followed scientific methods, a recent and exhaustive analysis of Leonardo as a scientist by Fritjof Capra argues that Leonardo was a fundamentally different kind of scientist from Galileo, Newton and other scientists who followed him in that, as a "Renaissance Man", his theorising and hypothesising integrated the arts and particularly painting.[154][page needed]
+Leonardo started his study in the anatomy of the human body under the apprenticeship of Andrea del Verrocchio, who demanded that his students develop a deep knowledge of the subject.[155] As an artist, he quickly became master of topographic anatomy, drawing many studies of muscles, tendons and other visible anatomical features.
+As a successful artist, Leonardo was given permission to dissect human corpses at the Hospital of Santa Maria Nuova in Florence and later at hospitals in Milan and Rome. From 1510 to 1511 he collaborated in his studies with the doctor Marcantonio della Torre. Leonardo made over 240 detailed drawings and wrote about 13,000 words towards a treatise on anatomy.[156] Only a small amount of the material on anatomy was published in Leonardo's Treatise on painting.[138] During the time that Melzi was ordering the material into chapters for publication, they were examined by a number of anatomists and artists, including Vasari, Cellini and Albrecht Dürer, who made a number of drawings from them.[138]
+Leonardo's anatomical drawings include many studies of the human skeleton and its parts, and of muscles and sinews. He studied the mechanical functions of the skeleton and the muscular forces that are applied to it in a manner that prefigured the modern science of biomechanics.[157] He drew the heart and vascular system, the sex organs and other internal organs, making one of the first scientific drawings of a fetus in utero.[133] The drawings and notation are far ahead of their time, and if published would undoubtedly have made a major contribution to medical science.[156]
+Leonardo also closely observed and recorded the effects of age and of human emotion on the physiology, studying in particular the effects of rage. He drew many figures who had significant facial deformities or signs of illness.[38][133] Leonardo also studied and drew the anatomy of many animals, dissecting cows, birds, monkeys, bears, and frogs, and comparing in his drawings their anatomical structure with that of humans. He also made a number of studies of horses.[133]
+Leonardo's dissections and documentation of muscles, nerves, and vessels helped to describe the physiology and mechanics of movement. He attempted to identify the source of 'emotions' and their expression. He found it difficult to incorporate the prevailing system and theories of bodily humours, but eventually he abandoned these physiological explanations of bodily functions. He made the observations that humours were not located in cerebral spaces or ventricles. He documented that the humours were not contained in the heart or the liver, and that it was the heart that defined the circulatory system. He was the first to define atherosclerosis and liver cirrhosis. He created models of the cerebral ventricles with the use of melted wax and constructed a glass aorta to observe the circulation of blood through the aortic valve by using water and grass seed to watch flow patterns. Vesalius published his work on anatomy and physiology in De humani corporis fabrica in 1543.[158]
+During his lifetime, Leonardo was also valued as an engineer. With the same rational and analytical approach that moved him to represent the human body and to investigate anatomy, Leonardo studied and designed a bewildering number of machines and devices. He drew their “anatomy” with unparalleled mastery, producing the first form of the modern technical drawing, including a perfected "exploded view" technique, to represent internal components. Those studies and projects collected in his codices fill more than 5,000 pages.[159] In a letter of 1482 to the lord of Milan Ludovico il Moro, he wrote that he could create all sorts of machines both for the protection of a city and for siege. When he fled from Milan to Venice in 1499, he found employment as an engineer and devised a system of moveable barricades to protect the city from attack. In 1502, he created a scheme for diverting the flow of the Arno river, a project on which Niccolò Machiavelli also worked.[160][161] He continued to contemplate the canalization of Lombardy's plains while in Louis XII's company[63] and of the Loire and its tributaries in the company of Francis I.[162] Leonardo's journals include a vast number of inventions, both practical and impractical. They include musical instruments, a mechanical knight, hydraulic pumps, reversible crank mechanisms, finned mortar shells, and a steam cannon.[27][38]
+Leonardo was fascinated by the phenomenon of flight for much of his life, producing many studies, including Codex on the Flight of Birds (c. 1505), as well as plans for several flying machines, such as a flapping ornithopter and a machine with a helical rotor.[38] A 2003 documentary by British television station Channel Four, titled Leonardo's Dream Machines, various designs by Leonardo, such as a parachute and a giant crossbow, were interpreted and constructed.[163][164] Some of those designs proved successful, whilst others fared less well when tested.
+Research performed by Marc van den Broek revealed older prototypes for more than 100 inventions that are ascribed to Leonardo. Similarities between Leonardo´s illustrations and drawings from the Middle Ages and from Ancient Greece and Rome, the Chinese and Persian Empires, and Egypt suggest that a large portion of Leonardo's inventions had been conceived before his lifetime. Leonardo´s innovation was to combine different functions from existing drafts and set them into scenes that illustrated their utility. By reconstituting technical inventions he created something new.[165]
+Leonardo's fame within his own lifetime was such that the King of France carried him away like a trophy, and was claimed to have supported him in his old age and held him in his arms as he died. Interest in Leonardo and his work has never diminished. Crowds still queue to see his best-known artworks, T-shirts still bear his most famous drawing, and writers continue to hail him as a genius while speculating about his private life, as well as about what one so intelligent actually believed in.[38]
+The continued admiration that Leonardo commanded from painters, critics and historians is reflected in many other written tributes. Baldassare Castiglione, author of Il Cortegiano (The Courtier), wrote in 1528: "...Another of the greatest painters in this world looks down on this art in which he is unequalled..."[166] while the biographer known as "Anonimo Gaddiano" wrote, c. 1540: "His genius was so rare and universal that it can be said that nature worked a miracle on his behalf..."[167]
+Giorgio Vasari, in the enlarged edition of Lives of the Artists (1568)[168] introduced his chapter on Leonardo with the following words:
+In the normal course of events many men and women are born with remarkable talents; but occasionally, in a way that transcends nature, a single person is marvellously endowed by Heaven with beauty, grace and talent in such abundance that he leaves other men far behind, all his actions seem inspired and indeed everything he does clearly comes from God rather than from human skill. Everyone acknowledged that this was true of Leonardo da Vinci, an artist of outstanding physical beauty, who displayed infinite grace in everything that he did and who cultivated his genius so brilliantly that all problems he studied he solved with ease.
+The 19th century brought a particular admiration for Leonardo's genius, causing Henry Fuseli to write in 1801: "Such was the dawn of modern art, when Leonardo da Vinci broke forth with a splendour that distanced former excellence: made up of all the elements that constitute the essence of genius..."[169] This is echoed by A.E. Rio who wrote in 1861: "He towered above all other artists through the strength and the nobility of his talents."[170]
+By the 19th century, the scope of Leonardo's notebooks was known, as well as his paintings. Hippolyte Taine wrote in 1866: "There may not be in the world an example of another genius so universal, so incapable of fulfilment, so full of yearning for the infinite, so naturally refined, so far ahead of his own century and the following centuries."[171]
+Art historian Bernard Berenson wrote in 1896: "Leonardo is the one artist of whom it may be said with perfect literalness: Nothing that he touched but turned into a thing of eternal beauty. Whether it be the cross section of a skull, the structure of a weed, or a study of muscles, he, with his feeling for line and for light and shade, forever transmuted it into life-communicating values."[172]
+The interest in Leonardo's genius has continued unabated; experts study and translate his writings, analyse his paintings using scientific techniques, argue over attributions and search for works which have been recorded but never found.[173] Liana Bortolon, writing in 1967, said: "Because of the multiplicity of interests that spurred him to pursue every field of knowledge...Leonardo can be considered, quite rightly, to have been the universal genius par excellence, and with all the disquieting overtones inherent in that term. Man is as uncomfortable today, faced with a genius, as he was in the 16th century. Five centuries have passed, yet we still view Leonardo with awe."[27]
+Twenty-first-century author Walter Isaacson based much of his biography of Leonardo[104] on thousands of notebook entries, studying the personal notes, sketches, budget notations, and musings of the man whom he considers the greatest of innovators. Isaacson was surprised to discover a "fun, joyous" side of Leonardo in addition to his limitless curiosity and creative genius.[174]
+On the 500th anniversary of Leonardo's death, the Louvre in Paris arranged for the largest ever single exhibit of his work, called Leonardo, between November 2019 and February 2020. The exhibit includes over 100 paintings, drawings and notebooks. Eleven of the paintings that Leonardo completed in his lifetime were included. Five of these are owned by the Louvre, but the Mona Lisa was not included because it is in such great demand among general visitors to the Louvre; it remains on display in its gallery. Vitruvian Man, however, is on display following a legal battle with its owner, the Gallerie dell'Accademia in Venice. Salvator Mundi was also not included because its Saudi owner did not agree to lease the work.[175][176]
+Much of the Collegiate Church of Saint Florentin at the Château d'Amboise, where Leonardo was buried, was damaged during the French Revolution, leading to the church's demolition in 1802.[177] Some of the graves were destroyed in the process, scattering the bones interred there and thereby leaving the whereabouts of Leonardo's remains subject to dispute. While excavating the site in 1863, fine-arts inspector general Arsène Houssaye found a partially complete skeleton with a bronze ring on one finger, some white hair, and stone fragments bearing the inscriptions "EO," "AR," "DUS," and "VINC"—interpreted as forming "Leonardus Vinci".[177][89][178][179] A silver shield found near the bones depicts a beardless Francis I, corresponding to the king's appearance during Leonardo's lifetime, and the skull's inclusion of eight teeth corresponds to someone of approximately the appropriate age.[178] The unusually large skull led Houssaye to believe that he had located Leonardo's remains,[179] but he thought the skeleton seemed too short.[178] Other art historians say that the 1.73 metres (5.7 ft) tall skeleton may well be Leonardo's.[180]
+The remains, except for the ring and a lock of hair which Houssaye kept,[89] were brought to Paris in a lead box, where the skull was allegedly presented to Napoleon III,[178] before being returned to the Château d'Amboise and re-interred in the Chapel of Saint Hubert in 1874.[177] A new memorial tombstone was added by sculptor Francesco La Monaca in the 1930s.[181] Reflecting doubts about the attribution, a plaque above the tomb states that the remains are only presumed to be those of Leonardo.[177] It has since been theorized that the folding of the skeleton's right arm over the head may correspond to the paralysis of Leonardo's right hand.[178][77][83]
+In 2016, it was announced that DNA tests would be conducted to determine whether the attribution is correct.[177] The DNA of the remains will be compared to that of samples collected from Leonardo's work and his half-brother Domenico's descendants;[177] it may also be sequenced.[182] The lock of hair and ring, now in a private US collection,[ac] were displayed in Vinci beginning on 2 May 2019, the 500th anniversary of the artist's death.[183][89]
+Salvator Mundi, a painting by Leonardo depicting Jesus holding an orb, sold for a world record US$450.3 million at a Christie's auction in New York, 15 November 2017.[184] The highest known sale price for any artwork was previously US$300 million, for Willem de Kooning's Interchange, which was sold privately in September 2015.[185] The highest price previously paid for a work of art at auction was for Pablo Picasso's Les Femmes d'Alger, which sold for US$179.4 million in May 2015 at Christie's New York.[185]
+Early sources
+Modern sources

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+Leonardo da Vinci (English: /ˌliːəˈnɑːrdoʊ də ˈvɪntʃi, ˌliːoʊˈ-, ˌleɪoʊˈ-/;[a] 14/15 April 1452[b] – 2 May 1519),[4] was an Italian polymath of the High Renaissance who is widely considered one of the greatest painters of all time (despite less than 25 of his paintings having survived).[c] The Mona Lisa is the most famous of Leonardo's works and the most famous portrait ever made.[5] The Last Supper is the most reproduced religious painting of all time[6] and his Vitruvian Man drawing is also regarded as a cultural icon.[7] He is also known for his his notebooks, in which he made drawings and notes on science and invention; these involve a variety of subjects including anatomy, cartography, painting, and paleontology. Leonardo's collective works compose a contribution to later generations of artists rivalled only by that of his contemporary Michelangelo.[8]
+Properly named Leonardo di ser Piero da Vinci,[d] Leonardo was born out of wedlock to a notary, Piero da Vinci, and a peasant woman, Caterina, in Vinci, in the region of Florence, Italy. Leonardo was educated in the studio of the renowned Italian painter Andrea del Verrocchio. Much of his earlier working life was spent in the service of Ludovico il Moro in Milan, and he later worked in Rome, Bologna and Venice. He spent his last three years in France, where he died in 1519.
+Although he had no formal academic training,[9] many historians and scholars regard Leonardo as the prime exemplar of the "Universal Genius" or "Renaissance Man", an individual of "unquenchable curiosity" and "feverishly inventive imagination."[6] He is widely considered one of the most diversely talented individuals ever to have lived.[10] According to art historian Helen Gardner, the scope and depth of his interests were without precedent in recorded history, and "his mind and personality seem to us superhuman, while the man himself mysterious and remote."[6] Scholars interpret his view of the world as being based in logic, though the empirical methods he used were unorthodox for his time.[11]
+Leonardo is revered for his technological ingenuity. He conceptualized flying machines, a type of armoured fighting vehicle, concentrated solar power, an adding machine,[12] and the double hull. Relatively few of his designs were constructed or even feasible during his lifetime, as the modern scientific approaches to metallurgy and engineering were only in their infancy during the Renaissance. Some of his smaller inventions, however, entered the world of manufacturing unheralded, such as an automated bobbin winder and a machine for testing the tensile strength of wire. He is also sometimes credited with the inventions of the parachute, helicopter, and tank.[13][14] He made substantial discoveries in anatomy, civil engineering, geology, optics, and hydrodynamics, but he did not publish his findings and they had little to no direct influence on subsequent science.[15]
+Leonardo was born on 14/15 April 1452[b] in the Tuscan hill town of Vinci, in the lower valley of the Arno river in the territory of the Medici-ruled Republic of Florence.[18] He was the out-of-wedlock son of Messer Piero Fruosino di Antonio da Vinci, a wealthy Florentine legal notary, and a peasant named Caterina,[e] identified as Caterina Buti del Vacca and more recently as Caterina di Meo Lippi by historian Martin Kemp. There have been many theories regarding Leonardo's mother's identity, including that she was a slave of foreign origin or an impoverished local youth.[17][20][21][22][f] Leonardo had no surname in the modern sense—da Vinci simply meaning "of Vinci"; his full birth name was Lionardo di ser Piero da Vinci,[2][25] meaning "Leonardo, (son) of ser Piero (from) Vinci."[18][d]
+Leonardo spent his first years in the hamlet of Anchiano in the home of his mother, and from at least 1457 lived in the household of his father, grandparents and uncle in the small town of Vinci.[26] His father had married a 16-year-old girl named Albiera Amadori, who loved Leonardo but died young[27] in 1465 without children. In 1468, when Leonardo was 16, his father married again to 20-year-old Francesca Lanfredini, who also died without children. Piero's legitimate heirs were born from his third wife Margherita di Guglielmo, who gave birth to six children, and his fourth and final wife, Lucrezia Cortigiani, who bore him another six heirs.[28][29]
+In all, Leonardo had 12 half-siblings, who were much younger than he was (the last was born when Leonardo was 40 years old) and with whom he had very little contact.[g]
+Leonardo received an informal education in Latin, geometry and mathematics. In later life, Leonardo recorded few distinct childhood incidents. One was of a kite coming to his cradle and opening his mouth with its tail; he regarded this as an omen of his writing on the subject.[31][32] The second occurred while he was exploring in the mountains: he discovered a cave and was both terrified that some great monster might lurk there and driven by curiosity to find out what was inside.[27] He also seems to have remembered some of his childhood observations of water, writing and crossing out the name of his hometown in one of his notebooks on the formation of rivers.[26]
+Leonardo's early life has been the subject of historical conjecture.[33] Vasari, the 16th-century biographer of Renaissance painters, tells a story of Leonardo as a very young man: A local peasant made himself a round shield and requested that Ser Piero have it painted for him. Leonardo, inspired by the story of Medusa, responded with a painting of a monster spitting fire that was so terrifying that his father bought a different shield to give to the peasant and sold Leonardo's to a Florentine art dealer for 100 ducats, who in turn sold it to the Duke of Milan.[34]
+In the mid-1460s, Leonardo's family moved to Florence, and around the age of 14,[30] he became a garzone (studio boy) in the workshop of Verrocchio, who was the leading Florentine painter and sculptor of his time.[35] Leonardo became an apprentice by the age of 17 and remained in training for seven years.[36][37] Other famous painters apprenticed in the workshop or associated with it include Ghirlandaio, Perugino, Botticelli, and Lorenzo di Credi.[27][38] Leonardo was exposed to both theoretical training and a wide range of technical skills,[39] including drafting, chemistry, metallurgy, metal working, plaster casting, leather working, mechanics, and wood-work, as well as the artistic skills of drawing, painting, sculpting, and modelling.[40][h]
+Much of the painting in Verrocchio's workshop was done by his employees. According to Vasari, Leonardo collaborated with Verrocchio on his The Baptism of Christ, painting the young angel holding Jesus' robe in a manner that was so far superior to his master's that Verrocchio put down his brush and never painted again, although this is believed to be an apocryphal story.[41] Close examination reveals areas of the work that have been painted or touched-up over the tempera, using the new technique of oil paint, including the landscape, the rocks seen through the brown mountain stream, and much of the figure of Jesus, bearing witness to the hand of Leonardo.[42] Leonardo may have been the model for two works by Verrocchio: the bronze statue of David in the Bargello, and the Archangel Raphael in Tobias and the Angel.[20]
+By 1472, at the age of 20, Leonardo qualified as a master in the Guild of Saint Luke, the guild of artists and doctors of medicine,[i] but even after his father set him up in his own workshop, his attachment to Verrocchio was such that he continued to collaborate and live with him.[27][43] Leonardo's earliest known dated work is a 1473 pen-and-ink drawing of the Arno valley,[38] which has been cited as the first "pure" landscape in the Occident.[j][44] According to Vasari, the young Leonardo was the first to suggest making the Arno river a navigable channel between Florence and Pisa.[45]
+In January 1478, Leonardo received an independent commission to paint an altarpiece for the Chapel of St. Bernard in the Palazzo Vecchio,[46] an indication of his independence from Verrocchio's studio. One anonymous writer claims that in 1480, Leonardo was living with the Medici and often worked in the garden of the Piazza San Marco, Florence, where a Neoplatonic academy of artists, poets and philosophers organized by the Medici met.[20] In March 1481, he received a commission from the monks of San Donato in Scopeto for The Adoration of the Magi.[47]
+Neither of these initial commissions were completed, being abandoned when Leonardo went to offer his services to Duke of Milan Ludovico Sforza. In 1482, he casted a silver stringed instrument[k] from a horse's skull and ram horns to bring to Sforza,[48] whom he wrote a letter describing the diverse things that he could achieve in the fields of engineering and weapon design, and mentioning that he could paint.[38][48][49]
+Leonardo worked in Milan from 1482 until 1499. He was commissioned to paint the Virgin of the Rocks for the Confraternity of the Immaculate Conception and The Last Supper for the monastery of Santa Maria delle Grazie.[51] In the spring of 1485, Leonardo travelled to Hungary on behalf of Sforza to meet king Matthias Corvinus, and was commissioned by him to paint a Madonna.[52]
+Leonardo was employed on many other projects for Sforza, including the preparation of floats and pageants for special occasions, a drawing and wooden model for a competition to design the cupola for Milan Cathedral (which he withdrew),[53] and a model for a huge equestrian monument to Ludovico's predecessor Francesco Sforza. This would have surpassed in size the only two large equestrian statues of the Renaissance, Donatello's Gattamelata in Padua and Verrocchio's Bartolomeo Colleoni in Venice, and became known as the Gran Cavallo.[38] Leonardo completed a model for the horse and made detailed plans for its casting,[38] but in November 1494, Ludovico gave the bronze to his brother-in-law to be used for a cannon to defend the city from Charles VIII.[38]
+With Ludovico Sforza overthrown at the dawn of the Second Italian War, Leonardo, with his assistant Salaì and friend, the mathematician Luca Pacioli, fled Milan for Venice.[54] There, he was employed as a military architect and engineer, devising methods to defend the city from naval attack.[27] On his return to Florence in 1500, he and his household were guests of the Servite monks at the monastery of Santissima Annunziata and were provided with a workshop where, according to Vasari, Leonardo created the cartoon of The Virgin and Child with St Anne and St John the Baptist, a work that won such admiration that "men and women, young and old" flocked to see it "as if they were attending a great festival."[55][l]
+In Cesena in 1502, Leonardo entered the service of Cesare Borgia, the son of Pope Alexander VI, acting as a military architect and engineer and travelling throughout Italy with his patron.[54] Leonardo created a map of Cesare Borgia's stronghold, a town plan of Imola in order to win his patronage. Maps were extremely rare at the time and it would have seemed like a new concept. Upon seeing it, Cesare hired Leonardo as his chief military engineer and architect. Later in the year, Leonardo produced another map for his patron, one of Chiana Valley, Tuscany, so as to give his patron a better overlay of the land and greater strategic position. He created this map in conjunction with his other project of constructing a dam from the sea to Florence, in order to allow a supply of water to sustain the canal during all seasons.
+Leonardo had left Borgia's service and returned to Florence by early 1503,[57] where he rejoined the Guild of Saint Luke on 18 October of that year. By this same month, Leonardo had begun working on a portrait of Lisa del Giocondo, the model for the Mona Lisa,[58][59] which he would continue working on until his twilight years. In January 1504, he was part of a committee formed to recommend where Michelangelo's statue of David should be placed.[60] He then spent two years in Florence designing and painting a mural of The Battle of Anghiari for the Signoria,[54] with Michelangelo designing its companion piece, The Battle of Cascina.[m]
+In 1506, Leonardo was summoned to Milan by Charles II d'Amboise, the acting French governor of the city. The Council of Florence wished Leonardo to return promptly to finish The Battle of Anghiari, but he was given leave at the behest of Louis XII, who considered commissioning the artist to make some portraits.[63] Leonardo may have commenced a project for an equestrian figure of d'Amboise;[64] a wax model survives and, if genuine, is the only extant example of Leonardo's sculpture. Leonardo was otherwise free to pursue his scientific interests.[63] Many of Leonardo's most prominent pupils either knew or worked with him in Milan,[27] including Bernardino Luini, Giovanni Antonio Boltraffio, and Marco d'Oggiono.
+In 1507, Leonardo was in Florence sorting out a dispute with his brothers over the estate of his father, who had died in 1504. By 1508, Leonardo was back in Milan, living in his own house in Porta Orientale in the parish of Santa Babila.[65]
+In 1512, Leonardo was working on plans for an equestrian monument for Gian Giacomo Trivulzio, but this was prevented by an invasion of a confederation of Swiss, Spanish and Venetian forces, which drove the French from Milan. Leonardo stayed in the city, spending several months in 1513 at the Medici's Vaprio d'Adda villa.[67] In March of that year, Lorenzo de' Medici's son Giovanni assumed the papacy (as Leo X); Leonardo went to Rome that September, where he was received by the pope's brother Giuliano.[67] From September 1513 to 1516, Leonardo spent much of his time living in the Belvedere Courtyard in the Apostolic Palace, where Michelangelo and Raphael were both active.[65] Leonardo was given an allowance of 33 ducats a month, and according to Vasari, decorated a lizard with scales dipped in quicksilver.[68] The pope gave him a painting commission of unknown subject matter, but cancelled it when the artist set about developing a new kind of varnish.[68][n] Leonardo became ill, in what may have been the first of multiple strokes leading to his death.[68] He practiced botany in the Gardens of Vatican City, and was commissioned to make plans for the pope's proposed draining of the Pontine Marshes.[69] He also dissected cadavers, making notes for a treatise on vocal cords;[70] these he gave to an official in hopes of regaining the pope's favor, but was unsuccessful.[68]
+In October 1515, King Francis I of France recaptured Milan.[47] Leonardo was present at the 19 December meeting of Francis I and Leo X, which took place in Bologna.[27][71][72] In 1516, Leonardo entered Francis' service, being given the use of the manor house Clos Lucé, near the king's residence at the royal Château d'Amboise. Being frequently visited by Francis, he drew plans for an immense castle town the king intended to erect at Romorantin, and made a mechanical lion, which during a pageant walked toward the king and—upon being struck by a wand—opened its chest to reveal a cluster of lilies.[73][74][o] Leonardo was accompanied during this time by his friend and apprentice, Francesco Melzi, and supported by a pension totalling 10,000 scudi.[65] At some point, Melzi drew a portrait of Leonardo; the only others known from his lifetime were a sketch by an unknown assistant on the back of one of Leonardo's studies (c. 1517)[76] and a drawing by Giovanni Ambrogio Figino depicting an elderly Leonardo with his right arm assuaged by cloth.[77][p][q] The latter, in addition to the record of an October 1517 visit by Louis d'Aragon,[r] confirms an account of Leonardo's right hand being paralytic at the age of 65,[80] which may indicate why he left works such as the Mona Lisa unfinished.[78][81][82] He continued to work at some capacity until eventually becoming ill and bedridden for several months.[80]
+Leonardo died at Clos Lucé on 2 May 1519 at the age of 67, possibly of a stroke.[83][82] Francis I had become a close friend. Vasari describes Leonardo as lamenting on his deathbed, full of repentance, that "he had offended against God and men by failing to practice his art as he should have done."[84] Vasari states that in his last days, Leonardo sent for a priest to make his confession and to receive the Holy Sacrament.[85] Vasari also records that the king held Leonardo's head in his arms as he died, although this story may be legend rather than fact.[s][t] In accordance with his will, sixty beggars carrying tapers followed Leonardo's casket.[87][u] Melzi was the principal heir and executor, receiving, as well as money, Leonardo's paintings, tools, library and personal effects. Leonardo also remembered his other long-time pupil and companion, Salaì, and his servant Baptista de Vilanis, who each received half of Leonardo's vineyards.[88] His brothers received land, and his serving woman received a fur-lined cloak. On 12 August 1519, Leonardo's remains were interred in the Collegiate Church of Saint Florentin at the Château d'Amboise.[89]
+Some 20 years after Leonardo's death, Francis was reported by the goldsmith and sculptor Benvenuto Cellini as saying: "There had never been another man born in the world who knew as much as Leonardo, not so much about painting, sculpture and architecture, as that he was a very great philosopher."[90]
+Florence at the time of Leonardo's youth was the centre of Christian Humanist thought and culture.[35] Leonardo commenced his apprenticeship with Verrocchio in 1466, the year that Verrocchio's master, the great sculptor Donatello, died.[v] The painter Uccello, whose early experiments with perspective were to influence the development of landscape painting, was a very old man. The painters Piero della Francesca and Filippo Lippi, sculptor Luca della Robbia, and architect and writer Leon Battista Alberti were in their sixties. The successful artists of the next generation were Leonardo's teacher Verrocchio, Antonio del Pollaiuolo, and the portrait sculptor Mino da Fiesole.[91][92][93][94]
+Leonardo's youth was spent in a Florence that was ornamented by the works of these artists and by Donatello's contemporaries, Masaccio, whose figurative frescoes were imbued with realism and emotion; and Ghiberti, whose Gates of Paradise, gleaming with gold leaf, displayed the art of combining complex figure compositions with detailed architectural backgrounds. Piero della Francesca had made a detailed study of perspective,[95] and was the first painter to make a scientific study of light. These studies and Alberti's treatise De pictura[96] were to have a profound effect on younger artists and in particular on Leonardo's own observations and artworks.[91][93][94]
+A prevalent tradition in Florence was the small altarpiece of the Virgin and Child. Many of these were created in tempera or glazed terracotta by the workshops of Filippo Lippi, Verrocchio and the prolific della Robbia family.[91] Leonardo's early Madonnas such as The Madonna with a carnation and the Benois Madonna followed this tradition while showing idiosyncratic departures, particularly in the latter in which the Virgin is set at an oblique angle to the picture space with the Christ Child at the opposite angle. This compositional theme was to emerge in Leonardo's later paintings such as The Virgin and Child with St. Anne.[27]
+Leonardo was a contemporary of Botticelli, Domenico Ghirlandaio and Perugino, who were all slightly older than he was.[92] He would have met them at the workshop of Verrocchio, with whom they had associations, and at the Academy of the Medici.[27] Botticelli was a particular favourite of the Medici family, and thus his success as a painter was assured. Ghirlandaio and Perugino were both prolific and ran large workshops. They competently delivered commissions to well-satisfied patrons who appreciated Ghirlandaio's ability to portray the wealthy citizens of Florence within large religious frescoes, and Perugino's ability to deliver a multitude of saints and angels of unfailing sweetness and innocence.[91]
+These three were among those commissioned to paint the walls of the Sistine Chapel, the work commencing with Perugino's employment in 1479. Leonardo was not part of this prestigious commission. His first significant commission, The Adoration of the Magi for the Monks of Scopeto, was never completed.[27]
+In 1476, during the time of Leonardo's association with Verrocchio's workshop, the Portinari Altarpiece by Hugo van der Goes arrived in Florence, bringing from Northern Europe new painterly techniques that were to profoundly affect Leonardo, Ghirlandaio, Perugino and others.[92] In 1479, the Sicilian painter Antonello da Messina, who worked exclusively in oils, travelled north on his way to Venice, where the leading painter Giovanni Bellini adopted the technique of oil painting, quickly making it the preferred method in Venice. Leonardo was also later to visit Venice.[92][94]
+Like the two contemporary architects Donato Bramante (who designed the Belvedere Courtyard) and Antonio da Sangallo the Elder, Leonardo experimented with designs for centrally planned churches, a number of which appear in his journals, as both plans and views, although none was ever realised.[92][97]
+Leonardo's political contemporaries were Lorenzo de' Medici (il Magnifico), who was three years older, and his younger brother Giuliano, who was slain in the Pazzi conspiracy in 1478. Leonardo was sent as an ambassador by the Medici court to Ludovico il Moro, who ruled Milan between 1479 and 1499.[92]
+With Alberti, Leonardo visited the home of the Medici and through them came to know the older Humanist philosophers of whom Marsiglio Ficino, proponent of Neo Platonism; Cristoforo Landino, writer of commentaries on Classical writings, and John Argyropoulos, teacher of Greek and translator of Aristotle were the foremost. Also associated with the Academy of the Medici was Leonardo's contemporary, the brilliant young poet and philosopher Pico della Mirandola.[92][94][87] Leonardo later wrote in the margin of a journal, "The Medici made me and the Medici destroyed me." While it was through the action of Lorenzo that Leonardo received his employment at the court of Milan, it is not known exactly what Leonardo meant by this cryptic comment.[27]
+Although usually named together as the three giants of the High Renaissance, Leonardo, Michelangelo and Raphael were not of the same generation. Leonardo was 23 when Michelangelo was born and 31 when Raphael was born.[92] Raphael died at the age of 37 in 1520, the year after Leonardo died, but Michelangelo went on creating for another 45 years.[93][94]
+Within Leonardo's lifetime, his extraordinary powers of invention, his "outstanding physical beauty," "infinite grace," "great strength and generosity," "regal spirit and tremendous breadth of mind," as described by Vasari,[98] as well as all other aspects of his life, attracted the curiosity of others. One such aspect was his love for animals, likely including vegetarianism and according to Vasari, a habit of purchasing caged birds and releasing them.[99][100]
+Leonardo had many friends who are now renowned either in their fields or for their historical significance. They included the mathematician Luca Pacioli,[101] with whom he collaborated on the book Divina proportione in the 1490s. Leonardo appears to have had no close relationships with women except for his friendship with Cecilia Gallerani and the two Este sisters, Beatrice and Isabella.[102] While on a journey that took him through Mantua, he drew a portrait of Isabella that appears to have been used to create a painted portrait, now lost.[27]
+Beyond friendship, Leonardo kept his private life secret. His sexuality has been the subject of satire, analysis, and speculation. This trend began in the mid-16th century and was revived in the 19th and 20th centuries, most notably by Sigmund Freud.[103] Leonardo's most intimate relationships were perhaps with his pupils Salaì and Melzi. Melzi, writing to inform Leonardo's brothers of his death, described Leonardo's feelings for his pupils as both loving and passionate. It has been claimed since the 16th century that these relationships were of a sexual or erotic nature. Court records of 1476, when he was aged twenty-four, show that Leonardo and three other young men were charged with sodomy in an incident involving a well-known male prostitute. The charges were dismissed for lack of evidence, and there is speculation that since one of the accused, Lionardo de Tornabuoni, was related to Lorenzo de' Medici, the family exerted its influence to secure the dismissal.[104] Since that date much has been written about his presumed homosexuality and its role in his art, particularly in the androgyny and eroticism manifested in Saint John the Baptist and Bacchus and more explicitly in a number of erotic drawings.[105]
+Gian Giacomo Caprotti da Oreno, nicknamed Salaì or Il Salaino ("The Little Unclean One," i.e., the devil), entered Leonardo's household in 1490. After only a year, Leonardo made a list of his misdemeanours, calling him "a thief, a liar, stubborn, and a glutton," after he had made off with money and valuables on at least five occasions and spent a fortune on clothes.[106] Nevertheless, Leonardo treated him with great indulgence, and he remained in Leonardo's household for the next thirty years.[107] Salaì executed a number of paintings under the name of Andrea Salaì, but although Vasari claims that Leonardo "taught him a great deal about painting,"[74] his work is generally considered to be of less artistic merit than others among Leonardo's pupils, such as Marco d'Oggiono and Boltraffio. In 1515, he painted a nude version of the Mona Lisa, known as Monna Vanna.[108] Salaì owned the Mona Lisa at the time of his death in 1524, and in his will it was assessed at 505 lire, an exceptionally high valuation for a small panel portrait.[109]
+In 1506, Leonardo took on another pupil, Count Francesco Melzi, the son of a Lombard aristocrat, who is considered to have been his favourite student. He travelled to France with Leonardo and remained with him until Leonardo's death.[27] Melzi inherited the artistic and scientific works, manuscripts, and collections of Leonardo and administered the estate.
+Despite the recent awareness and admiration of Leonardo as a scientist and inventor, for the better part of four hundred years his fame rested on his achievements as a painter. A handful of works that are either authenticated or attributed to him have been regarded as among the great masterpieces. These paintings are famous for a variety of qualities that have been much imitated by students and discussed at great length by connoisseurs and critics. By the 1490s Leonardo had already been described as a "Divine" painter.[110]
+Among the qualities that make Leonardo's work unique are his innovative techniques for laying on the paint; his detailed knowledge of anatomy, light, botany and geology; his interest in physiognomy and the way humans register emotion in expression and gesture; his innovative use of the human form in figurative composition; and his use of subtle gradation of tone. All these qualities come together in his most famous painted works, the Mona Lisa, the Last Supper, and the Virgin of the Rocks.[111]
+Leonardo first gained attention for his work on the Baptism of Christ, painted in conjunction with Verrocchio. Two other paintings appear to date from his time at Verrocchio's workshop, both of which are Annunciations. One is small, 59 centimetres (23 in) long and 14 centimetres (5.5 in) high. It is a "predella" to go at the base of a larger composition, a painting by Lorenzo di Credi from which it has become separated. The other is a much larger work, 217 centimetres (85 in) long.[112] In both Annunciations, Leonardo used a formal arrangement, like two well-known pictures by Fra Angelico of the same subject, of the Virgin Mary sitting or kneeling to the right of the picture, approached from the left by an angel in profile, with a rich flowing garment, raised wings and bearing a lily. Although previously attributed to Ghirlandaio, the larger work is now generally attributed to Leonardo.[113]
+In the smaller painting, Mary averts her eyes and folds her hands in a gesture that symbolised submission to God's will. Mary is not submissive, however, in the larger piece. The girl, interrupted in her reading by this unexpected messenger, puts a finger in her bible to mark the place and raises her hand in a formal gesture of greeting or surprise.[91] This calm young woman appears to accept her role as the Mother of God, not with resignation but with confidence. In this painting, the young Leonardo presents the humanist face of the Virgin Mary, recognising humanity's role in God's incarnation.
+In the 1480s, Leonardo received two very important commissions and commenced another work that was of ground-breaking importance in terms of composition. Two of the three were never finished, and the third took so long that it was subject to lengthy negotiations over completion and payment.
+One of these paintings was Saint Jerome in the Wilderness, which Bortolon associates with a difficult period of Leonardo's life, as evidenced in his diary: "I thought I was learning to live; I was only learning to die."[27] Although the painting is barely begun, the composition can be seen and is very unusual.[w] Jerome, as a penitent, occupies the middle of the picture, set on a slight diagonal and viewed somewhat from above. His kneeling form takes on a trapezoid shape, with one arm stretched to the outer edge of the painting and his gaze looking in the opposite direction. J. Wasserman points out the link between this painting and Leonardo's anatomical studies.[114] Across the foreground sprawls his symbol, a great lion whose body and tail make a double spiral across the base of the picture space. The other remarkable feature is the sketchy landscape of craggy rocks against which the figure is silhouetted.
+The daring display of figure composition, the landscape elements and personal drama also appear in the great unfinished masterpiece, the Adoration of the Magi, a commission from the Monks of San Donato a Scopeto. It is a complex composition, of about 250 x 250 centimetres. Leonardo did numerous drawings and preparatory studies, including a detailed one in linear perspective of the ruined classical architecture that forms part of the background. In 1482 Leonardo went to Milan at the behest of Lorenzo de' Medici in order to win favour with Ludovico il Moro, and the painting was abandoned.[20][113]
+The third important work of this period is the Virgin of the Rocks, commissioned in Milan for the Confraternity of the Immaculate Conception. The painting, to be done with the assistance of the de Predis brothers, was to fill a large complex altarpiece.[115] Leonardo chose to paint an apocryphal moment of the infancy of Christ when the infant John the Baptist, in protection of an angel, met the Holy Family on the road to Egypt. The painting demonstrates an eerie beauty as the graceful figures kneel in adoration around the infant Christ in a wild landscape of tumbling rock and whirling water.[116] While the painting is quite large, about 200×120 centimetres, it is not nearly as complex as the painting ordered by the monks of St Donato, having only four figures rather than about fifty and a rocky landscape rather than architectural details. The painting was eventually finished; in fact, two versions of the painting were finished: one remained at the chapel of the Confraternity, while Leonardo took the other to France. The Brothers did not get their painting, however, nor the de Predis their payment, until the next century.[38][54]
+Leonardo's most remarkable portrait of this period is the Lady with an Ermine, presumed to be  Cecilia Gallerani (c. 1483–1490), lover of Ludovico Sforza.[117][118] The painting is charcterised by the pose of the figure with the head turned at a very different angle to the torso, unusual at a date when many portraits were still rigidly in profile. The ermine plainly carries symbolic meaning, relating either to the sitter, or to Ludovico who belonged to the prestigious Order of the Ermine.[117]
+Leonardo's most famous painting of the 1490s is The Last Supper, commissioned for the refectory of the Convent of Santa Maria della Grazie in Milan. It represents the last meal shared by Jesus with his disciples before his capture and death, and shows the moment when Jesus has just said "one of you will betray me", and the consternation that this statement caused.[38]
+The novelist Matteo Bandello observed Leonardo at work and wrote that some days he would paint from dawn till dusk without stopping to eat and then not paint for three or four days at a time.[119] This was beyond the comprehension of the prior of the convent, who hounded him until Leonardo asked Ludovico to intervene. Vasari describes how Leonardo, troubled over his ability to adequately depict the faces of Christ and the traitor Judas, told the Duke that he might be obliged to use the prior as his model.[120]
+When finished, the painting was acclaimed as a masterpiece of design and characterization,[121] but it deteriorated rapidly, so that within a hundred years it was described by one viewer as "completely ruined."[122] Leonardo, instead of using the reliable technique of fresco, had used tempera over a ground that was mainly gesso, resulting in a surface subject to mould and to flaking.[123] Despite this, the painting remains one of the most reproduced works of art; countless copies have been made in various mediums.
+It is recorded that in 1492, Leonardo, with assistants painted the Sala delle Asse in the Sforza Castle in Milan,  with a trompe-l'œil depicting trees, with an intricate labyrinth of leaves and knots on the ceiling.[124]
+In 1505 Leonardo was commissioned to paint The Battle of Anghiari in the Salone dei Cinquecento (Hall of the Five Hundred) in the Palazzo Vecchio, Florence. Leonardo devised a dynamic composition depicting four men riding raging war horses engaged in a battle for possession of a standard, at the Battle of Anghiari in 1440.  Michelangelo was assigned the opposite wall to depict the Battle of Cascina. Leonardo's painting deteriorated rapidly and is now known from a copy by Rubens.[125]
+Among the works created by Leonardo in the 16th century is the small portrait known as the Mona Lisa or La Gioconda, the laughing one. In the present era, it is arguably the most famous painting in the world. Its fame rests, in particular, on the elusive smile on the woman's face, its mysterious quality perhaps due to the subtly shadowed corners of the mouth and eyes such that the exact nature of the smile cannot be determined. The shadowy quality for which the work is renowned came to be called "sfumato," or Leonardo's smoke. Vasari, who is generally thought to have known the painting only by repute, said that "the smile was so pleasing that it seemed divine rather than human; and those who saw it were amazed to find that it was as alive as the original."[126][x]
+Other characteristics of the painting are the unadorned dress, in which the eyes and hands have no competition from other details; the dramatic landscape background, in which the world seems to be in a state of flux; the subdued colouring; and the extremely smooth nature of the painterly technique, employing oils laid on much like tempera, and blended on the surface so that the brushstrokes are indistinguishable.[y] Vasari expressed the opinion that the manner of painting would make even "the most confident master...despair and lose heart."[129] The perfect state of preservation and the fact that there is no sign of repair or overpainting is rare in a panel painting of this date.[130]
+In the painting Virgin and Child with St. Anne, the composition again picks up the theme of figures in a landscape, which Wasserman describes as "breathtakingly beautiful"[131] and harkens back to the St Jerome picture with the figure set at an oblique angle. What makes this painting unusual is that there are two obliquely set figures superimposed. Mary is seated on the knee of her mother, St Anne. She leans forward to restrain the Christ Child as he plays roughly with a lamb, the sign of his own impending sacrifice.[38] This painting, which was copied many times, influenced Michelangelo, Raphael, and Andrea del Sarto,[132] and through them Pontormo and Correggio. The trends in composition were adopted in particular by the Venetian painters Tintoretto and Veronese.
+Leonardo was not a prolific painter, but he was a most prolific draftsman, keeping journals full of small sketches and detailed drawings recording all manner of things that took his attention. As well as the journals there exist many studies for paintings, some of which can be identified as preparatory to particular works such as The Adoration of the Magi, The Virgin of the Rocks and The Last Supper.[133]
+His earliest dated drawing is a Landscape of the Arno Valley, 1473, which shows the river, the mountains, Montelupo Castle and the farmlands beyond it in great detail.[27][133][z] According to art historian Ludwig Heydenreich, this is "The first true landscape in art."[134] Massimo Polidoro says that it was the first landscape "not to be the background of some religious scene or a portrait. It is the first [documented] time where a landscape was drawn just for the sake of it."[44]
+Among his famous drawings are the Vitruvian Man, a study of the proportions of the human body; the Head of an Angel, for The Virgin of the Rocks in the Louvre; a botanical study of Star of Bethlehem; and a large drawing (160×100 cm) in black chalk on coloured paper of The Virgin and Child with St. Anne and St. John the Baptist in the National Gallery, London.[133] This drawing employs the subtle sfumato technique of shading, in the manner of the Mona Lisa. It is thought that Leonardo never made a painting from it, the closest similarity being to The Virgin and Child with St. Anne in the Louvre.[135]
+Other drawings of interest include numerous studies generally referred to as "caricatures" because, although exaggerated, they appear to be based upon observation of live models. Vasari relates that if Leonardo saw a person with an interesting face he would follow them around all day observing them.[136] There are numerous studies of beautiful young men, often associated with Salaì, with the rare and much admired facial feature, the so-called "Grecian profile."[aa] These faces are often contrasted with that of a warrior.[133] Salaì is often depicted in fancy-dress costume. Leonardo is known to have designed sets for pageants with which these may be associated. Other, often meticulous, drawings show studies of drapery. A marked development in Leonardo's ability to draw drapery occurred in his early works. Another often-reproduced drawing is a macabre sketch that was done by Leonardo in Florence in 1479 showing the body of Bernardo Baroncelli, hanged in connection with the murder of Giuliano, brother of Lorenzo de' Medici, in the Pazzi conspiracy.[133] In his notes, Leonardo recorded the colours of the robes that Baroncelli was wearing when he died.
+Renaissance humanism recognised no mutually exclusive polarities between the sciences and the arts, and Leonardo's studies in science and engineering are sometimes considered as impressive and innovative as his artistic work.[38] These studies were recorded in 13,000 pages of notes and drawings, which fuse art and natural philosophy (the forerunner of modern science). They were made and maintained daily throughout Leonardo's life and travels, as he made continual observations of the world around him.[38]
+Leonardo's notes and drawings display an enormous range of interests and preoccupations, some as mundane as lists of groceries and people who owed him money and some as intriguing as designs for wings and shoes for walking on water. There are compositions for paintings, studies of details and drapery, studies of faces and emotions, of animals, babies, dissections, plant studies, rock formations, whirlpools, war machines, flying machines and architecture.[38]
+These notebooks—originally loose papers of different types and sizes, were largely entrusted to Leonardo's pupil and heir Francesco Melzi after the master's death.[137] These were to be published, a task of overwhelming difficulty because of its scope and Leonardo's idiosyncratic writing.[138] Some of Leonardo's drawings were copied by an anonymous Milanese artist for a planned treatise on art c. 1570.[139] After Melzi's death in 1570, the collection passed to his son, the lawyer Orazio, who initially took little interest in the journals.[137] In 1587, a Melzi household tutor named Lelio Gavardi took 13 of the manuscripts to Pisa; there, the architect Giovanni Magenta reproached Gavardi for having taken the manuscripts illicitly and returned them to Orazio. Having many more such works in his possession, Orazio gifted the volumes to Magenta. News spread of these lost works of Leonardo's, and Orazio retrieved seven of the 13 manuscripts, which he then gave to Pompeo Leoni for publication in two volumes; one of these was the Codex Atlanticus. The other six works had been distributed to a few others.[140] After Orazio's death, his heirs sold the rest of Leonardo's possessions, and thus began their dispersal.[141]
+Some works have found their way into major collections such as the Royal Library at Windsor Castle, the Louvre, the Biblioteca Nacional de España, the Victoria and Albert Museum, the Biblioteca Ambrosiana in Milan, which holds the 12-volume Codex Atlanticus, and the British Library in London, which has put a selection from the Codex Arundel (BL Arundel MS 263) online.[142] Works have also been at Holkham Hall, the Metropolitan Museum of Art, and in the private hands of John Nicholas Brown I and Robert Lehman.[137] The Codex Leicester is the only privately owned major scientific work of Leonardo; it is owned by Bill Gates and displayed once a year in different cities around the world.
+Most of Leonardo's writings are in mirror-image cursive.[143][44] Since Leonardo wrote with his left hand, it was probably easier for him to write from right to left.[144][ab] Leonardo used a variety of shorthand and symbols, and states in his notes that he intended to prepare them for publication.[143] In many cases a single topic is covered in detail in both words and pictures on a single sheet, together conveying information that would not be lost if the pages were published out of order.[147] Why they were not published during Leonardo's lifetime is unknown.[38]
+Leonardo's approach to science was observational: he tried to understand a phenomenon by describing and depicting it in utmost detail and did not emphasise experiments or theoretical explanation. Since he lacked formal education in Latin and mathematics, contemporary scholars mostly ignored Leonardo the scientist, although he did teach himself Latin. In the 1490s he studied mathematics under Luca Pacioli and prepared a series of drawings of regular solids in a skeletal form to be engraved as plates for Pacioli's book Divina proportione, published in 1509.[38] While living in Milan, he studied light from the summit of Monte Rosa.[63] Scientific writings in his notebook on fossils have been considered as influential on early palaeontology[148][149] and he has been called the father of ichnology.[150]
+The content of his journals suggest that he was planning a series of treatises on a variety of subjects. A coherent treatise on anatomy is said to have been observed during a visit by Cardinal Louis d'Aragon's secretary in 1517.[151] Aspects of his work on the studies of anatomy, light and the landscape were assembled for publication by Melzi and eventually published as A Treatise on Painting in France and Italy in 1651 and Germany in 1724,[152] with engravings based upon drawings by the Classical painter Nicolas Poussin.[153] According to Arasse, the treatise, which in France went into 62 editions in fifty years, caused Leonardo to be seen as "the precursor of French academic thought on art."[38]
+While Leonardo's experimentation followed scientific methods, a recent and exhaustive analysis of Leonardo as a scientist by Fritjof Capra argues that Leonardo was a fundamentally different kind of scientist from Galileo, Newton and other scientists who followed him in that, as a "Renaissance Man", his theorising and hypothesising integrated the arts and particularly painting.[154][page needed]
+Leonardo started his study in the anatomy of the human body under the apprenticeship of Andrea del Verrocchio, who demanded that his students develop a deep knowledge of the subject.[155] As an artist, he quickly became master of topographic anatomy, drawing many studies of muscles, tendons and other visible anatomical features.
+As a successful artist, Leonardo was given permission to dissect human corpses at the Hospital of Santa Maria Nuova in Florence and later at hospitals in Milan and Rome. From 1510 to 1511 he collaborated in his studies with the doctor Marcantonio della Torre. Leonardo made over 240 detailed drawings and wrote about 13,000 words towards a treatise on anatomy.[156] Only a small amount of the material on anatomy was published in Leonardo's Treatise on painting.[138] During the time that Melzi was ordering the material into chapters for publication, they were examined by a number of anatomists and artists, including Vasari, Cellini and Albrecht Dürer, who made a number of drawings from them.[138]
+Leonardo's anatomical drawings include many studies of the human skeleton and its parts, and of muscles and sinews. He studied the mechanical functions of the skeleton and the muscular forces that are applied to it in a manner that prefigured the modern science of biomechanics.[157] He drew the heart and vascular system, the sex organs and other internal organs, making one of the first scientific drawings of a fetus in utero.[133] The drawings and notation are far ahead of their time, and if published would undoubtedly have made a major contribution to medical science.[156]
+Leonardo also closely observed and recorded the effects of age and of human emotion on the physiology, studying in particular the effects of rage. He drew many figures who had significant facial deformities or signs of illness.[38][133] Leonardo also studied and drew the anatomy of many animals, dissecting cows, birds, monkeys, bears, and frogs, and comparing in his drawings their anatomical structure with that of humans. He also made a number of studies of horses.[133]
+Leonardo's dissections and documentation of muscles, nerves, and vessels helped to describe the physiology and mechanics of movement. He attempted to identify the source of 'emotions' and their expression. He found it difficult to incorporate the prevailing system and theories of bodily humours, but eventually he abandoned these physiological explanations of bodily functions. He made the observations that humours were not located in cerebral spaces or ventricles. He documented that the humours were not contained in the heart or the liver, and that it was the heart that defined the circulatory system. He was the first to define atherosclerosis and liver cirrhosis. He created models of the cerebral ventricles with the use of melted wax and constructed a glass aorta to observe the circulation of blood through the aortic valve by using water and grass seed to watch flow patterns. Vesalius published his work on anatomy and physiology in De humani corporis fabrica in 1543.[158]
+During his lifetime, Leonardo was also valued as an engineer. With the same rational and analytical approach that moved him to represent the human body and to investigate anatomy, Leonardo studied and designed a bewildering number of machines and devices. He drew their “anatomy” with unparalleled mastery, producing the first form of the modern technical drawing, including a perfected "exploded view" technique, to represent internal components. Those studies and projects collected in his codices fill more than 5,000 pages.[159] In a letter of 1482 to the lord of Milan Ludovico il Moro, he wrote that he could create all sorts of machines both for the protection of a city and for siege. When he fled from Milan to Venice in 1499, he found employment as an engineer and devised a system of moveable barricades to protect the city from attack. In 1502, he created a scheme for diverting the flow of the Arno river, a project on which Niccolò Machiavelli also worked.[160][161] He continued to contemplate the canalization of Lombardy's plains while in Louis XII's company[63] and of the Loire and its tributaries in the company of Francis I.[162] Leonardo's journals include a vast number of inventions, both practical and impractical. They include musical instruments, a mechanical knight, hydraulic pumps, reversible crank mechanisms, finned mortar shells, and a steam cannon.[27][38]
+Leonardo was fascinated by the phenomenon of flight for much of his life, producing many studies, including Codex on the Flight of Birds (c. 1505), as well as plans for several flying machines, such as a flapping ornithopter and a machine with a helical rotor.[38] A 2003 documentary by British television station Channel Four, titled Leonardo's Dream Machines, various designs by Leonardo, such as a parachute and a giant crossbow, were interpreted and constructed.[163][164] Some of those designs proved successful, whilst others fared less well when tested.
+Research performed by Marc van den Broek revealed older prototypes for more than 100 inventions that are ascribed to Leonardo. Similarities between Leonardo´s illustrations and drawings from the Middle Ages and from Ancient Greece and Rome, the Chinese and Persian Empires, and Egypt suggest that a large portion of Leonardo's inventions had been conceived before his lifetime. Leonardo´s innovation was to combine different functions from existing drafts and set them into scenes that illustrated their utility. By reconstituting technical inventions he created something new.[165]
+Leonardo's fame within his own lifetime was such that the King of France carried him away like a trophy, and was claimed to have supported him in his old age and held him in his arms as he died. Interest in Leonardo and his work has never diminished. Crowds still queue to see his best-known artworks, T-shirts still bear his most famous drawing, and writers continue to hail him as a genius while speculating about his private life, as well as about what one so intelligent actually believed in.[38]
+The continued admiration that Leonardo commanded from painters, critics and historians is reflected in many other written tributes. Baldassare Castiglione, author of Il Cortegiano (The Courtier), wrote in 1528: "...Another of the greatest painters in this world looks down on this art in which he is unequalled..."[166] while the biographer known as "Anonimo Gaddiano" wrote, c. 1540: "His genius was so rare and universal that it can be said that nature worked a miracle on his behalf..."[167]
+Giorgio Vasari, in the enlarged edition of Lives of the Artists (1568)[168] introduced his chapter on Leonardo with the following words:
+In the normal course of events many men and women are born with remarkable talents; but occasionally, in a way that transcends nature, a single person is marvellously endowed by Heaven with beauty, grace and talent in such abundance that he leaves other men far behind, all his actions seem inspired and indeed everything he does clearly comes from God rather than from human skill. Everyone acknowledged that this was true of Leonardo da Vinci, an artist of outstanding physical beauty, who displayed infinite grace in everything that he did and who cultivated his genius so brilliantly that all problems he studied he solved with ease.
+The 19th century brought a particular admiration for Leonardo's genius, causing Henry Fuseli to write in 1801: "Such was the dawn of modern art, when Leonardo da Vinci broke forth with a splendour that distanced former excellence: made up of all the elements that constitute the essence of genius..."[169] This is echoed by A.E. Rio who wrote in 1861: "He towered above all other artists through the strength and the nobility of his talents."[170]
+By the 19th century, the scope of Leonardo's notebooks was known, as well as his paintings. Hippolyte Taine wrote in 1866: "There may not be in the world an example of another genius so universal, so incapable of fulfilment, so full of yearning for the infinite, so naturally refined, so far ahead of his own century and the following centuries."[171]
+Art historian Bernard Berenson wrote in 1896: "Leonardo is the one artist of whom it may be said with perfect literalness: Nothing that he touched but turned into a thing of eternal beauty. Whether it be the cross section of a skull, the structure of a weed, or a study of muscles, he, with his feeling for line and for light and shade, forever transmuted it into life-communicating values."[172]
+The interest in Leonardo's genius has continued unabated; experts study and translate his writings, analyse his paintings using scientific techniques, argue over attributions and search for works which have been recorded but never found.[173] Liana Bortolon, writing in 1967, said: "Because of the multiplicity of interests that spurred him to pursue every field of knowledge...Leonardo can be considered, quite rightly, to have been the universal genius par excellence, and with all the disquieting overtones inherent in that term. Man is as uncomfortable today, faced with a genius, as he was in the 16th century. Five centuries have passed, yet we still view Leonardo with awe."[27]
+Twenty-first-century author Walter Isaacson based much of his biography of Leonardo[104] on thousands of notebook entries, studying the personal notes, sketches, budget notations, and musings of the man whom he considers the greatest of innovators. Isaacson was surprised to discover a "fun, joyous" side of Leonardo in addition to his limitless curiosity and creative genius.[174]
+On the 500th anniversary of Leonardo's death, the Louvre in Paris arranged for the largest ever single exhibit of his work, called Leonardo, between November 2019 and February 2020. The exhibit includes over 100 paintings, drawings and notebooks. Eleven of the paintings that Leonardo completed in his lifetime were included. Five of these are owned by the Louvre, but the Mona Lisa was not included because it is in such great demand among general visitors to the Louvre; it remains on display in its gallery. Vitruvian Man, however, is on display following a legal battle with its owner, the Gallerie dell'Accademia in Venice. Salvator Mundi was also not included because its Saudi owner did not agree to lease the work.[175][176]
+Much of the Collegiate Church of Saint Florentin at the Château d'Amboise, where Leonardo was buried, was damaged during the French Revolution, leading to the church's demolition in 1802.[177] Some of the graves were destroyed in the process, scattering the bones interred there and thereby leaving the whereabouts of Leonardo's remains subject to dispute. While excavating the site in 1863, fine-arts inspector general Arsène Houssaye found a partially complete skeleton with a bronze ring on one finger, some white hair, and stone fragments bearing the inscriptions "EO," "AR," "DUS," and "VINC"—interpreted as forming "Leonardus Vinci".[177][89][178][179] A silver shield found near the bones depicts a beardless Francis I, corresponding to the king's appearance during Leonardo's lifetime, and the skull's inclusion of eight teeth corresponds to someone of approximately the appropriate age.[178] The unusually large skull led Houssaye to believe that he had located Leonardo's remains,[179] but he thought the skeleton seemed too short.[178] Other art historians say that the 1.73 metres (5.7 ft) tall skeleton may well be Leonardo's.[180]
+The remains, except for the ring and a lock of hair which Houssaye kept,[89] were brought to Paris in a lead box, where the skull was allegedly presented to Napoleon III,[178] before being returned to the Château d'Amboise and re-interred in the Chapel of Saint Hubert in 1874.[177] A new memorial tombstone was added by sculptor Francesco La Monaca in the 1930s.[181] Reflecting doubts about the attribution, a plaque above the tomb states that the remains are only presumed to be those of Leonardo.[177] It has since been theorized that the folding of the skeleton's right arm over the head may correspond to the paralysis of Leonardo's right hand.[178][77][83]
+In 2016, it was announced that DNA tests would be conducted to determine whether the attribution is correct.[177] The DNA of the remains will be compared to that of samples collected from Leonardo's work and his half-brother Domenico's descendants;[177] it may also be sequenced.[182] The lock of hair and ring, now in a private US collection,[ac] were displayed in Vinci beginning on 2 May 2019, the 500th anniversary of the artist's death.[183][89]
+Salvator Mundi, a painting by Leonardo depicting Jesus holding an orb, sold for a world record US$450.3 million at a Christie's auction in New York, 15 November 2017.[184] The highest known sale price for any artwork was previously US$300 million, for Willem de Kooning's Interchange, which was sold privately in September 2015.[185] The highest price previously paid for a work of art at auction was for Pablo Picasso's Les Femmes d'Alger, which sold for US$179.4 million in May 2015 at Christie's New York.[185]
+Early sources
+Modern sources

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+Leonardo da Vinci (English: /ˌliːəˈnɑːrdoʊ də ˈvɪntʃi, ˌliːoʊˈ-, ˌleɪoʊˈ-/;[a] 14/15 April 1452[b] – 2 May 1519),[4] was an Italian polymath of the High Renaissance who is widely considered one of the greatest painters of all time (despite less than 25 of his paintings having survived).[c] The Mona Lisa is the most famous of Leonardo's works and the most famous portrait ever made.[5] The Last Supper is the most reproduced religious painting of all time[6] and his Vitruvian Man drawing is also regarded as a cultural icon.[7] He is also known for his his notebooks, in which he made drawings and notes on science and invention; these involve a variety of subjects including anatomy, cartography, painting, and paleontology. Leonardo's collective works compose a contribution to later generations of artists rivalled only by that of his contemporary Michelangelo.[8]
+Properly named Leonardo di ser Piero da Vinci,[d] Leonardo was born out of wedlock to a notary, Piero da Vinci, and a peasant woman, Caterina, in Vinci, in the region of Florence, Italy. Leonardo was educated in the studio of the renowned Italian painter Andrea del Verrocchio. Much of his earlier working life was spent in the service of Ludovico il Moro in Milan, and he later worked in Rome, Bologna and Venice. He spent his last three years in France, where he died in 1519.
+Although he had no formal academic training,[9] many historians and scholars regard Leonardo as the prime exemplar of the "Universal Genius" or "Renaissance Man", an individual of "unquenchable curiosity" and "feverishly inventive imagination."[6] He is widely considered one of the most diversely talented individuals ever to have lived.[10] According to art historian Helen Gardner, the scope and depth of his interests were without precedent in recorded history, and "his mind and personality seem to us superhuman, while the man himself mysterious and remote."[6] Scholars interpret his view of the world as being based in logic, though the empirical methods he used were unorthodox for his time.[11]
+Leonardo is revered for his technological ingenuity. He conceptualized flying machines, a type of armoured fighting vehicle, concentrated solar power, an adding machine,[12] and the double hull. Relatively few of his designs were constructed or even feasible during his lifetime, as the modern scientific approaches to metallurgy and engineering were only in their infancy during the Renaissance. Some of his smaller inventions, however, entered the world of manufacturing unheralded, such as an automated bobbin winder and a machine for testing the tensile strength of wire. He is also sometimes credited with the inventions of the parachute, helicopter, and tank.[13][14] He made substantial discoveries in anatomy, civil engineering, geology, optics, and hydrodynamics, but he did not publish his findings and they had little to no direct influence on subsequent science.[15]
+Leonardo was born on 14/15 April 1452[b] in the Tuscan hill town of Vinci, in the lower valley of the Arno river in the territory of the Medici-ruled Republic of Florence.[18] He was the out-of-wedlock son of Messer Piero Fruosino di Antonio da Vinci, a wealthy Florentine legal notary, and a peasant named Caterina,[e] identified as Caterina Buti del Vacca and more recently as Caterina di Meo Lippi by historian Martin Kemp. There have been many theories regarding Leonardo's mother's identity, including that she was a slave of foreign origin or an impoverished local youth.[17][20][21][22][f] Leonardo had no surname in the modern sense—da Vinci simply meaning "of Vinci"; his full birth name was Lionardo di ser Piero da Vinci,[2][25] meaning "Leonardo, (son) of ser Piero (from) Vinci."[18][d]
+Leonardo spent his first years in the hamlet of Anchiano in the home of his mother, and from at least 1457 lived in the household of his father, grandparents and uncle in the small town of Vinci.[26] His father had married a 16-year-old girl named Albiera Amadori, who loved Leonardo but died young[27] in 1465 without children. In 1468, when Leonardo was 16, his father married again to 20-year-old Francesca Lanfredini, who also died without children. Piero's legitimate heirs were born from his third wife Margherita di Guglielmo, who gave birth to six children, and his fourth and final wife, Lucrezia Cortigiani, who bore him another six heirs.[28][29]
+In all, Leonardo had 12 half-siblings, who were much younger than he was (the last was born when Leonardo was 40 years old) and with whom he had very little contact.[g]
+Leonardo received an informal education in Latin, geometry and mathematics. In later life, Leonardo recorded few distinct childhood incidents. One was of a kite coming to his cradle and opening his mouth with its tail; he regarded this as an omen of his writing on the subject.[31][32] The second occurred while he was exploring in the mountains: he discovered a cave and was both terrified that some great monster might lurk there and driven by curiosity to find out what was inside.[27] He also seems to have remembered some of his childhood observations of water, writing and crossing out the name of his hometown in one of his notebooks on the formation of rivers.[26]
+Leonardo's early life has been the subject of historical conjecture.[33] Vasari, the 16th-century biographer of Renaissance painters, tells a story of Leonardo as a very young man: A local peasant made himself a round shield and requested that Ser Piero have it painted for him. Leonardo, inspired by the story of Medusa, responded with a painting of a monster spitting fire that was so terrifying that his father bought a different shield to give to the peasant and sold Leonardo's to a Florentine art dealer for 100 ducats, who in turn sold it to the Duke of Milan.[34]
+In the mid-1460s, Leonardo's family moved to Florence, and around the age of 14,[30] he became a garzone (studio boy) in the workshop of Verrocchio, who was the leading Florentine painter and sculptor of his time.[35] Leonardo became an apprentice by the age of 17 and remained in training for seven years.[36][37] Other famous painters apprenticed in the workshop or associated with it include Ghirlandaio, Perugino, Botticelli, and Lorenzo di Credi.[27][38] Leonardo was exposed to both theoretical training and a wide range of technical skills,[39] including drafting, chemistry, metallurgy, metal working, plaster casting, leather working, mechanics, and wood-work, as well as the artistic skills of drawing, painting, sculpting, and modelling.[40][h]
+Much of the painting in Verrocchio's workshop was done by his employees. According to Vasari, Leonardo collaborated with Verrocchio on his The Baptism of Christ, painting the young angel holding Jesus' robe in a manner that was so far superior to his master's that Verrocchio put down his brush and never painted again, although this is believed to be an apocryphal story.[41] Close examination reveals areas of the work that have been painted or touched-up over the tempera, using the new technique of oil paint, including the landscape, the rocks seen through the brown mountain stream, and much of the figure of Jesus, bearing witness to the hand of Leonardo.[42] Leonardo may have been the model for two works by Verrocchio: the bronze statue of David in the Bargello, and the Archangel Raphael in Tobias and the Angel.[20]
+By 1472, at the age of 20, Leonardo qualified as a master in the Guild of Saint Luke, the guild of artists and doctors of medicine,[i] but even after his father set him up in his own workshop, his attachment to Verrocchio was such that he continued to collaborate and live with him.[27][43] Leonardo's earliest known dated work is a 1473 pen-and-ink drawing of the Arno valley,[38] which has been cited as the first "pure" landscape in the Occident.[j][44] According to Vasari, the young Leonardo was the first to suggest making the Arno river a navigable channel between Florence and Pisa.[45]
+In January 1478, Leonardo received an independent commission to paint an altarpiece for the Chapel of St. Bernard in the Palazzo Vecchio,[46] an indication of his independence from Verrocchio's studio. One anonymous writer claims that in 1480, Leonardo was living with the Medici and often worked in the garden of the Piazza San Marco, Florence, where a Neoplatonic academy of artists, poets and philosophers organized by the Medici met.[20] In March 1481, he received a commission from the monks of San Donato in Scopeto for The Adoration of the Magi.[47]
+Neither of these initial commissions were completed, being abandoned when Leonardo went to offer his services to Duke of Milan Ludovico Sforza. In 1482, he casted a silver stringed instrument[k] from a horse's skull and ram horns to bring to Sforza,[48] whom he wrote a letter describing the diverse things that he could achieve in the fields of engineering and weapon design, and mentioning that he could paint.[38][48][49]
+Leonardo worked in Milan from 1482 until 1499. He was commissioned to paint the Virgin of the Rocks for the Confraternity of the Immaculate Conception and The Last Supper for the monastery of Santa Maria delle Grazie.[51] In the spring of 1485, Leonardo travelled to Hungary on behalf of Sforza to meet king Matthias Corvinus, and was commissioned by him to paint a Madonna.[52]
+Leonardo was employed on many other projects for Sforza, including the preparation of floats and pageants for special occasions, a drawing and wooden model for a competition to design the cupola for Milan Cathedral (which he withdrew),[53] and a model for a huge equestrian monument to Ludovico's predecessor Francesco Sforza. This would have surpassed in size the only two large equestrian statues of the Renaissance, Donatello's Gattamelata in Padua and Verrocchio's Bartolomeo Colleoni in Venice, and became known as the Gran Cavallo.[38] Leonardo completed a model for the horse and made detailed plans for its casting,[38] but in November 1494, Ludovico gave the bronze to his brother-in-law to be used for a cannon to defend the city from Charles VIII.[38]
+With Ludovico Sforza overthrown at the dawn of the Second Italian War, Leonardo, with his assistant Salaì and friend, the mathematician Luca Pacioli, fled Milan for Venice.[54] There, he was employed as a military architect and engineer, devising methods to defend the city from naval attack.[27] On his return to Florence in 1500, he and his household were guests of the Servite monks at the monastery of Santissima Annunziata and were provided with a workshop where, according to Vasari, Leonardo created the cartoon of The Virgin and Child with St Anne and St John the Baptist, a work that won such admiration that "men and women, young and old" flocked to see it "as if they were attending a great festival."[55][l]
+In Cesena in 1502, Leonardo entered the service of Cesare Borgia, the son of Pope Alexander VI, acting as a military architect and engineer and travelling throughout Italy with his patron.[54] Leonardo created a map of Cesare Borgia's stronghold, a town plan of Imola in order to win his patronage. Maps were extremely rare at the time and it would have seemed like a new concept. Upon seeing it, Cesare hired Leonardo as his chief military engineer and architect. Later in the year, Leonardo produced another map for his patron, one of Chiana Valley, Tuscany, so as to give his patron a better overlay of the land and greater strategic position. He created this map in conjunction with his other project of constructing a dam from the sea to Florence, in order to allow a supply of water to sustain the canal during all seasons.
+Leonardo had left Borgia's service and returned to Florence by early 1503,[57] where he rejoined the Guild of Saint Luke on 18 October of that year. By this same month, Leonardo had begun working on a portrait of Lisa del Giocondo, the model for the Mona Lisa,[58][59] which he would continue working on until his twilight years. In January 1504, he was part of a committee formed to recommend where Michelangelo's statue of David should be placed.[60] He then spent two years in Florence designing and painting a mural of The Battle of Anghiari for the Signoria,[54] with Michelangelo designing its companion piece, The Battle of Cascina.[m]
+In 1506, Leonardo was summoned to Milan by Charles II d'Amboise, the acting French governor of the city. The Council of Florence wished Leonardo to return promptly to finish The Battle of Anghiari, but he was given leave at the behest of Louis XII, who considered commissioning the artist to make some portraits.[63] Leonardo may have commenced a project for an equestrian figure of d'Amboise;[64] a wax model survives and, if genuine, is the only extant example of Leonardo's sculpture. Leonardo was otherwise free to pursue his scientific interests.[63] Many of Leonardo's most prominent pupils either knew or worked with him in Milan,[27] including Bernardino Luini, Giovanni Antonio Boltraffio, and Marco d'Oggiono.
+In 1507, Leonardo was in Florence sorting out a dispute with his brothers over the estate of his father, who had died in 1504. By 1508, Leonardo was back in Milan, living in his own house in Porta Orientale in the parish of Santa Babila.[65]
+In 1512, Leonardo was working on plans for an equestrian monument for Gian Giacomo Trivulzio, but this was prevented by an invasion of a confederation of Swiss, Spanish and Venetian forces, which drove the French from Milan. Leonardo stayed in the city, spending several months in 1513 at the Medici's Vaprio d'Adda villa.[67] In March of that year, Lorenzo de' Medici's son Giovanni assumed the papacy (as Leo X); Leonardo went to Rome that September, where he was received by the pope's brother Giuliano.[67] From September 1513 to 1516, Leonardo spent much of his time living in the Belvedere Courtyard in the Apostolic Palace, where Michelangelo and Raphael were both active.[65] Leonardo was given an allowance of 33 ducats a month, and according to Vasari, decorated a lizard with scales dipped in quicksilver.[68] The pope gave him a painting commission of unknown subject matter, but cancelled it when the artist set about developing a new kind of varnish.[68][n] Leonardo became ill, in what may have been the first of multiple strokes leading to his death.[68] He practiced botany in the Gardens of Vatican City, and was commissioned to make plans for the pope's proposed draining of the Pontine Marshes.[69] He also dissected cadavers, making notes for a treatise on vocal cords;[70] these he gave to an official in hopes of regaining the pope's favor, but was unsuccessful.[68]
+In October 1515, King Francis I of France recaptured Milan.[47] Leonardo was present at the 19 December meeting of Francis I and Leo X, which took place in Bologna.[27][71][72] In 1516, Leonardo entered Francis' service, being given the use of the manor house Clos Lucé, near the king's residence at the royal Château d'Amboise. Being frequently visited by Francis, he drew plans for an immense castle town the king intended to erect at Romorantin, and made a mechanical lion, which during a pageant walked toward the king and—upon being struck by a wand—opened its chest to reveal a cluster of lilies.[73][74][o] Leonardo was accompanied during this time by his friend and apprentice, Francesco Melzi, and supported by a pension totalling 10,000 scudi.[65] At some point, Melzi drew a portrait of Leonardo; the only others known from his lifetime were a sketch by an unknown assistant on the back of one of Leonardo's studies (c. 1517)[76] and a drawing by Giovanni Ambrogio Figino depicting an elderly Leonardo with his right arm assuaged by cloth.[77][p][q] The latter, in addition to the record of an October 1517 visit by Louis d'Aragon,[r] confirms an account of Leonardo's right hand being paralytic at the age of 65,[80] which may indicate why he left works such as the Mona Lisa unfinished.[78][81][82] He continued to work at some capacity until eventually becoming ill and bedridden for several months.[80]
+Leonardo died at Clos Lucé on 2 May 1519 at the age of 67, possibly of a stroke.[83][82] Francis I had become a close friend. Vasari describes Leonardo as lamenting on his deathbed, full of repentance, that "he had offended against God and men by failing to practice his art as he should have done."[84] Vasari states that in his last days, Leonardo sent for a priest to make his confession and to receive the Holy Sacrament.[85] Vasari also records that the king held Leonardo's head in his arms as he died, although this story may be legend rather than fact.[s][t] In accordance with his will, sixty beggars carrying tapers followed Leonardo's casket.[87][u] Melzi was the principal heir and executor, receiving, as well as money, Leonardo's paintings, tools, library and personal effects. Leonardo also remembered his other long-time pupil and companion, Salaì, and his servant Baptista de Vilanis, who each received half of Leonardo's vineyards.[88] His brothers received land, and his serving woman received a fur-lined cloak. On 12 August 1519, Leonardo's remains were interred in the Collegiate Church of Saint Florentin at the Château d'Amboise.[89]
+Some 20 years after Leonardo's death, Francis was reported by the goldsmith and sculptor Benvenuto Cellini as saying: "There had never been another man born in the world who knew as much as Leonardo, not so much about painting, sculpture and architecture, as that he was a very great philosopher."[90]
+Florence at the time of Leonardo's youth was the centre of Christian Humanist thought and culture.[35] Leonardo commenced his apprenticeship with Verrocchio in 1466, the year that Verrocchio's master, the great sculptor Donatello, died.[v] The painter Uccello, whose early experiments with perspective were to influence the development of landscape painting, was a very old man. The painters Piero della Francesca and Filippo Lippi, sculptor Luca della Robbia, and architect and writer Leon Battista Alberti were in their sixties. The successful artists of the next generation were Leonardo's teacher Verrocchio, Antonio del Pollaiuolo, and the portrait sculptor Mino da Fiesole.[91][92][93][94]
+Leonardo's youth was spent in a Florence that was ornamented by the works of these artists and by Donatello's contemporaries, Masaccio, whose figurative frescoes were imbued with realism and emotion; and Ghiberti, whose Gates of Paradise, gleaming with gold leaf, displayed the art of combining complex figure compositions with detailed architectural backgrounds. Piero della Francesca had made a detailed study of perspective,[95] and was the first painter to make a scientific study of light. These studies and Alberti's treatise De pictura[96] were to have a profound effect on younger artists and in particular on Leonardo's own observations and artworks.[91][93][94]
+A prevalent tradition in Florence was the small altarpiece of the Virgin and Child. Many of these were created in tempera or glazed terracotta by the workshops of Filippo Lippi, Verrocchio and the prolific della Robbia family.[91] Leonardo's early Madonnas such as The Madonna with a carnation and the Benois Madonna followed this tradition while showing idiosyncratic departures, particularly in the latter in which the Virgin is set at an oblique angle to the picture space with the Christ Child at the opposite angle. This compositional theme was to emerge in Leonardo's later paintings such as The Virgin and Child with St. Anne.[27]
+Leonardo was a contemporary of Botticelli, Domenico Ghirlandaio and Perugino, who were all slightly older than he was.[92] He would have met them at the workshop of Verrocchio, with whom they had associations, and at the Academy of the Medici.[27] Botticelli was a particular favourite of the Medici family, and thus his success as a painter was assured. Ghirlandaio and Perugino were both prolific and ran large workshops. They competently delivered commissions to well-satisfied patrons who appreciated Ghirlandaio's ability to portray the wealthy citizens of Florence within large religious frescoes, and Perugino's ability to deliver a multitude of saints and angels of unfailing sweetness and innocence.[91]
+These three were among those commissioned to paint the walls of the Sistine Chapel, the work commencing with Perugino's employment in 1479. Leonardo was not part of this prestigious commission. His first significant commission, The Adoration of the Magi for the Monks of Scopeto, was never completed.[27]
+In 1476, during the time of Leonardo's association with Verrocchio's workshop, the Portinari Altarpiece by Hugo van der Goes arrived in Florence, bringing from Northern Europe new painterly techniques that were to profoundly affect Leonardo, Ghirlandaio, Perugino and others.[92] In 1479, the Sicilian painter Antonello da Messina, who worked exclusively in oils, travelled north on his way to Venice, where the leading painter Giovanni Bellini adopted the technique of oil painting, quickly making it the preferred method in Venice. Leonardo was also later to visit Venice.[92][94]
+Like the two contemporary architects Donato Bramante (who designed the Belvedere Courtyard) and Antonio da Sangallo the Elder, Leonardo experimented with designs for centrally planned churches, a number of which appear in his journals, as both plans and views, although none was ever realised.[92][97]
+Leonardo's political contemporaries were Lorenzo de' Medici (il Magnifico), who was three years older, and his younger brother Giuliano, who was slain in the Pazzi conspiracy in 1478. Leonardo was sent as an ambassador by the Medici court to Ludovico il Moro, who ruled Milan between 1479 and 1499.[92]
+With Alberti, Leonardo visited the home of the Medici and through them came to know the older Humanist philosophers of whom Marsiglio Ficino, proponent of Neo Platonism; Cristoforo Landino, writer of commentaries on Classical writings, and John Argyropoulos, teacher of Greek and translator of Aristotle were the foremost. Also associated with the Academy of the Medici was Leonardo's contemporary, the brilliant young poet and philosopher Pico della Mirandola.[92][94][87] Leonardo later wrote in the margin of a journal, "The Medici made me and the Medici destroyed me." While it was through the action of Lorenzo that Leonardo received his employment at the court of Milan, it is not known exactly what Leonardo meant by this cryptic comment.[27]
+Although usually named together as the three giants of the High Renaissance, Leonardo, Michelangelo and Raphael were not of the same generation. Leonardo was 23 when Michelangelo was born and 31 when Raphael was born.[92] Raphael died at the age of 37 in 1520, the year after Leonardo died, but Michelangelo went on creating for another 45 years.[93][94]
+Within Leonardo's lifetime, his extraordinary powers of invention, his "outstanding physical beauty," "infinite grace," "great strength and generosity," "regal spirit and tremendous breadth of mind," as described by Vasari,[98] as well as all other aspects of his life, attracted the curiosity of others. One such aspect was his love for animals, likely including vegetarianism and according to Vasari, a habit of purchasing caged birds and releasing them.[99][100]
+Leonardo had many friends who are now renowned either in their fields or for their historical significance. They included the mathematician Luca Pacioli,[101] with whom he collaborated on the book Divina proportione in the 1490s. Leonardo appears to have had no close relationships with women except for his friendship with Cecilia Gallerani and the two Este sisters, Beatrice and Isabella.[102] While on a journey that took him through Mantua, he drew a portrait of Isabella that appears to have been used to create a painted portrait, now lost.[27]
+Beyond friendship, Leonardo kept his private life secret. His sexuality has been the subject of satire, analysis, and speculation. This trend began in the mid-16th century and was revived in the 19th and 20th centuries, most notably by Sigmund Freud.[103] Leonardo's most intimate relationships were perhaps with his pupils Salaì and Melzi. Melzi, writing to inform Leonardo's brothers of his death, described Leonardo's feelings for his pupils as both loving and passionate. It has been claimed since the 16th century that these relationships were of a sexual or erotic nature. Court records of 1476, when he was aged twenty-four, show that Leonardo and three other young men were charged with sodomy in an incident involving a well-known male prostitute. The charges were dismissed for lack of evidence, and there is speculation that since one of the accused, Lionardo de Tornabuoni, was related to Lorenzo de' Medici, the family exerted its influence to secure the dismissal.[104] Since that date much has been written about his presumed homosexuality and its role in his art, particularly in the androgyny and eroticism manifested in Saint John the Baptist and Bacchus and more explicitly in a number of erotic drawings.[105]
+Gian Giacomo Caprotti da Oreno, nicknamed Salaì or Il Salaino ("The Little Unclean One," i.e., the devil), entered Leonardo's household in 1490. After only a year, Leonardo made a list of his misdemeanours, calling him "a thief, a liar, stubborn, and a glutton," after he had made off with money and valuables on at least five occasions and spent a fortune on clothes.[106] Nevertheless, Leonardo treated him with great indulgence, and he remained in Leonardo's household for the next thirty years.[107] Salaì executed a number of paintings under the name of Andrea Salaì, but although Vasari claims that Leonardo "taught him a great deal about painting,"[74] his work is generally considered to be of less artistic merit than others among Leonardo's pupils, such as Marco d'Oggiono and Boltraffio. In 1515, he painted a nude version of the Mona Lisa, known as Monna Vanna.[108] Salaì owned the Mona Lisa at the time of his death in 1524, and in his will it was assessed at 505 lire, an exceptionally high valuation for a small panel portrait.[109]
+In 1506, Leonardo took on another pupil, Count Francesco Melzi, the son of a Lombard aristocrat, who is considered to have been his favourite student. He travelled to France with Leonardo and remained with him until Leonardo's death.[27] Melzi inherited the artistic and scientific works, manuscripts, and collections of Leonardo and administered the estate.
+Despite the recent awareness and admiration of Leonardo as a scientist and inventor, for the better part of four hundred years his fame rested on his achievements as a painter. A handful of works that are either authenticated or attributed to him have been regarded as among the great masterpieces. These paintings are famous for a variety of qualities that have been much imitated by students and discussed at great length by connoisseurs and critics. By the 1490s Leonardo had already been described as a "Divine" painter.[110]
+Among the qualities that make Leonardo's work unique are his innovative techniques for laying on the paint; his detailed knowledge of anatomy, light, botany and geology; his interest in physiognomy and the way humans register emotion in expression and gesture; his innovative use of the human form in figurative composition; and his use of subtle gradation of tone. All these qualities come together in his most famous painted works, the Mona Lisa, the Last Supper, and the Virgin of the Rocks.[111]
+Leonardo first gained attention for his work on the Baptism of Christ, painted in conjunction with Verrocchio. Two other paintings appear to date from his time at Verrocchio's workshop, both of which are Annunciations. One is small, 59 centimetres (23 in) long and 14 centimetres (5.5 in) high. It is a "predella" to go at the base of a larger composition, a painting by Lorenzo di Credi from which it has become separated. The other is a much larger work, 217 centimetres (85 in) long.[112] In both Annunciations, Leonardo used a formal arrangement, like two well-known pictures by Fra Angelico of the same subject, of the Virgin Mary sitting or kneeling to the right of the picture, approached from the left by an angel in profile, with a rich flowing garment, raised wings and bearing a lily. Although previously attributed to Ghirlandaio, the larger work is now generally attributed to Leonardo.[113]
+In the smaller painting, Mary averts her eyes and folds her hands in a gesture that symbolised submission to God's will. Mary is not submissive, however, in the larger piece. The girl, interrupted in her reading by this unexpected messenger, puts a finger in her bible to mark the place and raises her hand in a formal gesture of greeting or surprise.[91] This calm young woman appears to accept her role as the Mother of God, not with resignation but with confidence. In this painting, the young Leonardo presents the humanist face of the Virgin Mary, recognising humanity's role in God's incarnation.
+In the 1480s, Leonardo received two very important commissions and commenced another work that was of ground-breaking importance in terms of composition. Two of the three were never finished, and the third took so long that it was subject to lengthy negotiations over completion and payment.
+One of these paintings was Saint Jerome in the Wilderness, which Bortolon associates with a difficult period of Leonardo's life, as evidenced in his diary: "I thought I was learning to live; I was only learning to die."[27] Although the painting is barely begun, the composition can be seen and is very unusual.[w] Jerome, as a penitent, occupies the middle of the picture, set on a slight diagonal and viewed somewhat from above. His kneeling form takes on a trapezoid shape, with one arm stretched to the outer edge of the painting and his gaze looking in the opposite direction. J. Wasserman points out the link between this painting and Leonardo's anatomical studies.[114] Across the foreground sprawls his symbol, a great lion whose body and tail make a double spiral across the base of the picture space. The other remarkable feature is the sketchy landscape of craggy rocks against which the figure is silhouetted.
+The daring display of figure composition, the landscape elements and personal drama also appear in the great unfinished masterpiece, the Adoration of the Magi, a commission from the Monks of San Donato a Scopeto. It is a complex composition, of about 250 x 250 centimetres. Leonardo did numerous drawings and preparatory studies, including a detailed one in linear perspective of the ruined classical architecture that forms part of the background. In 1482 Leonardo went to Milan at the behest of Lorenzo de' Medici in order to win favour with Ludovico il Moro, and the painting was abandoned.[20][113]
+The third important work of this period is the Virgin of the Rocks, commissioned in Milan for the Confraternity of the Immaculate Conception. The painting, to be done with the assistance of the de Predis brothers, was to fill a large complex altarpiece.[115] Leonardo chose to paint an apocryphal moment of the infancy of Christ when the infant John the Baptist, in protection of an angel, met the Holy Family on the road to Egypt. The painting demonstrates an eerie beauty as the graceful figures kneel in adoration around the infant Christ in a wild landscape of tumbling rock and whirling water.[116] While the painting is quite large, about 200×120 centimetres, it is not nearly as complex as the painting ordered by the monks of St Donato, having only four figures rather than about fifty and a rocky landscape rather than architectural details. The painting was eventually finished; in fact, two versions of the painting were finished: one remained at the chapel of the Confraternity, while Leonardo took the other to France. The Brothers did not get their painting, however, nor the de Predis their payment, until the next century.[38][54]
+Leonardo's most remarkable portrait of this period is the Lady with an Ermine, presumed to be  Cecilia Gallerani (c. 1483–1490), lover of Ludovico Sforza.[117][118] The painting is charcterised by the pose of the figure with the head turned at a very different angle to the torso, unusual at a date when many portraits were still rigidly in profile. The ermine plainly carries symbolic meaning, relating either to the sitter, or to Ludovico who belonged to the prestigious Order of the Ermine.[117]
+Leonardo's most famous painting of the 1490s is The Last Supper, commissioned for the refectory of the Convent of Santa Maria della Grazie in Milan. It represents the last meal shared by Jesus with his disciples before his capture and death, and shows the moment when Jesus has just said "one of you will betray me", and the consternation that this statement caused.[38]
+The novelist Matteo Bandello observed Leonardo at work and wrote that some days he would paint from dawn till dusk without stopping to eat and then not paint for three or four days at a time.[119] This was beyond the comprehension of the prior of the convent, who hounded him until Leonardo asked Ludovico to intervene. Vasari describes how Leonardo, troubled over his ability to adequately depict the faces of Christ and the traitor Judas, told the Duke that he might be obliged to use the prior as his model.[120]
+When finished, the painting was acclaimed as a masterpiece of design and characterization,[121] but it deteriorated rapidly, so that within a hundred years it was described by one viewer as "completely ruined."[122] Leonardo, instead of using the reliable technique of fresco, had used tempera over a ground that was mainly gesso, resulting in a surface subject to mould and to flaking.[123] Despite this, the painting remains one of the most reproduced works of art; countless copies have been made in various mediums.
+It is recorded that in 1492, Leonardo, with assistants painted the Sala delle Asse in the Sforza Castle in Milan,  with a trompe-l'œil depicting trees, with an intricate labyrinth of leaves and knots on the ceiling.[124]
+In 1505 Leonardo was commissioned to paint The Battle of Anghiari in the Salone dei Cinquecento (Hall of the Five Hundred) in the Palazzo Vecchio, Florence. Leonardo devised a dynamic composition depicting four men riding raging war horses engaged in a battle for possession of a standard, at the Battle of Anghiari in 1440.  Michelangelo was assigned the opposite wall to depict the Battle of Cascina. Leonardo's painting deteriorated rapidly and is now known from a copy by Rubens.[125]
+Among the works created by Leonardo in the 16th century is the small portrait known as the Mona Lisa or La Gioconda, the laughing one. In the present era, it is arguably the most famous painting in the world. Its fame rests, in particular, on the elusive smile on the woman's face, its mysterious quality perhaps due to the subtly shadowed corners of the mouth and eyes such that the exact nature of the smile cannot be determined. The shadowy quality for which the work is renowned came to be called "sfumato," or Leonardo's smoke. Vasari, who is generally thought to have known the painting only by repute, said that "the smile was so pleasing that it seemed divine rather than human; and those who saw it were amazed to find that it was as alive as the original."[126][x]
+Other characteristics of the painting are the unadorned dress, in which the eyes and hands have no competition from other details; the dramatic landscape background, in which the world seems to be in a state of flux; the subdued colouring; and the extremely smooth nature of the painterly technique, employing oils laid on much like tempera, and blended on the surface so that the brushstrokes are indistinguishable.[y] Vasari expressed the opinion that the manner of painting would make even "the most confident master...despair and lose heart."[129] The perfect state of preservation and the fact that there is no sign of repair or overpainting is rare in a panel painting of this date.[130]
+In the painting Virgin and Child with St. Anne, the composition again picks up the theme of figures in a landscape, which Wasserman describes as "breathtakingly beautiful"[131] and harkens back to the St Jerome picture with the figure set at an oblique angle. What makes this painting unusual is that there are two obliquely set figures superimposed. Mary is seated on the knee of her mother, St Anne. She leans forward to restrain the Christ Child as he plays roughly with a lamb, the sign of his own impending sacrifice.[38] This painting, which was copied many times, influenced Michelangelo, Raphael, and Andrea del Sarto,[132] and through them Pontormo and Correggio. The trends in composition were adopted in particular by the Venetian painters Tintoretto and Veronese.
+Leonardo was not a prolific painter, but he was a most prolific draftsman, keeping journals full of small sketches and detailed drawings recording all manner of things that took his attention. As well as the journals there exist many studies for paintings, some of which can be identified as preparatory to particular works such as The Adoration of the Magi, The Virgin of the Rocks and The Last Supper.[133]
+His earliest dated drawing is a Landscape of the Arno Valley, 1473, which shows the river, the mountains, Montelupo Castle and the farmlands beyond it in great detail.[27][133][z] According to art historian Ludwig Heydenreich, this is "The first true landscape in art."[134] Massimo Polidoro says that it was the first landscape "not to be the background of some religious scene or a portrait. It is the first [documented] time where a landscape was drawn just for the sake of it."[44]
+Among his famous drawings are the Vitruvian Man, a study of the proportions of the human body; the Head of an Angel, for The Virgin of the Rocks in the Louvre; a botanical study of Star of Bethlehem; and a large drawing (160×100 cm) in black chalk on coloured paper of The Virgin and Child with St. Anne and St. John the Baptist in the National Gallery, London.[133] This drawing employs the subtle sfumato technique of shading, in the manner of the Mona Lisa. It is thought that Leonardo never made a painting from it, the closest similarity being to The Virgin and Child with St. Anne in the Louvre.[135]
+Other drawings of interest include numerous studies generally referred to as "caricatures" because, although exaggerated, they appear to be based upon observation of live models. Vasari relates that if Leonardo saw a person with an interesting face he would follow them around all day observing them.[136] There are numerous studies of beautiful young men, often associated with Salaì, with the rare and much admired facial feature, the so-called "Grecian profile."[aa] These faces are often contrasted with that of a warrior.[133] Salaì is often depicted in fancy-dress costume. Leonardo is known to have designed sets for pageants with which these may be associated. Other, often meticulous, drawings show studies of drapery. A marked development in Leonardo's ability to draw drapery occurred in his early works. Another often-reproduced drawing is a macabre sketch that was done by Leonardo in Florence in 1479 showing the body of Bernardo Baroncelli, hanged in connection with the murder of Giuliano, brother of Lorenzo de' Medici, in the Pazzi conspiracy.[133] In his notes, Leonardo recorded the colours of the robes that Baroncelli was wearing when he died.
+Renaissance humanism recognised no mutually exclusive polarities between the sciences and the arts, and Leonardo's studies in science and engineering are sometimes considered as impressive and innovative as his artistic work.[38] These studies were recorded in 13,000 pages of notes and drawings, which fuse art and natural philosophy (the forerunner of modern science). They were made and maintained daily throughout Leonardo's life and travels, as he made continual observations of the world around him.[38]
+Leonardo's notes and drawings display an enormous range of interests and preoccupations, some as mundane as lists of groceries and people who owed him money and some as intriguing as designs for wings and shoes for walking on water. There are compositions for paintings, studies of details and drapery, studies of faces and emotions, of animals, babies, dissections, plant studies, rock formations, whirlpools, war machines, flying machines and architecture.[38]
+These notebooks—originally loose papers of different types and sizes, were largely entrusted to Leonardo's pupil and heir Francesco Melzi after the master's death.[137] These were to be published, a task of overwhelming difficulty because of its scope and Leonardo's idiosyncratic writing.[138] Some of Leonardo's drawings were copied by an anonymous Milanese artist for a planned treatise on art c. 1570.[139] After Melzi's death in 1570, the collection passed to his son, the lawyer Orazio, who initially took little interest in the journals.[137] In 1587, a Melzi household tutor named Lelio Gavardi took 13 of the manuscripts to Pisa; there, the architect Giovanni Magenta reproached Gavardi for having taken the manuscripts illicitly and returned them to Orazio. Having many more such works in his possession, Orazio gifted the volumes to Magenta. News spread of these lost works of Leonardo's, and Orazio retrieved seven of the 13 manuscripts, which he then gave to Pompeo Leoni for publication in two volumes; one of these was the Codex Atlanticus. The other six works had been distributed to a few others.[140] After Orazio's death, his heirs sold the rest of Leonardo's possessions, and thus began their dispersal.[141]
+Some works have found their way into major collections such as the Royal Library at Windsor Castle, the Louvre, the Biblioteca Nacional de España, the Victoria and Albert Museum, the Biblioteca Ambrosiana in Milan, which holds the 12-volume Codex Atlanticus, and the British Library in London, which has put a selection from the Codex Arundel (BL Arundel MS 263) online.[142] Works have also been at Holkham Hall, the Metropolitan Museum of Art, and in the private hands of John Nicholas Brown I and Robert Lehman.[137] The Codex Leicester is the only privately owned major scientific work of Leonardo; it is owned by Bill Gates and displayed once a year in different cities around the world.
+Most of Leonardo's writings are in mirror-image cursive.[143][44] Since Leonardo wrote with his left hand, it was probably easier for him to write from right to left.[144][ab] Leonardo used a variety of shorthand and symbols, and states in his notes that he intended to prepare them for publication.[143] In many cases a single topic is covered in detail in both words and pictures on a single sheet, together conveying information that would not be lost if the pages were published out of order.[147] Why they were not published during Leonardo's lifetime is unknown.[38]
+Leonardo's approach to science was observational: he tried to understand a phenomenon by describing and depicting it in utmost detail and did not emphasise experiments or theoretical explanation. Since he lacked formal education in Latin and mathematics, contemporary scholars mostly ignored Leonardo the scientist, although he did teach himself Latin. In the 1490s he studied mathematics under Luca Pacioli and prepared a series of drawings of regular solids in a skeletal form to be engraved as plates for Pacioli's book Divina proportione, published in 1509.[38] While living in Milan, he studied light from the summit of Monte Rosa.[63] Scientific writings in his notebook on fossils have been considered as influential on early palaeontology[148][149] and he has been called the father of ichnology.[150]
+The content of his journals suggest that he was planning a series of treatises on a variety of subjects. A coherent treatise on anatomy is said to have been observed during a visit by Cardinal Louis d'Aragon's secretary in 1517.[151] Aspects of his work on the studies of anatomy, light and the landscape were assembled for publication by Melzi and eventually published as A Treatise on Painting in France and Italy in 1651 and Germany in 1724,[152] with engravings based upon drawings by the Classical painter Nicolas Poussin.[153] According to Arasse, the treatise, which in France went into 62 editions in fifty years, caused Leonardo to be seen as "the precursor of French academic thought on art."[38]
+While Leonardo's experimentation followed scientific methods, a recent and exhaustive analysis of Leonardo as a scientist by Fritjof Capra argues that Leonardo was a fundamentally different kind of scientist from Galileo, Newton and other scientists who followed him in that, as a "Renaissance Man", his theorising and hypothesising integrated the arts and particularly painting.[154][page needed]
+Leonardo started his study in the anatomy of the human body under the apprenticeship of Andrea del Verrocchio, who demanded that his students develop a deep knowledge of the subject.[155] As an artist, he quickly became master of topographic anatomy, drawing many studies of muscles, tendons and other visible anatomical features.
+As a successful artist, Leonardo was given permission to dissect human corpses at the Hospital of Santa Maria Nuova in Florence and later at hospitals in Milan and Rome. From 1510 to 1511 he collaborated in his studies with the doctor Marcantonio della Torre. Leonardo made over 240 detailed drawings and wrote about 13,000 words towards a treatise on anatomy.[156] Only a small amount of the material on anatomy was published in Leonardo's Treatise on painting.[138] During the time that Melzi was ordering the material into chapters for publication, they were examined by a number of anatomists and artists, including Vasari, Cellini and Albrecht Dürer, who made a number of drawings from them.[138]
+Leonardo's anatomical drawings include many studies of the human skeleton and its parts, and of muscles and sinews. He studied the mechanical functions of the skeleton and the muscular forces that are applied to it in a manner that prefigured the modern science of biomechanics.[157] He drew the heart and vascular system, the sex organs and other internal organs, making one of the first scientific drawings of a fetus in utero.[133] The drawings and notation are far ahead of their time, and if published would undoubtedly have made a major contribution to medical science.[156]
+Leonardo also closely observed and recorded the effects of age and of human emotion on the physiology, studying in particular the effects of rage. He drew many figures who had significant facial deformities or signs of illness.[38][133] Leonardo also studied and drew the anatomy of many animals, dissecting cows, birds, monkeys, bears, and frogs, and comparing in his drawings their anatomical structure with that of humans. He also made a number of studies of horses.[133]
+Leonardo's dissections and documentation of muscles, nerves, and vessels helped to describe the physiology and mechanics of movement. He attempted to identify the source of 'emotions' and their expression. He found it difficult to incorporate the prevailing system and theories of bodily humours, but eventually he abandoned these physiological explanations of bodily functions. He made the observations that humours were not located in cerebral spaces or ventricles. He documented that the humours were not contained in the heart or the liver, and that it was the heart that defined the circulatory system. He was the first to define atherosclerosis and liver cirrhosis. He created models of the cerebral ventricles with the use of melted wax and constructed a glass aorta to observe the circulation of blood through the aortic valve by using water and grass seed to watch flow patterns. Vesalius published his work on anatomy and physiology in De humani corporis fabrica in 1543.[158]
+During his lifetime, Leonardo was also valued as an engineer. With the same rational and analytical approach that moved him to represent the human body and to investigate anatomy, Leonardo studied and designed a bewildering number of machines and devices. He drew their “anatomy” with unparalleled mastery, producing the first form of the modern technical drawing, including a perfected "exploded view" technique, to represent internal components. Those studies and projects collected in his codices fill more than 5,000 pages.[159] In a letter of 1482 to the lord of Milan Ludovico il Moro, he wrote that he could create all sorts of machines both for the protection of a city and for siege. When he fled from Milan to Venice in 1499, he found employment as an engineer and devised a system of moveable barricades to protect the city from attack. In 1502, he created a scheme for diverting the flow of the Arno river, a project on which Niccolò Machiavelli also worked.[160][161] He continued to contemplate the canalization of Lombardy's plains while in Louis XII's company[63] and of the Loire and its tributaries in the company of Francis I.[162] Leonardo's journals include a vast number of inventions, both practical and impractical. They include musical instruments, a mechanical knight, hydraulic pumps, reversible crank mechanisms, finned mortar shells, and a steam cannon.[27][38]
+Leonardo was fascinated by the phenomenon of flight for much of his life, producing many studies, including Codex on the Flight of Birds (c. 1505), as well as plans for several flying machines, such as a flapping ornithopter and a machine with a helical rotor.[38] A 2003 documentary by British television station Channel Four, titled Leonardo's Dream Machines, various designs by Leonardo, such as a parachute and a giant crossbow, were interpreted and constructed.[163][164] Some of those designs proved successful, whilst others fared less well when tested.
+Research performed by Marc van den Broek revealed older prototypes for more than 100 inventions that are ascribed to Leonardo. Similarities between Leonardo´s illustrations and drawings from the Middle Ages and from Ancient Greece and Rome, the Chinese and Persian Empires, and Egypt suggest that a large portion of Leonardo's inventions had been conceived before his lifetime. Leonardo´s innovation was to combine different functions from existing drafts and set them into scenes that illustrated their utility. By reconstituting technical inventions he created something new.[165]
+Leonardo's fame within his own lifetime was such that the King of France carried him away like a trophy, and was claimed to have supported him in his old age and held him in his arms as he died. Interest in Leonardo and his work has never diminished. Crowds still queue to see his best-known artworks, T-shirts still bear his most famous drawing, and writers continue to hail him as a genius while speculating about his private life, as well as about what one so intelligent actually believed in.[38]
+The continued admiration that Leonardo commanded from painters, critics and historians is reflected in many other written tributes. Baldassare Castiglione, author of Il Cortegiano (The Courtier), wrote in 1528: "...Another of the greatest painters in this world looks down on this art in which he is unequalled..."[166] while the biographer known as "Anonimo Gaddiano" wrote, c. 1540: "His genius was so rare and universal that it can be said that nature worked a miracle on his behalf..."[167]
+Giorgio Vasari, in the enlarged edition of Lives of the Artists (1568)[168] introduced his chapter on Leonardo with the following words:
+In the normal course of events many men and women are born with remarkable talents; but occasionally, in a way that transcends nature, a single person is marvellously endowed by Heaven with beauty, grace and talent in such abundance that he leaves other men far behind, all his actions seem inspired and indeed everything he does clearly comes from God rather than from human skill. Everyone acknowledged that this was true of Leonardo da Vinci, an artist of outstanding physical beauty, who displayed infinite grace in everything that he did and who cultivated his genius so brilliantly that all problems he studied he solved with ease.
+The 19th century brought a particular admiration for Leonardo's genius, causing Henry Fuseli to write in 1801: "Such was the dawn of modern art, when Leonardo da Vinci broke forth with a splendour that distanced former excellence: made up of all the elements that constitute the essence of genius..."[169] This is echoed by A.E. Rio who wrote in 1861: "He towered above all other artists through the strength and the nobility of his talents."[170]
+By the 19th century, the scope of Leonardo's notebooks was known, as well as his paintings. Hippolyte Taine wrote in 1866: "There may not be in the world an example of another genius so universal, so incapable of fulfilment, so full of yearning for the infinite, so naturally refined, so far ahead of his own century and the following centuries."[171]
+Art historian Bernard Berenson wrote in 1896: "Leonardo is the one artist of whom it may be said with perfect literalness: Nothing that he touched but turned into a thing of eternal beauty. Whether it be the cross section of a skull, the structure of a weed, or a study of muscles, he, with his feeling for line and for light and shade, forever transmuted it into life-communicating values."[172]
+The interest in Leonardo's genius has continued unabated; experts study and translate his writings, analyse his paintings using scientific techniques, argue over attributions and search for works which have been recorded but never found.[173] Liana Bortolon, writing in 1967, said: "Because of the multiplicity of interests that spurred him to pursue every field of knowledge...Leonardo can be considered, quite rightly, to have been the universal genius par excellence, and with all the disquieting overtones inherent in that term. Man is as uncomfortable today, faced with a genius, as he was in the 16th century. Five centuries have passed, yet we still view Leonardo with awe."[27]
+Twenty-first-century author Walter Isaacson based much of his biography of Leonardo[104] on thousands of notebook entries, studying the personal notes, sketches, budget notations, and musings of the man whom he considers the greatest of innovators. Isaacson was surprised to discover a "fun, joyous" side of Leonardo in addition to his limitless curiosity and creative genius.[174]
+On the 500th anniversary of Leonardo's death, the Louvre in Paris arranged for the largest ever single exhibit of his work, called Leonardo, between November 2019 and February 2020. The exhibit includes over 100 paintings, drawings and notebooks. Eleven of the paintings that Leonardo completed in his lifetime were included. Five of these are owned by the Louvre, but the Mona Lisa was not included because it is in such great demand among general visitors to the Louvre; it remains on display in its gallery. Vitruvian Man, however, is on display following a legal battle with its owner, the Gallerie dell'Accademia in Venice. Salvator Mundi was also not included because its Saudi owner did not agree to lease the work.[175][176]
+Much of the Collegiate Church of Saint Florentin at the Château d'Amboise, where Leonardo was buried, was damaged during the French Revolution, leading to the church's demolition in 1802.[177] Some of the graves were destroyed in the process, scattering the bones interred there and thereby leaving the whereabouts of Leonardo's remains subject to dispute. While excavating the site in 1863, fine-arts inspector general Arsène Houssaye found a partially complete skeleton with a bronze ring on one finger, some white hair, and stone fragments bearing the inscriptions "EO," "AR," "DUS," and "VINC"—interpreted as forming "Leonardus Vinci".[177][89][178][179] A silver shield found near the bones depicts a beardless Francis I, corresponding to the king's appearance during Leonardo's lifetime, and the skull's inclusion of eight teeth corresponds to someone of approximately the appropriate age.[178] The unusually large skull led Houssaye to believe that he had located Leonardo's remains,[179] but he thought the skeleton seemed too short.[178] Other art historians say that the 1.73 metres (5.7 ft) tall skeleton may well be Leonardo's.[180]
+The remains, except for the ring and a lock of hair which Houssaye kept,[89] were brought to Paris in a lead box, where the skull was allegedly presented to Napoleon III,[178] before being returned to the Château d'Amboise and re-interred in the Chapel of Saint Hubert in 1874.[177] A new memorial tombstone was added by sculptor Francesco La Monaca in the 1930s.[181] Reflecting doubts about the attribution, a plaque above the tomb states that the remains are only presumed to be those of Leonardo.[177] It has since been theorized that the folding of the skeleton's right arm over the head may correspond to the paralysis of Leonardo's right hand.[178][77][83]
+In 2016, it was announced that DNA tests would be conducted to determine whether the attribution is correct.[177] The DNA of the remains will be compared to that of samples collected from Leonardo's work and his half-brother Domenico's descendants;[177] it may also be sequenced.[182] The lock of hair and ring, now in a private US collection,[ac] were displayed in Vinci beginning on 2 May 2019, the 500th anniversary of the artist's death.[183][89]
+Salvator Mundi, a painting by Leonardo depicting Jesus holding an orb, sold for a world record US$450.3 million at a Christie's auction in New York, 15 November 2017.[184] The highest known sale price for any artwork was previously US$300 million, for Willem de Kooning's Interchange, which was sold privately in September 2015.[185] The highest price previously paid for a work of art at auction was for Pablo Picasso's Les Femmes d'Alger, which sold for US$179.4 million in May 2015 at Christie's New York.[185]
+Early sources
+Modern sources

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+Leonardo da Vinci (English: /ˌliːəˈnɑːrdoʊ də ˈvɪntʃi, ˌliːoʊˈ-, ˌleɪoʊˈ-/;[a] 14/15 April 1452[b] – 2 May 1519),[4] was an Italian polymath of the High Renaissance who is widely considered one of the greatest painters of all time (despite less than 25 of his paintings having survived).[c] The Mona Lisa is the most famous of Leonardo's works and the most famous portrait ever made.[5] The Last Supper is the most reproduced religious painting of all time[6] and his Vitruvian Man drawing is also regarded as a cultural icon.[7] He is also known for his his notebooks, in which he made drawings and notes on science and invention; these involve a variety of subjects including anatomy, cartography, painting, and paleontology. Leonardo's collective works compose a contribution to later generations of artists rivalled only by that of his contemporary Michelangelo.[8]
+Properly named Leonardo di ser Piero da Vinci,[d] Leonardo was born out of wedlock to a notary, Piero da Vinci, and a peasant woman, Caterina, in Vinci, in the region of Florence, Italy. Leonardo was educated in the studio of the renowned Italian painter Andrea del Verrocchio. Much of his earlier working life was spent in the service of Ludovico il Moro in Milan, and he later worked in Rome, Bologna and Venice. He spent his last three years in France, where he died in 1519.
+Although he had no formal academic training,[9] many historians and scholars regard Leonardo as the prime exemplar of the "Universal Genius" or "Renaissance Man", an individual of "unquenchable curiosity" and "feverishly inventive imagination."[6] He is widely considered one of the most diversely talented individuals ever to have lived.[10] According to art historian Helen Gardner, the scope and depth of his interests were without precedent in recorded history, and "his mind and personality seem to us superhuman, while the man himself mysterious and remote."[6] Scholars interpret his view of the world as being based in logic, though the empirical methods he used were unorthodox for his time.[11]
+Leonardo is revered for his technological ingenuity. He conceptualized flying machines, a type of armoured fighting vehicle, concentrated solar power, an adding machine,[12] and the double hull. Relatively few of his designs were constructed or even feasible during his lifetime, as the modern scientific approaches to metallurgy and engineering were only in their infancy during the Renaissance. Some of his smaller inventions, however, entered the world of manufacturing unheralded, such as an automated bobbin winder and a machine for testing the tensile strength of wire. He is also sometimes credited with the inventions of the parachute, helicopter, and tank.[13][14] He made substantial discoveries in anatomy, civil engineering, geology, optics, and hydrodynamics, but he did not publish his findings and they had little to no direct influence on subsequent science.[15]
+Leonardo was born on 14/15 April 1452[b] in the Tuscan hill town of Vinci, in the lower valley of the Arno river in the territory of the Medici-ruled Republic of Florence.[18] He was the out-of-wedlock son of Messer Piero Fruosino di Antonio da Vinci, a wealthy Florentine legal notary, and a peasant named Caterina,[e] identified as Caterina Buti del Vacca and more recently as Caterina di Meo Lippi by historian Martin Kemp. There have been many theories regarding Leonardo's mother's identity, including that she was a slave of foreign origin or an impoverished local youth.[17][20][21][22][f] Leonardo had no surname in the modern sense—da Vinci simply meaning "of Vinci"; his full birth name was Lionardo di ser Piero da Vinci,[2][25] meaning "Leonardo, (son) of ser Piero (from) Vinci."[18][d]
+Leonardo spent his first years in the hamlet of Anchiano in the home of his mother, and from at least 1457 lived in the household of his father, grandparents and uncle in the small town of Vinci.[26] His father had married a 16-year-old girl named Albiera Amadori, who loved Leonardo but died young[27] in 1465 without children. In 1468, when Leonardo was 16, his father married again to 20-year-old Francesca Lanfredini, who also died without children. Piero's legitimate heirs were born from his third wife Margherita di Guglielmo, who gave birth to six children, and his fourth and final wife, Lucrezia Cortigiani, who bore him another six heirs.[28][29]
+In all, Leonardo had 12 half-siblings, who were much younger than he was (the last was born when Leonardo was 40 years old) and with whom he had very little contact.[g]
+Leonardo received an informal education in Latin, geometry and mathematics. In later life, Leonardo recorded few distinct childhood incidents. One was of a kite coming to his cradle and opening his mouth with its tail; he regarded this as an omen of his writing on the subject.[31][32] The second occurred while he was exploring in the mountains: he discovered a cave and was both terrified that some great monster might lurk there and driven by curiosity to find out what was inside.[27] He also seems to have remembered some of his childhood observations of water, writing and crossing out the name of his hometown in one of his notebooks on the formation of rivers.[26]
+Leonardo's early life has been the subject of historical conjecture.[33] Vasari, the 16th-century biographer of Renaissance painters, tells a story of Leonardo as a very young man: A local peasant made himself a round shield and requested that Ser Piero have it painted for him. Leonardo, inspired by the story of Medusa, responded with a painting of a monster spitting fire that was so terrifying that his father bought a different shield to give to the peasant and sold Leonardo's to a Florentine art dealer for 100 ducats, who in turn sold it to the Duke of Milan.[34]
+In the mid-1460s, Leonardo's family moved to Florence, and around the age of 14,[30] he became a garzone (studio boy) in the workshop of Verrocchio, who was the leading Florentine painter and sculptor of his time.[35] Leonardo became an apprentice by the age of 17 and remained in training for seven years.[36][37] Other famous painters apprenticed in the workshop or associated with it include Ghirlandaio, Perugino, Botticelli, and Lorenzo di Credi.[27][38] Leonardo was exposed to both theoretical training and a wide range of technical skills,[39] including drafting, chemistry, metallurgy, metal working, plaster casting, leather working, mechanics, and wood-work, as well as the artistic skills of drawing, painting, sculpting, and modelling.[40][h]
+Much of the painting in Verrocchio's workshop was done by his employees. According to Vasari, Leonardo collaborated with Verrocchio on his The Baptism of Christ, painting the young angel holding Jesus' robe in a manner that was so far superior to his master's that Verrocchio put down his brush and never painted again, although this is believed to be an apocryphal story.[41] Close examination reveals areas of the work that have been painted or touched-up over the tempera, using the new technique of oil paint, including the landscape, the rocks seen through the brown mountain stream, and much of the figure of Jesus, bearing witness to the hand of Leonardo.[42] Leonardo may have been the model for two works by Verrocchio: the bronze statue of David in the Bargello, and the Archangel Raphael in Tobias and the Angel.[20]
+By 1472, at the age of 20, Leonardo qualified as a master in the Guild of Saint Luke, the guild of artists and doctors of medicine,[i] but even after his father set him up in his own workshop, his attachment to Verrocchio was such that he continued to collaborate and live with him.[27][43] Leonardo's earliest known dated work is a 1473 pen-and-ink drawing of the Arno valley,[38] which has been cited as the first "pure" landscape in the Occident.[j][44] According to Vasari, the young Leonardo was the first to suggest making the Arno river a navigable channel between Florence and Pisa.[45]
+In January 1478, Leonardo received an independent commission to paint an altarpiece for the Chapel of St. Bernard in the Palazzo Vecchio,[46] an indication of his independence from Verrocchio's studio. One anonymous writer claims that in 1480, Leonardo was living with the Medici and often worked in the garden of the Piazza San Marco, Florence, where a Neoplatonic academy of artists, poets and philosophers organized by the Medici met.[20] In March 1481, he received a commission from the monks of San Donato in Scopeto for The Adoration of the Magi.[47]
+Neither of these initial commissions were completed, being abandoned when Leonardo went to offer his services to Duke of Milan Ludovico Sforza. In 1482, he casted a silver stringed instrument[k] from a horse's skull and ram horns to bring to Sforza,[48] whom he wrote a letter describing the diverse things that he could achieve in the fields of engineering and weapon design, and mentioning that he could paint.[38][48][49]
+Leonardo worked in Milan from 1482 until 1499. He was commissioned to paint the Virgin of the Rocks for the Confraternity of the Immaculate Conception and The Last Supper for the monastery of Santa Maria delle Grazie.[51] In the spring of 1485, Leonardo travelled to Hungary on behalf of Sforza to meet king Matthias Corvinus, and was commissioned by him to paint a Madonna.[52]
+Leonardo was employed on many other projects for Sforza, including the preparation of floats and pageants for special occasions, a drawing and wooden model for a competition to design the cupola for Milan Cathedral (which he withdrew),[53] and a model for a huge equestrian monument to Ludovico's predecessor Francesco Sforza. This would have surpassed in size the only two large equestrian statues of the Renaissance, Donatello's Gattamelata in Padua and Verrocchio's Bartolomeo Colleoni in Venice, and became known as the Gran Cavallo.[38] Leonardo completed a model for the horse and made detailed plans for its casting,[38] but in November 1494, Ludovico gave the bronze to his brother-in-law to be used for a cannon to defend the city from Charles VIII.[38]
+With Ludovico Sforza overthrown at the dawn of the Second Italian War, Leonardo, with his assistant Salaì and friend, the mathematician Luca Pacioli, fled Milan for Venice.[54] There, he was employed as a military architect and engineer, devising methods to defend the city from naval attack.[27] On his return to Florence in 1500, he and his household were guests of the Servite monks at the monastery of Santissima Annunziata and were provided with a workshop where, according to Vasari, Leonardo created the cartoon of The Virgin and Child with St Anne and St John the Baptist, a work that won such admiration that "men and women, young and old" flocked to see it "as if they were attending a great festival."[55][l]
+In Cesena in 1502, Leonardo entered the service of Cesare Borgia, the son of Pope Alexander VI, acting as a military architect and engineer and travelling throughout Italy with his patron.[54] Leonardo created a map of Cesare Borgia's stronghold, a town plan of Imola in order to win his patronage. Maps were extremely rare at the time and it would have seemed like a new concept. Upon seeing it, Cesare hired Leonardo as his chief military engineer and architect. Later in the year, Leonardo produced another map for his patron, one of Chiana Valley, Tuscany, so as to give his patron a better overlay of the land and greater strategic position. He created this map in conjunction with his other project of constructing a dam from the sea to Florence, in order to allow a supply of water to sustain the canal during all seasons.
+Leonardo had left Borgia's service and returned to Florence by early 1503,[57] where he rejoined the Guild of Saint Luke on 18 October of that year. By this same month, Leonardo had begun working on a portrait of Lisa del Giocondo, the model for the Mona Lisa,[58][59] which he would continue working on until his twilight years. In January 1504, he was part of a committee formed to recommend where Michelangelo's statue of David should be placed.[60] He then spent two years in Florence designing and painting a mural of The Battle of Anghiari for the Signoria,[54] with Michelangelo designing its companion piece, The Battle of Cascina.[m]
+In 1506, Leonardo was summoned to Milan by Charles II d'Amboise, the acting French governor of the city. The Council of Florence wished Leonardo to return promptly to finish The Battle of Anghiari, but he was given leave at the behest of Louis XII, who considered commissioning the artist to make some portraits.[63] Leonardo may have commenced a project for an equestrian figure of d'Amboise;[64] a wax model survives and, if genuine, is the only extant example of Leonardo's sculpture. Leonardo was otherwise free to pursue his scientific interests.[63] Many of Leonardo's most prominent pupils either knew or worked with him in Milan,[27] including Bernardino Luini, Giovanni Antonio Boltraffio, and Marco d'Oggiono.
+In 1507, Leonardo was in Florence sorting out a dispute with his brothers over the estate of his father, who had died in 1504. By 1508, Leonardo was back in Milan, living in his own house in Porta Orientale in the parish of Santa Babila.[65]
+In 1512, Leonardo was working on plans for an equestrian monument for Gian Giacomo Trivulzio, but this was prevented by an invasion of a confederation of Swiss, Spanish and Venetian forces, which drove the French from Milan. Leonardo stayed in the city, spending several months in 1513 at the Medici's Vaprio d'Adda villa.[67] In March of that year, Lorenzo de' Medici's son Giovanni assumed the papacy (as Leo X); Leonardo went to Rome that September, where he was received by the pope's brother Giuliano.[67] From September 1513 to 1516, Leonardo spent much of his time living in the Belvedere Courtyard in the Apostolic Palace, where Michelangelo and Raphael were both active.[65] Leonardo was given an allowance of 33 ducats a month, and according to Vasari, decorated a lizard with scales dipped in quicksilver.[68] The pope gave him a painting commission of unknown subject matter, but cancelled it when the artist set about developing a new kind of varnish.[68][n] Leonardo became ill, in what may have been the first of multiple strokes leading to his death.[68] He practiced botany in the Gardens of Vatican City, and was commissioned to make plans for the pope's proposed draining of the Pontine Marshes.[69] He also dissected cadavers, making notes for a treatise on vocal cords;[70] these he gave to an official in hopes of regaining the pope's favor, but was unsuccessful.[68]
+In October 1515, King Francis I of France recaptured Milan.[47] Leonardo was present at the 19 December meeting of Francis I and Leo X, which took place in Bologna.[27][71][72] In 1516, Leonardo entered Francis' service, being given the use of the manor house Clos Lucé, near the king's residence at the royal Château d'Amboise. Being frequently visited by Francis, he drew plans for an immense castle town the king intended to erect at Romorantin, and made a mechanical lion, which during a pageant walked toward the king and—upon being struck by a wand—opened its chest to reveal a cluster of lilies.[73][74][o] Leonardo was accompanied during this time by his friend and apprentice, Francesco Melzi, and supported by a pension totalling 10,000 scudi.[65] At some point, Melzi drew a portrait of Leonardo; the only others known from his lifetime were a sketch by an unknown assistant on the back of one of Leonardo's studies (c. 1517)[76] and a drawing by Giovanni Ambrogio Figino depicting an elderly Leonardo with his right arm assuaged by cloth.[77][p][q] The latter, in addition to the record of an October 1517 visit by Louis d'Aragon,[r] confirms an account of Leonardo's right hand being paralytic at the age of 65,[80] which may indicate why he left works such as the Mona Lisa unfinished.[78][81][82] He continued to work at some capacity until eventually becoming ill and bedridden for several months.[80]
+Leonardo died at Clos Lucé on 2 May 1519 at the age of 67, possibly of a stroke.[83][82] Francis I had become a close friend. Vasari describes Leonardo as lamenting on his deathbed, full of repentance, that "he had offended against God and men by failing to practice his art as he should have done."[84] Vasari states that in his last days, Leonardo sent for a priest to make his confession and to receive the Holy Sacrament.[85] Vasari also records that the king held Leonardo's head in his arms as he died, although this story may be legend rather than fact.[s][t] In accordance with his will, sixty beggars carrying tapers followed Leonardo's casket.[87][u] Melzi was the principal heir and executor, receiving, as well as money, Leonardo's paintings, tools, library and personal effects. Leonardo also remembered his other long-time pupil and companion, Salaì, and his servant Baptista de Vilanis, who each received half of Leonardo's vineyards.[88] His brothers received land, and his serving woman received a fur-lined cloak. On 12 August 1519, Leonardo's remains were interred in the Collegiate Church of Saint Florentin at the Château d'Amboise.[89]
+Some 20 years after Leonardo's death, Francis was reported by the goldsmith and sculptor Benvenuto Cellini as saying: "There had never been another man born in the world who knew as much as Leonardo, not so much about painting, sculpture and architecture, as that he was a very great philosopher."[90]
+Florence at the time of Leonardo's youth was the centre of Christian Humanist thought and culture.[35] Leonardo commenced his apprenticeship with Verrocchio in 1466, the year that Verrocchio's master, the great sculptor Donatello, died.[v] The painter Uccello, whose early experiments with perspective were to influence the development of landscape painting, was a very old man. The painters Piero della Francesca and Filippo Lippi, sculptor Luca della Robbia, and architect and writer Leon Battista Alberti were in their sixties. The successful artists of the next generation were Leonardo's teacher Verrocchio, Antonio del Pollaiuolo, and the portrait sculptor Mino da Fiesole.[91][92][93][94]
+Leonardo's youth was spent in a Florence that was ornamented by the works of these artists and by Donatello's contemporaries, Masaccio, whose figurative frescoes were imbued with realism and emotion; and Ghiberti, whose Gates of Paradise, gleaming with gold leaf, displayed the art of combining complex figure compositions with detailed architectural backgrounds. Piero della Francesca had made a detailed study of perspective,[95] and was the first painter to make a scientific study of light. These studies and Alberti's treatise De pictura[96] were to have a profound effect on younger artists and in particular on Leonardo's own observations and artworks.[91][93][94]
+A prevalent tradition in Florence was the small altarpiece of the Virgin and Child. Many of these were created in tempera or glazed terracotta by the workshops of Filippo Lippi, Verrocchio and the prolific della Robbia family.[91] Leonardo's early Madonnas such as The Madonna with a carnation and the Benois Madonna followed this tradition while showing idiosyncratic departures, particularly in the latter in which the Virgin is set at an oblique angle to the picture space with the Christ Child at the opposite angle. This compositional theme was to emerge in Leonardo's later paintings such as The Virgin and Child with St. Anne.[27]
+Leonardo was a contemporary of Botticelli, Domenico Ghirlandaio and Perugino, who were all slightly older than he was.[92] He would have met them at the workshop of Verrocchio, with whom they had associations, and at the Academy of the Medici.[27] Botticelli was a particular favourite of the Medici family, and thus his success as a painter was assured. Ghirlandaio and Perugino were both prolific and ran large workshops. They competently delivered commissions to well-satisfied patrons who appreciated Ghirlandaio's ability to portray the wealthy citizens of Florence within large religious frescoes, and Perugino's ability to deliver a multitude of saints and angels of unfailing sweetness and innocence.[91]
+These three were among those commissioned to paint the walls of the Sistine Chapel, the work commencing with Perugino's employment in 1479. Leonardo was not part of this prestigious commission. His first significant commission, The Adoration of the Magi for the Monks of Scopeto, was never completed.[27]
+In 1476, during the time of Leonardo's association with Verrocchio's workshop, the Portinari Altarpiece by Hugo van der Goes arrived in Florence, bringing from Northern Europe new painterly techniques that were to profoundly affect Leonardo, Ghirlandaio, Perugino and others.[92] In 1479, the Sicilian painter Antonello da Messina, who worked exclusively in oils, travelled north on his way to Venice, where the leading painter Giovanni Bellini adopted the technique of oil painting, quickly making it the preferred method in Venice. Leonardo was also later to visit Venice.[92][94]
+Like the two contemporary architects Donato Bramante (who designed the Belvedere Courtyard) and Antonio da Sangallo the Elder, Leonardo experimented with designs for centrally planned churches, a number of which appear in his journals, as both plans and views, although none was ever realised.[92][97]
+Leonardo's political contemporaries were Lorenzo de' Medici (il Magnifico), who was three years older, and his younger brother Giuliano, who was slain in the Pazzi conspiracy in 1478. Leonardo was sent as an ambassador by the Medici court to Ludovico il Moro, who ruled Milan between 1479 and 1499.[92]
+With Alberti, Leonardo visited the home of the Medici and through them came to know the older Humanist philosophers of whom Marsiglio Ficino, proponent of Neo Platonism; Cristoforo Landino, writer of commentaries on Classical writings, and John Argyropoulos, teacher of Greek and translator of Aristotle were the foremost. Also associated with the Academy of the Medici was Leonardo's contemporary, the brilliant young poet and philosopher Pico della Mirandola.[92][94][87] Leonardo later wrote in the margin of a journal, "The Medici made me and the Medici destroyed me." While it was through the action of Lorenzo that Leonardo received his employment at the court of Milan, it is not known exactly what Leonardo meant by this cryptic comment.[27]
+Although usually named together as the three giants of the High Renaissance, Leonardo, Michelangelo and Raphael were not of the same generation. Leonardo was 23 when Michelangelo was born and 31 when Raphael was born.[92] Raphael died at the age of 37 in 1520, the year after Leonardo died, but Michelangelo went on creating for another 45 years.[93][94]
+Within Leonardo's lifetime, his extraordinary powers of invention, his "outstanding physical beauty," "infinite grace," "great strength and generosity," "regal spirit and tremendous breadth of mind," as described by Vasari,[98] as well as all other aspects of his life, attracted the curiosity of others. One such aspect was his love for animals, likely including vegetarianism and according to Vasari, a habit of purchasing caged birds and releasing them.[99][100]
+Leonardo had many friends who are now renowned either in their fields or for their historical significance. They included the mathematician Luca Pacioli,[101] with whom he collaborated on the book Divina proportione in the 1490s. Leonardo appears to have had no close relationships with women except for his friendship with Cecilia Gallerani and the two Este sisters, Beatrice and Isabella.[102] While on a journey that took him through Mantua, he drew a portrait of Isabella that appears to have been used to create a painted portrait, now lost.[27]
+Beyond friendship, Leonardo kept his private life secret. His sexuality has been the subject of satire, analysis, and speculation. This trend began in the mid-16th century and was revived in the 19th and 20th centuries, most notably by Sigmund Freud.[103] Leonardo's most intimate relationships were perhaps with his pupils Salaì and Melzi. Melzi, writing to inform Leonardo's brothers of his death, described Leonardo's feelings for his pupils as both loving and passionate. It has been claimed since the 16th century that these relationships were of a sexual or erotic nature. Court records of 1476, when he was aged twenty-four, show that Leonardo and three other young men were charged with sodomy in an incident involving a well-known male prostitute. The charges were dismissed for lack of evidence, and there is speculation that since one of the accused, Lionardo de Tornabuoni, was related to Lorenzo de' Medici, the family exerted its influence to secure the dismissal.[104] Since that date much has been written about his presumed homosexuality and its role in his art, particularly in the androgyny and eroticism manifested in Saint John the Baptist and Bacchus and more explicitly in a number of erotic drawings.[105]
+Gian Giacomo Caprotti da Oreno, nicknamed Salaì or Il Salaino ("The Little Unclean One," i.e., the devil), entered Leonardo's household in 1490. After only a year, Leonardo made a list of his misdemeanours, calling him "a thief, a liar, stubborn, and a glutton," after he had made off with money and valuables on at least five occasions and spent a fortune on clothes.[106] Nevertheless, Leonardo treated him with great indulgence, and he remained in Leonardo's household for the next thirty years.[107] Salaì executed a number of paintings under the name of Andrea Salaì, but although Vasari claims that Leonardo "taught him a great deal about painting,"[74] his work is generally considered to be of less artistic merit than others among Leonardo's pupils, such as Marco d'Oggiono and Boltraffio. In 1515, he painted a nude version of the Mona Lisa, known as Monna Vanna.[108] Salaì owned the Mona Lisa at the time of his death in 1524, and in his will it was assessed at 505 lire, an exceptionally high valuation for a small panel portrait.[109]
+In 1506, Leonardo took on another pupil, Count Francesco Melzi, the son of a Lombard aristocrat, who is considered to have been his favourite student. He travelled to France with Leonardo and remained with him until Leonardo's death.[27] Melzi inherited the artistic and scientific works, manuscripts, and collections of Leonardo and administered the estate.
+Despite the recent awareness and admiration of Leonardo as a scientist and inventor, for the better part of four hundred years his fame rested on his achievements as a painter. A handful of works that are either authenticated or attributed to him have been regarded as among the great masterpieces. These paintings are famous for a variety of qualities that have been much imitated by students and discussed at great length by connoisseurs and critics. By the 1490s Leonardo had already been described as a "Divine" painter.[110]
+Among the qualities that make Leonardo's work unique are his innovative techniques for laying on the paint; his detailed knowledge of anatomy, light, botany and geology; his interest in physiognomy and the way humans register emotion in expression and gesture; his innovative use of the human form in figurative composition; and his use of subtle gradation of tone. All these qualities come together in his most famous painted works, the Mona Lisa, the Last Supper, and the Virgin of the Rocks.[111]
+Leonardo first gained attention for his work on the Baptism of Christ, painted in conjunction with Verrocchio. Two other paintings appear to date from his time at Verrocchio's workshop, both of which are Annunciations. One is small, 59 centimetres (23 in) long and 14 centimetres (5.5 in) high. It is a "predella" to go at the base of a larger composition, a painting by Lorenzo di Credi from which it has become separated. The other is a much larger work, 217 centimetres (85 in) long.[112] In both Annunciations, Leonardo used a formal arrangement, like two well-known pictures by Fra Angelico of the same subject, of the Virgin Mary sitting or kneeling to the right of the picture, approached from the left by an angel in profile, with a rich flowing garment, raised wings and bearing a lily. Although previously attributed to Ghirlandaio, the larger work is now generally attributed to Leonardo.[113]
+In the smaller painting, Mary averts her eyes and folds her hands in a gesture that symbolised submission to God's will. Mary is not submissive, however, in the larger piece. The girl, interrupted in her reading by this unexpected messenger, puts a finger in her bible to mark the place and raises her hand in a formal gesture of greeting or surprise.[91] This calm young woman appears to accept her role as the Mother of God, not with resignation but with confidence. In this painting, the young Leonardo presents the humanist face of the Virgin Mary, recognising humanity's role in God's incarnation.
+In the 1480s, Leonardo received two very important commissions and commenced another work that was of ground-breaking importance in terms of composition. Two of the three were never finished, and the third took so long that it was subject to lengthy negotiations over completion and payment.
+One of these paintings was Saint Jerome in the Wilderness, which Bortolon associates with a difficult period of Leonardo's life, as evidenced in his diary: "I thought I was learning to live; I was only learning to die."[27] Although the painting is barely begun, the composition can be seen and is very unusual.[w] Jerome, as a penitent, occupies the middle of the picture, set on a slight diagonal and viewed somewhat from above. His kneeling form takes on a trapezoid shape, with one arm stretched to the outer edge of the painting and his gaze looking in the opposite direction. J. Wasserman points out the link between this painting and Leonardo's anatomical studies.[114] Across the foreground sprawls his symbol, a great lion whose body and tail make a double spiral across the base of the picture space. The other remarkable feature is the sketchy landscape of craggy rocks against which the figure is silhouetted.
+The daring display of figure composition, the landscape elements and personal drama also appear in the great unfinished masterpiece, the Adoration of the Magi, a commission from the Monks of San Donato a Scopeto. It is a complex composition, of about 250 x 250 centimetres. Leonardo did numerous drawings and preparatory studies, including a detailed one in linear perspective of the ruined classical architecture that forms part of the background. In 1482 Leonardo went to Milan at the behest of Lorenzo de' Medici in order to win favour with Ludovico il Moro, and the painting was abandoned.[20][113]
+The third important work of this period is the Virgin of the Rocks, commissioned in Milan for the Confraternity of the Immaculate Conception. The painting, to be done with the assistance of the de Predis brothers, was to fill a large complex altarpiece.[115] Leonardo chose to paint an apocryphal moment of the infancy of Christ when the infant John the Baptist, in protection of an angel, met the Holy Family on the road to Egypt. The painting demonstrates an eerie beauty as the graceful figures kneel in adoration around the infant Christ in a wild landscape of tumbling rock and whirling water.[116] While the painting is quite large, about 200×120 centimetres, it is not nearly as complex as the painting ordered by the monks of St Donato, having only four figures rather than about fifty and a rocky landscape rather than architectural details. The painting was eventually finished; in fact, two versions of the painting were finished: one remained at the chapel of the Confraternity, while Leonardo took the other to France. The Brothers did not get their painting, however, nor the de Predis their payment, until the next century.[38][54]
+Leonardo's most remarkable portrait of this period is the Lady with an Ermine, presumed to be  Cecilia Gallerani (c. 1483–1490), lover of Ludovico Sforza.[117][118] The painting is charcterised by the pose of the figure with the head turned at a very different angle to the torso, unusual at a date when many portraits were still rigidly in profile. The ermine plainly carries symbolic meaning, relating either to the sitter, or to Ludovico who belonged to the prestigious Order of the Ermine.[117]
+Leonardo's most famous painting of the 1490s is The Last Supper, commissioned for the refectory of the Convent of Santa Maria della Grazie in Milan. It represents the last meal shared by Jesus with his disciples before his capture and death, and shows the moment when Jesus has just said "one of you will betray me", and the consternation that this statement caused.[38]
+The novelist Matteo Bandello observed Leonardo at work and wrote that some days he would paint from dawn till dusk without stopping to eat and then not paint for three or four days at a time.[119] This was beyond the comprehension of the prior of the convent, who hounded him until Leonardo asked Ludovico to intervene. Vasari describes how Leonardo, troubled over his ability to adequately depict the faces of Christ and the traitor Judas, told the Duke that he might be obliged to use the prior as his model.[120]
+When finished, the painting was acclaimed as a masterpiece of design and characterization,[121] but it deteriorated rapidly, so that within a hundred years it was described by one viewer as "completely ruined."[122] Leonardo, instead of using the reliable technique of fresco, had used tempera over a ground that was mainly gesso, resulting in a surface subject to mould and to flaking.[123] Despite this, the painting remains one of the most reproduced works of art; countless copies have been made in various mediums.
+It is recorded that in 1492, Leonardo, with assistants painted the Sala delle Asse in the Sforza Castle in Milan,  with a trompe-l'œil depicting trees, with an intricate labyrinth of leaves and knots on the ceiling.[124]
+In 1505 Leonardo was commissioned to paint The Battle of Anghiari in the Salone dei Cinquecento (Hall of the Five Hundred) in the Palazzo Vecchio, Florence. Leonardo devised a dynamic composition depicting four men riding raging war horses engaged in a battle for possession of a standard, at the Battle of Anghiari in 1440.  Michelangelo was assigned the opposite wall to depict the Battle of Cascina. Leonardo's painting deteriorated rapidly and is now known from a copy by Rubens.[125]
+Among the works created by Leonardo in the 16th century is the small portrait known as the Mona Lisa or La Gioconda, the laughing one. In the present era, it is arguably the most famous painting in the world. Its fame rests, in particular, on the elusive smile on the woman's face, its mysterious quality perhaps due to the subtly shadowed corners of the mouth and eyes such that the exact nature of the smile cannot be determined. The shadowy quality for which the work is renowned came to be called "sfumato," or Leonardo's smoke. Vasari, who is generally thought to have known the painting only by repute, said that "the smile was so pleasing that it seemed divine rather than human; and those who saw it were amazed to find that it was as alive as the original."[126][x]
+Other characteristics of the painting are the unadorned dress, in which the eyes and hands have no competition from other details; the dramatic landscape background, in which the world seems to be in a state of flux; the subdued colouring; and the extremely smooth nature of the painterly technique, employing oils laid on much like tempera, and blended on the surface so that the brushstrokes are indistinguishable.[y] Vasari expressed the opinion that the manner of painting would make even "the most confident master...despair and lose heart."[129] The perfect state of preservation and the fact that there is no sign of repair or overpainting is rare in a panel painting of this date.[130]
+In the painting Virgin and Child with St. Anne, the composition again picks up the theme of figures in a landscape, which Wasserman describes as "breathtakingly beautiful"[131] and harkens back to the St Jerome picture with the figure set at an oblique angle. What makes this painting unusual is that there are two obliquely set figures superimposed. Mary is seated on the knee of her mother, St Anne. She leans forward to restrain the Christ Child as he plays roughly with a lamb, the sign of his own impending sacrifice.[38] This painting, which was copied many times, influenced Michelangelo, Raphael, and Andrea del Sarto,[132] and through them Pontormo and Correggio. The trends in composition were adopted in particular by the Venetian painters Tintoretto and Veronese.
+Leonardo was not a prolific painter, but he was a most prolific draftsman, keeping journals full of small sketches and detailed drawings recording all manner of things that took his attention. As well as the journals there exist many studies for paintings, some of which can be identified as preparatory to particular works such as The Adoration of the Magi, The Virgin of the Rocks and The Last Supper.[133]
+His earliest dated drawing is a Landscape of the Arno Valley, 1473, which shows the river, the mountains, Montelupo Castle and the farmlands beyond it in great detail.[27][133][z] According to art historian Ludwig Heydenreich, this is "The first true landscape in art."[134] Massimo Polidoro says that it was the first landscape "not to be the background of some religious scene or a portrait. It is the first [documented] time where a landscape was drawn just for the sake of it."[44]
+Among his famous drawings are the Vitruvian Man, a study of the proportions of the human body; the Head of an Angel, for The Virgin of the Rocks in the Louvre; a botanical study of Star of Bethlehem; and a large drawing (160×100 cm) in black chalk on coloured paper of The Virgin and Child with St. Anne and St. John the Baptist in the National Gallery, London.[133] This drawing employs the subtle sfumato technique of shading, in the manner of the Mona Lisa. It is thought that Leonardo never made a painting from it, the closest similarity being to The Virgin and Child with St. Anne in the Louvre.[135]
+Other drawings of interest include numerous studies generally referred to as "caricatures" because, although exaggerated, they appear to be based upon observation of live models. Vasari relates that if Leonardo saw a person with an interesting face he would follow them around all day observing them.[136] There are numerous studies of beautiful young men, often associated with Salaì, with the rare and much admired facial feature, the so-called "Grecian profile."[aa] These faces are often contrasted with that of a warrior.[133] Salaì is often depicted in fancy-dress costume. Leonardo is known to have designed sets for pageants with which these may be associated. Other, often meticulous, drawings show studies of drapery. A marked development in Leonardo's ability to draw drapery occurred in his early works. Another often-reproduced drawing is a macabre sketch that was done by Leonardo in Florence in 1479 showing the body of Bernardo Baroncelli, hanged in connection with the murder of Giuliano, brother of Lorenzo de' Medici, in the Pazzi conspiracy.[133] In his notes, Leonardo recorded the colours of the robes that Baroncelli was wearing when he died.
+Renaissance humanism recognised no mutually exclusive polarities between the sciences and the arts, and Leonardo's studies in science and engineering are sometimes considered as impressive and innovative as his artistic work.[38] These studies were recorded in 13,000 pages of notes and drawings, which fuse art and natural philosophy (the forerunner of modern science). They were made and maintained daily throughout Leonardo's life and travels, as he made continual observations of the world around him.[38]
+Leonardo's notes and drawings display an enormous range of interests and preoccupations, some as mundane as lists of groceries and people who owed him money and some as intriguing as designs for wings and shoes for walking on water. There are compositions for paintings, studies of details and drapery, studies of faces and emotions, of animals, babies, dissections, plant studies, rock formations, whirlpools, war machines, flying machines and architecture.[38]
+These notebooks—originally loose papers of different types and sizes, were largely entrusted to Leonardo's pupil and heir Francesco Melzi after the master's death.[137] These were to be published, a task of overwhelming difficulty because of its scope and Leonardo's idiosyncratic writing.[138] Some of Leonardo's drawings were copied by an anonymous Milanese artist for a planned treatise on art c. 1570.[139] After Melzi's death in 1570, the collection passed to his son, the lawyer Orazio, who initially took little interest in the journals.[137] In 1587, a Melzi household tutor named Lelio Gavardi took 13 of the manuscripts to Pisa; there, the architect Giovanni Magenta reproached Gavardi for having taken the manuscripts illicitly and returned them to Orazio. Having many more such works in his possession, Orazio gifted the volumes to Magenta. News spread of these lost works of Leonardo's, and Orazio retrieved seven of the 13 manuscripts, which he then gave to Pompeo Leoni for publication in two volumes; one of these was the Codex Atlanticus. The other six works had been distributed to a few others.[140] After Orazio's death, his heirs sold the rest of Leonardo's possessions, and thus began their dispersal.[141]
+Some works have found their way into major collections such as the Royal Library at Windsor Castle, the Louvre, the Biblioteca Nacional de España, the Victoria and Albert Museum, the Biblioteca Ambrosiana in Milan, which holds the 12-volume Codex Atlanticus, and the British Library in London, which has put a selection from the Codex Arundel (BL Arundel MS 263) online.[142] Works have also been at Holkham Hall, the Metropolitan Museum of Art, and in the private hands of John Nicholas Brown I and Robert Lehman.[137] The Codex Leicester is the only privately owned major scientific work of Leonardo; it is owned by Bill Gates and displayed once a year in different cities around the world.
+Most of Leonardo's writings are in mirror-image cursive.[143][44] Since Leonardo wrote with his left hand, it was probably easier for him to write from right to left.[144][ab] Leonardo used a variety of shorthand and symbols, and states in his notes that he intended to prepare them for publication.[143] In many cases a single topic is covered in detail in both words and pictures on a single sheet, together conveying information that would not be lost if the pages were published out of order.[147] Why they were not published during Leonardo's lifetime is unknown.[38]
+Leonardo's approach to science was observational: he tried to understand a phenomenon by describing and depicting it in utmost detail and did not emphasise experiments or theoretical explanation. Since he lacked formal education in Latin and mathematics, contemporary scholars mostly ignored Leonardo the scientist, although he did teach himself Latin. In the 1490s he studied mathematics under Luca Pacioli and prepared a series of drawings of regular solids in a skeletal form to be engraved as plates for Pacioli's book Divina proportione, published in 1509.[38] While living in Milan, he studied light from the summit of Monte Rosa.[63] Scientific writings in his notebook on fossils have been considered as influential on early palaeontology[148][149] and he has been called the father of ichnology.[150]
+The content of his journals suggest that he was planning a series of treatises on a variety of subjects. A coherent treatise on anatomy is said to have been observed during a visit by Cardinal Louis d'Aragon's secretary in 1517.[151] Aspects of his work on the studies of anatomy, light and the landscape were assembled for publication by Melzi and eventually published as A Treatise on Painting in France and Italy in 1651 and Germany in 1724,[152] with engravings based upon drawings by the Classical painter Nicolas Poussin.[153] According to Arasse, the treatise, which in France went into 62 editions in fifty years, caused Leonardo to be seen as "the precursor of French academic thought on art."[38]
+While Leonardo's experimentation followed scientific methods, a recent and exhaustive analysis of Leonardo as a scientist by Fritjof Capra argues that Leonardo was a fundamentally different kind of scientist from Galileo, Newton and other scientists who followed him in that, as a "Renaissance Man", his theorising and hypothesising integrated the arts and particularly painting.[154][page needed]
+Leonardo started his study in the anatomy of the human body under the apprenticeship of Andrea del Verrocchio, who demanded that his students develop a deep knowledge of the subject.[155] As an artist, he quickly became master of topographic anatomy, drawing many studies of muscles, tendons and other visible anatomical features.
+As a successful artist, Leonardo was given permission to dissect human corpses at the Hospital of Santa Maria Nuova in Florence and later at hospitals in Milan and Rome. From 1510 to 1511 he collaborated in his studies with the doctor Marcantonio della Torre. Leonardo made over 240 detailed drawings and wrote about 13,000 words towards a treatise on anatomy.[156] Only a small amount of the material on anatomy was published in Leonardo's Treatise on painting.[138] During the time that Melzi was ordering the material into chapters for publication, they were examined by a number of anatomists and artists, including Vasari, Cellini and Albrecht Dürer, who made a number of drawings from them.[138]
+Leonardo's anatomical drawings include many studies of the human skeleton and its parts, and of muscles and sinews. He studied the mechanical functions of the skeleton and the muscular forces that are applied to it in a manner that prefigured the modern science of biomechanics.[157] He drew the heart and vascular system, the sex organs and other internal organs, making one of the first scientific drawings of a fetus in utero.[133] The drawings and notation are far ahead of their time, and if published would undoubtedly have made a major contribution to medical science.[156]
+Leonardo also closely observed and recorded the effects of age and of human emotion on the physiology, studying in particular the effects of rage. He drew many figures who had significant facial deformities or signs of illness.[38][133] Leonardo also studied and drew the anatomy of many animals, dissecting cows, birds, monkeys, bears, and frogs, and comparing in his drawings their anatomical structure with that of humans. He also made a number of studies of horses.[133]
+Leonardo's dissections and documentation of muscles, nerves, and vessels helped to describe the physiology and mechanics of movement. He attempted to identify the source of 'emotions' and their expression. He found it difficult to incorporate the prevailing system and theories of bodily humours, but eventually he abandoned these physiological explanations of bodily functions. He made the observations that humours were not located in cerebral spaces or ventricles. He documented that the humours were not contained in the heart or the liver, and that it was the heart that defined the circulatory system. He was the first to define atherosclerosis and liver cirrhosis. He created models of the cerebral ventricles with the use of melted wax and constructed a glass aorta to observe the circulation of blood through the aortic valve by using water and grass seed to watch flow patterns. Vesalius published his work on anatomy and physiology in De humani corporis fabrica in 1543.[158]
+During his lifetime, Leonardo was also valued as an engineer. With the same rational and analytical approach that moved him to represent the human body and to investigate anatomy, Leonardo studied and designed a bewildering number of machines and devices. He drew their “anatomy” with unparalleled mastery, producing the first form of the modern technical drawing, including a perfected "exploded view" technique, to represent internal components. Those studies and projects collected in his codices fill more than 5,000 pages.[159] In a letter of 1482 to the lord of Milan Ludovico il Moro, he wrote that he could create all sorts of machines both for the protection of a city and for siege. When he fled from Milan to Venice in 1499, he found employment as an engineer and devised a system of moveable barricades to protect the city from attack. In 1502, he created a scheme for diverting the flow of the Arno river, a project on which Niccolò Machiavelli also worked.[160][161] He continued to contemplate the canalization of Lombardy's plains while in Louis XII's company[63] and of the Loire and its tributaries in the company of Francis I.[162] Leonardo's journals include a vast number of inventions, both practical and impractical. They include musical instruments, a mechanical knight, hydraulic pumps, reversible crank mechanisms, finned mortar shells, and a steam cannon.[27][38]
+Leonardo was fascinated by the phenomenon of flight for much of his life, producing many studies, including Codex on the Flight of Birds (c. 1505), as well as plans for several flying machines, such as a flapping ornithopter and a machine with a helical rotor.[38] A 2003 documentary by British television station Channel Four, titled Leonardo's Dream Machines, various designs by Leonardo, such as a parachute and a giant crossbow, were interpreted and constructed.[163][164] Some of those designs proved successful, whilst others fared less well when tested.
+Research performed by Marc van den Broek revealed older prototypes for more than 100 inventions that are ascribed to Leonardo. Similarities between Leonardo´s illustrations and drawings from the Middle Ages and from Ancient Greece and Rome, the Chinese and Persian Empires, and Egypt suggest that a large portion of Leonardo's inventions had been conceived before his lifetime. Leonardo´s innovation was to combine different functions from existing drafts and set them into scenes that illustrated their utility. By reconstituting technical inventions he created something new.[165]
+Leonardo's fame within his own lifetime was such that the King of France carried him away like a trophy, and was claimed to have supported him in his old age and held him in his arms as he died. Interest in Leonardo and his work has never diminished. Crowds still queue to see his best-known artworks, T-shirts still bear his most famous drawing, and writers continue to hail him as a genius while speculating about his private life, as well as about what one so intelligent actually believed in.[38]
+The continued admiration that Leonardo commanded from painters, critics and historians is reflected in many other written tributes. Baldassare Castiglione, author of Il Cortegiano (The Courtier), wrote in 1528: "...Another of the greatest painters in this world looks down on this art in which he is unequalled..."[166] while the biographer known as "Anonimo Gaddiano" wrote, c. 1540: "His genius was so rare and universal that it can be said that nature worked a miracle on his behalf..."[167]
+Giorgio Vasari, in the enlarged edition of Lives of the Artists (1568)[168] introduced his chapter on Leonardo with the following words:
+In the normal course of events many men and women are born with remarkable talents; but occasionally, in a way that transcends nature, a single person is marvellously endowed by Heaven with beauty, grace and talent in such abundance that he leaves other men far behind, all his actions seem inspired and indeed everything he does clearly comes from God rather than from human skill. Everyone acknowledged that this was true of Leonardo da Vinci, an artist of outstanding physical beauty, who displayed infinite grace in everything that he did and who cultivated his genius so brilliantly that all problems he studied he solved with ease.
+The 19th century brought a particular admiration for Leonardo's genius, causing Henry Fuseli to write in 1801: "Such was the dawn of modern art, when Leonardo da Vinci broke forth with a splendour that distanced former excellence: made up of all the elements that constitute the essence of genius..."[169] This is echoed by A.E. Rio who wrote in 1861: "He towered above all other artists through the strength and the nobility of his talents."[170]
+By the 19th century, the scope of Leonardo's notebooks was known, as well as his paintings. Hippolyte Taine wrote in 1866: "There may not be in the world an example of another genius so universal, so incapable of fulfilment, so full of yearning for the infinite, so naturally refined, so far ahead of his own century and the following centuries."[171]
+Art historian Bernard Berenson wrote in 1896: "Leonardo is the one artist of whom it may be said with perfect literalness: Nothing that he touched but turned into a thing of eternal beauty. Whether it be the cross section of a skull, the structure of a weed, or a study of muscles, he, with his feeling for line and for light and shade, forever transmuted it into life-communicating values."[172]
+The interest in Leonardo's genius has continued unabated; experts study and translate his writings, analyse his paintings using scientific techniques, argue over attributions and search for works which have been recorded but never found.[173] Liana Bortolon, writing in 1967, said: "Because of the multiplicity of interests that spurred him to pursue every field of knowledge...Leonardo can be considered, quite rightly, to have been the universal genius par excellence, and with all the disquieting overtones inherent in that term. Man is as uncomfortable today, faced with a genius, as he was in the 16th century. Five centuries have passed, yet we still view Leonardo with awe."[27]
+Twenty-first-century author Walter Isaacson based much of his biography of Leonardo[104] on thousands of notebook entries, studying the personal notes, sketches, budget notations, and musings of the man whom he considers the greatest of innovators. Isaacson was surprised to discover a "fun, joyous" side of Leonardo in addition to his limitless curiosity and creative genius.[174]
+On the 500th anniversary of Leonardo's death, the Louvre in Paris arranged for the largest ever single exhibit of his work, called Leonardo, between November 2019 and February 2020. The exhibit includes over 100 paintings, drawings and notebooks. Eleven of the paintings that Leonardo completed in his lifetime were included. Five of these are owned by the Louvre, but the Mona Lisa was not included because it is in such great demand among general visitors to the Louvre; it remains on display in its gallery. Vitruvian Man, however, is on display following a legal battle with its owner, the Gallerie dell'Accademia in Venice. Salvator Mundi was also not included because its Saudi owner did not agree to lease the work.[175][176]
+Much of the Collegiate Church of Saint Florentin at the Château d'Amboise, where Leonardo was buried, was damaged during the French Revolution, leading to the church's demolition in 1802.[177] Some of the graves were destroyed in the process, scattering the bones interred there and thereby leaving the whereabouts of Leonardo's remains subject to dispute. While excavating the site in 1863, fine-arts inspector general Arsène Houssaye found a partially complete skeleton with a bronze ring on one finger, some white hair, and stone fragments bearing the inscriptions "EO," "AR," "DUS," and "VINC"—interpreted as forming "Leonardus Vinci".[177][89][178][179] A silver shield found near the bones depicts a beardless Francis I, corresponding to the king's appearance during Leonardo's lifetime, and the skull's inclusion of eight teeth corresponds to someone of approximately the appropriate age.[178] The unusually large skull led Houssaye to believe that he had located Leonardo's remains,[179] but he thought the skeleton seemed too short.[178] Other art historians say that the 1.73 metres (5.7 ft) tall skeleton may well be Leonardo's.[180]
+The remains, except for the ring and a lock of hair which Houssaye kept,[89] were brought to Paris in a lead box, where the skull was allegedly presented to Napoleon III,[178] before being returned to the Château d'Amboise and re-interred in the Chapel of Saint Hubert in 1874.[177] A new memorial tombstone was added by sculptor Francesco La Monaca in the 1930s.[181] Reflecting doubts about the attribution, a plaque above the tomb states that the remains are only presumed to be those of Leonardo.[177] It has since been theorized that the folding of the skeleton's right arm over the head may correspond to the paralysis of Leonardo's right hand.[178][77][83]
+In 2016, it was announced that DNA tests would be conducted to determine whether the attribution is correct.[177] The DNA of the remains will be compared to that of samples collected from Leonardo's work and his half-brother Domenico's descendants;[177] it may also be sequenced.[182] The lock of hair and ring, now in a private US collection,[ac] were displayed in Vinci beginning on 2 May 2019, the 500th anniversary of the artist's death.[183][89]
+Salvator Mundi, a painting by Leonardo depicting Jesus holding an orb, sold for a world record US$450.3 million at a Christie's auction in New York, 15 November 2017.[184] The highest known sale price for any artwork was previously US$300 million, for Willem de Kooning's Interchange, which was sold privately in September 2015.[185] The highest price previously paid for a work of art at auction was for Pablo Picasso's Les Femmes d'Alger, which sold for US$179.4 million in May 2015 at Christie's New York.[185]
+Early sources
+Modern sources

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+Leonardo da Vinci (English: /ˌliːəˈnɑːrdoʊ də ˈvɪntʃi, ˌliːoʊˈ-, ˌleɪoʊˈ-/;[a] 14/15 April 1452[b] – 2 May 1519),[4] was an Italian polymath of the High Renaissance who is widely considered one of the greatest painters of all time (despite less than 25 of his paintings having survived).[c] The Mona Lisa is the most famous of Leonardo's works and the most famous portrait ever made.[5] The Last Supper is the most reproduced religious painting of all time[6] and his Vitruvian Man drawing is also regarded as a cultural icon.[7] He is also known for his his notebooks, in which he made drawings and notes on science and invention; these involve a variety of subjects including anatomy, cartography, painting, and paleontology. Leonardo's collective works compose a contribution to later generations of artists rivalled only by that of his contemporary Michelangelo.[8]
+Properly named Leonardo di ser Piero da Vinci,[d] Leonardo was born out of wedlock to a notary, Piero da Vinci, and a peasant woman, Caterina, in Vinci, in the region of Florence, Italy. Leonardo was educated in the studio of the renowned Italian painter Andrea del Verrocchio. Much of his earlier working life was spent in the service of Ludovico il Moro in Milan, and he later worked in Rome, Bologna and Venice. He spent his last three years in France, where he died in 1519.
+Although he had no formal academic training,[9] many historians and scholars regard Leonardo as the prime exemplar of the "Universal Genius" or "Renaissance Man", an individual of "unquenchable curiosity" and "feverishly inventive imagination."[6] He is widely considered one of the most diversely talented individuals ever to have lived.[10] According to art historian Helen Gardner, the scope and depth of his interests were without precedent in recorded history, and "his mind and personality seem to us superhuman, while the man himself mysterious and remote."[6] Scholars interpret his view of the world as being based in logic, though the empirical methods he used were unorthodox for his time.[11]
+Leonardo is revered for his technological ingenuity. He conceptualized flying machines, a type of armoured fighting vehicle, concentrated solar power, an adding machine,[12] and the double hull. Relatively few of his designs were constructed or even feasible during his lifetime, as the modern scientific approaches to metallurgy and engineering were only in their infancy during the Renaissance. Some of his smaller inventions, however, entered the world of manufacturing unheralded, such as an automated bobbin winder and a machine for testing the tensile strength of wire. He is also sometimes credited with the inventions of the parachute, helicopter, and tank.[13][14] He made substantial discoveries in anatomy, civil engineering, geology, optics, and hydrodynamics, but he did not publish his findings and they had little to no direct influence on subsequent science.[15]
+Leonardo was born on 14/15 April 1452[b] in the Tuscan hill town of Vinci, in the lower valley of the Arno river in the territory of the Medici-ruled Republic of Florence.[18] He was the out-of-wedlock son of Messer Piero Fruosino di Antonio da Vinci, a wealthy Florentine legal notary, and a peasant named Caterina,[e] identified as Caterina Buti del Vacca and more recently as Caterina di Meo Lippi by historian Martin Kemp. There have been many theories regarding Leonardo's mother's identity, including that she was a slave of foreign origin or an impoverished local youth.[17][20][21][22][f] Leonardo had no surname in the modern sense—da Vinci simply meaning "of Vinci"; his full birth name was Lionardo di ser Piero da Vinci,[2][25] meaning "Leonardo, (son) of ser Piero (from) Vinci."[18][d]
+Leonardo spent his first years in the hamlet of Anchiano in the home of his mother, and from at least 1457 lived in the household of his father, grandparents and uncle in the small town of Vinci.[26] His father had married a 16-year-old girl named Albiera Amadori, who loved Leonardo but died young[27] in 1465 without children. In 1468, when Leonardo was 16, his father married again to 20-year-old Francesca Lanfredini, who also died without children. Piero's legitimate heirs were born from his third wife Margherita di Guglielmo, who gave birth to six children, and his fourth and final wife, Lucrezia Cortigiani, who bore him another six heirs.[28][29]
+In all, Leonardo had 12 half-siblings, who were much younger than he was (the last was born when Leonardo was 40 years old) and with whom he had very little contact.[g]
+Leonardo received an informal education in Latin, geometry and mathematics. In later life, Leonardo recorded few distinct childhood incidents. One was of a kite coming to his cradle and opening his mouth with its tail; he regarded this as an omen of his writing on the subject.[31][32] The second occurred while he was exploring in the mountains: he discovered a cave and was both terrified that some great monster might lurk there and driven by curiosity to find out what was inside.[27] He also seems to have remembered some of his childhood observations of water, writing and crossing out the name of his hometown in one of his notebooks on the formation of rivers.[26]
+Leonardo's early life has been the subject of historical conjecture.[33] Vasari, the 16th-century biographer of Renaissance painters, tells a story of Leonardo as a very young man: A local peasant made himself a round shield and requested that Ser Piero have it painted for him. Leonardo, inspired by the story of Medusa, responded with a painting of a monster spitting fire that was so terrifying that his father bought a different shield to give to the peasant and sold Leonardo's to a Florentine art dealer for 100 ducats, who in turn sold it to the Duke of Milan.[34]
+In the mid-1460s, Leonardo's family moved to Florence, and around the age of 14,[30] he became a garzone (studio boy) in the workshop of Verrocchio, who was the leading Florentine painter and sculptor of his time.[35] Leonardo became an apprentice by the age of 17 and remained in training for seven years.[36][37] Other famous painters apprenticed in the workshop or associated with it include Ghirlandaio, Perugino, Botticelli, and Lorenzo di Credi.[27][38] Leonardo was exposed to both theoretical training and a wide range of technical skills,[39] including drafting, chemistry, metallurgy, metal working, plaster casting, leather working, mechanics, and wood-work, as well as the artistic skills of drawing, painting, sculpting, and modelling.[40][h]
+Much of the painting in Verrocchio's workshop was done by his employees. According to Vasari, Leonardo collaborated with Verrocchio on his The Baptism of Christ, painting the young angel holding Jesus' robe in a manner that was so far superior to his master's that Verrocchio put down his brush and never painted again, although this is believed to be an apocryphal story.[41] Close examination reveals areas of the work that have been painted or touched-up over the tempera, using the new technique of oil paint, including the landscape, the rocks seen through the brown mountain stream, and much of the figure of Jesus, bearing witness to the hand of Leonardo.[42] Leonardo may have been the model for two works by Verrocchio: the bronze statue of David in the Bargello, and the Archangel Raphael in Tobias and the Angel.[20]
+By 1472, at the age of 20, Leonardo qualified as a master in the Guild of Saint Luke, the guild of artists and doctors of medicine,[i] but even after his father set him up in his own workshop, his attachment to Verrocchio was such that he continued to collaborate and live with him.[27][43] Leonardo's earliest known dated work is a 1473 pen-and-ink drawing of the Arno valley,[38] which has been cited as the first "pure" landscape in the Occident.[j][44] According to Vasari, the young Leonardo was the first to suggest making the Arno river a navigable channel between Florence and Pisa.[45]
+In January 1478, Leonardo received an independent commission to paint an altarpiece for the Chapel of St. Bernard in the Palazzo Vecchio,[46] an indication of his independence from Verrocchio's studio. One anonymous writer claims that in 1480, Leonardo was living with the Medici and often worked in the garden of the Piazza San Marco, Florence, where a Neoplatonic academy of artists, poets and philosophers organized by the Medici met.[20] In March 1481, he received a commission from the monks of San Donato in Scopeto for The Adoration of the Magi.[47]
+Neither of these initial commissions were completed, being abandoned when Leonardo went to offer his services to Duke of Milan Ludovico Sforza. In 1482, he casted a silver stringed instrument[k] from a horse's skull and ram horns to bring to Sforza,[48] whom he wrote a letter describing the diverse things that he could achieve in the fields of engineering and weapon design, and mentioning that he could paint.[38][48][49]
+Leonardo worked in Milan from 1482 until 1499. He was commissioned to paint the Virgin of the Rocks for the Confraternity of the Immaculate Conception and The Last Supper for the monastery of Santa Maria delle Grazie.[51] In the spring of 1485, Leonardo travelled to Hungary on behalf of Sforza to meet king Matthias Corvinus, and was commissioned by him to paint a Madonna.[52]
+Leonardo was employed on many other projects for Sforza, including the preparation of floats and pageants for special occasions, a drawing and wooden model for a competition to design the cupola for Milan Cathedral (which he withdrew),[53] and a model for a huge equestrian monument to Ludovico's predecessor Francesco Sforza. This would have surpassed in size the only two large equestrian statues of the Renaissance, Donatello's Gattamelata in Padua and Verrocchio's Bartolomeo Colleoni in Venice, and became known as the Gran Cavallo.[38] Leonardo completed a model for the horse and made detailed plans for its casting,[38] but in November 1494, Ludovico gave the bronze to his brother-in-law to be used for a cannon to defend the city from Charles VIII.[38]
+With Ludovico Sforza overthrown at the dawn of the Second Italian War, Leonardo, with his assistant Salaì and friend, the mathematician Luca Pacioli, fled Milan for Venice.[54] There, he was employed as a military architect and engineer, devising methods to defend the city from naval attack.[27] On his return to Florence in 1500, he and his household were guests of the Servite monks at the monastery of Santissima Annunziata and were provided with a workshop where, according to Vasari, Leonardo created the cartoon of The Virgin and Child with St Anne and St John the Baptist, a work that won such admiration that "men and women, young and old" flocked to see it "as if they were attending a great festival."[55][l]
+In Cesena in 1502, Leonardo entered the service of Cesare Borgia, the son of Pope Alexander VI, acting as a military architect and engineer and travelling throughout Italy with his patron.[54] Leonardo created a map of Cesare Borgia's stronghold, a town plan of Imola in order to win his patronage. Maps were extremely rare at the time and it would have seemed like a new concept. Upon seeing it, Cesare hired Leonardo as his chief military engineer and architect. Later in the year, Leonardo produced another map for his patron, one of Chiana Valley, Tuscany, so as to give his patron a better overlay of the land and greater strategic position. He created this map in conjunction with his other project of constructing a dam from the sea to Florence, in order to allow a supply of water to sustain the canal during all seasons.
+Leonardo had left Borgia's service and returned to Florence by early 1503,[57] where he rejoined the Guild of Saint Luke on 18 October of that year. By this same month, Leonardo had begun working on a portrait of Lisa del Giocondo, the model for the Mona Lisa,[58][59] which he would continue working on until his twilight years. In January 1504, he was part of a committee formed to recommend where Michelangelo's statue of David should be placed.[60] He then spent two years in Florence designing and painting a mural of The Battle of Anghiari for the Signoria,[54] with Michelangelo designing its companion piece, The Battle of Cascina.[m]
+In 1506, Leonardo was summoned to Milan by Charles II d'Amboise, the acting French governor of the city. The Council of Florence wished Leonardo to return promptly to finish The Battle of Anghiari, but he was given leave at the behest of Louis XII, who considered commissioning the artist to make some portraits.[63] Leonardo may have commenced a project for an equestrian figure of d'Amboise;[64] a wax model survives and, if genuine, is the only extant example of Leonardo's sculpture. Leonardo was otherwise free to pursue his scientific interests.[63] Many of Leonardo's most prominent pupils either knew or worked with him in Milan,[27] including Bernardino Luini, Giovanni Antonio Boltraffio, and Marco d'Oggiono.
+In 1507, Leonardo was in Florence sorting out a dispute with his brothers over the estate of his father, who had died in 1504. By 1508, Leonardo was back in Milan, living in his own house in Porta Orientale in the parish of Santa Babila.[65]
+In 1512, Leonardo was working on plans for an equestrian monument for Gian Giacomo Trivulzio, but this was prevented by an invasion of a confederation of Swiss, Spanish and Venetian forces, which drove the French from Milan. Leonardo stayed in the city, spending several months in 1513 at the Medici's Vaprio d'Adda villa.[67] In March of that year, Lorenzo de' Medici's son Giovanni assumed the papacy (as Leo X); Leonardo went to Rome that September, where he was received by the pope's brother Giuliano.[67] From September 1513 to 1516, Leonardo spent much of his time living in the Belvedere Courtyard in the Apostolic Palace, where Michelangelo and Raphael were both active.[65] Leonardo was given an allowance of 33 ducats a month, and according to Vasari, decorated a lizard with scales dipped in quicksilver.[68] The pope gave him a painting commission of unknown subject matter, but cancelled it when the artist set about developing a new kind of varnish.[68][n] Leonardo became ill, in what may have been the first of multiple strokes leading to his death.[68] He practiced botany in the Gardens of Vatican City, and was commissioned to make plans for the pope's proposed draining of the Pontine Marshes.[69] He also dissected cadavers, making notes for a treatise on vocal cords;[70] these he gave to an official in hopes of regaining the pope's favor, but was unsuccessful.[68]
+In October 1515, King Francis I of France recaptured Milan.[47] Leonardo was present at the 19 December meeting of Francis I and Leo X, which took place in Bologna.[27][71][72] In 1516, Leonardo entered Francis' service, being given the use of the manor house Clos Lucé, near the king's residence at the royal Château d'Amboise. Being frequently visited by Francis, he drew plans for an immense castle town the king intended to erect at Romorantin, and made a mechanical lion, which during a pageant walked toward the king and—upon being struck by a wand—opened its chest to reveal a cluster of lilies.[73][74][o] Leonardo was accompanied during this time by his friend and apprentice, Francesco Melzi, and supported by a pension totalling 10,000 scudi.[65] At some point, Melzi drew a portrait of Leonardo; the only others known from his lifetime were a sketch by an unknown assistant on the back of one of Leonardo's studies (c. 1517)[76] and a drawing by Giovanni Ambrogio Figino depicting an elderly Leonardo with his right arm assuaged by cloth.[77][p][q] The latter, in addition to the record of an October 1517 visit by Louis d'Aragon,[r] confirms an account of Leonardo's right hand being paralytic at the age of 65,[80] which may indicate why he left works such as the Mona Lisa unfinished.[78][81][82] He continued to work at some capacity until eventually becoming ill and bedridden for several months.[80]
+Leonardo died at Clos Lucé on 2 May 1519 at the age of 67, possibly of a stroke.[83][82] Francis I had become a close friend. Vasari describes Leonardo as lamenting on his deathbed, full of repentance, that "he had offended against God and men by failing to practice his art as he should have done."[84] Vasari states that in his last days, Leonardo sent for a priest to make his confession and to receive the Holy Sacrament.[85] Vasari also records that the king held Leonardo's head in his arms as he died, although this story may be legend rather than fact.[s][t] In accordance with his will, sixty beggars carrying tapers followed Leonardo's casket.[87][u] Melzi was the principal heir and executor, receiving, as well as money, Leonardo's paintings, tools, library and personal effects. Leonardo also remembered his other long-time pupil and companion, Salaì, and his servant Baptista de Vilanis, who each received half of Leonardo's vineyards.[88] His brothers received land, and his serving woman received a fur-lined cloak. On 12 August 1519, Leonardo's remains were interred in the Collegiate Church of Saint Florentin at the Château d'Amboise.[89]
+Some 20 years after Leonardo's death, Francis was reported by the goldsmith and sculptor Benvenuto Cellini as saying: "There had never been another man born in the world who knew as much as Leonardo, not so much about painting, sculpture and architecture, as that he was a very great philosopher."[90]
+Florence at the time of Leonardo's youth was the centre of Christian Humanist thought and culture.[35] Leonardo commenced his apprenticeship with Verrocchio in 1466, the year that Verrocchio's master, the great sculptor Donatello, died.[v] The painter Uccello, whose early experiments with perspective were to influence the development of landscape painting, was a very old man. The painters Piero della Francesca and Filippo Lippi, sculptor Luca della Robbia, and architect and writer Leon Battista Alberti were in their sixties. The successful artists of the next generation were Leonardo's teacher Verrocchio, Antonio del Pollaiuolo, and the portrait sculptor Mino da Fiesole.[91][92][93][94]
+Leonardo's youth was spent in a Florence that was ornamented by the works of these artists and by Donatello's contemporaries, Masaccio, whose figurative frescoes were imbued with realism and emotion; and Ghiberti, whose Gates of Paradise, gleaming with gold leaf, displayed the art of combining complex figure compositions with detailed architectural backgrounds. Piero della Francesca had made a detailed study of perspective,[95] and was the first painter to make a scientific study of light. These studies and Alberti's treatise De pictura[96] were to have a profound effect on younger artists and in particular on Leonardo's own observations and artworks.[91][93][94]
+A prevalent tradition in Florence was the small altarpiece of the Virgin and Child. Many of these were created in tempera or glazed terracotta by the workshops of Filippo Lippi, Verrocchio and the prolific della Robbia family.[91] Leonardo's early Madonnas such as The Madonna with a carnation and the Benois Madonna followed this tradition while showing idiosyncratic departures, particularly in the latter in which the Virgin is set at an oblique angle to the picture space with the Christ Child at the opposite angle. This compositional theme was to emerge in Leonardo's later paintings such as The Virgin and Child with St. Anne.[27]
+Leonardo was a contemporary of Botticelli, Domenico Ghirlandaio and Perugino, who were all slightly older than he was.[92] He would have met them at the workshop of Verrocchio, with whom they had associations, and at the Academy of the Medici.[27] Botticelli was a particular favourite of the Medici family, and thus his success as a painter was assured. Ghirlandaio and Perugino were both prolific and ran large workshops. They competently delivered commissions to well-satisfied patrons who appreciated Ghirlandaio's ability to portray the wealthy citizens of Florence within large religious frescoes, and Perugino's ability to deliver a multitude of saints and angels of unfailing sweetness and innocence.[91]
+These three were among those commissioned to paint the walls of the Sistine Chapel, the work commencing with Perugino's employment in 1479. Leonardo was not part of this prestigious commission. His first significant commission, The Adoration of the Magi for the Monks of Scopeto, was never completed.[27]
+In 1476, during the time of Leonardo's association with Verrocchio's workshop, the Portinari Altarpiece by Hugo van der Goes arrived in Florence, bringing from Northern Europe new painterly techniques that were to profoundly affect Leonardo, Ghirlandaio, Perugino and others.[92] In 1479, the Sicilian painter Antonello da Messina, who worked exclusively in oils, travelled north on his way to Venice, where the leading painter Giovanni Bellini adopted the technique of oil painting, quickly making it the preferred method in Venice. Leonardo was also later to visit Venice.[92][94]
+Like the two contemporary architects Donato Bramante (who designed the Belvedere Courtyard) and Antonio da Sangallo the Elder, Leonardo experimented with designs for centrally planned churches, a number of which appear in his journals, as both plans and views, although none was ever realised.[92][97]
+Leonardo's political contemporaries were Lorenzo de' Medici (il Magnifico), who was three years older, and his younger brother Giuliano, who was slain in the Pazzi conspiracy in 1478. Leonardo was sent as an ambassador by the Medici court to Ludovico il Moro, who ruled Milan between 1479 and 1499.[92]
+With Alberti, Leonardo visited the home of the Medici and through them came to know the older Humanist philosophers of whom Marsiglio Ficino, proponent of Neo Platonism; Cristoforo Landino, writer of commentaries on Classical writings, and John Argyropoulos, teacher of Greek and translator of Aristotle were the foremost. Also associated with the Academy of the Medici was Leonardo's contemporary, the brilliant young poet and philosopher Pico della Mirandola.[92][94][87] Leonardo later wrote in the margin of a journal, "The Medici made me and the Medici destroyed me." While it was through the action of Lorenzo that Leonardo received his employment at the court of Milan, it is not known exactly what Leonardo meant by this cryptic comment.[27]
+Although usually named together as the three giants of the High Renaissance, Leonardo, Michelangelo and Raphael were not of the same generation. Leonardo was 23 when Michelangelo was born and 31 when Raphael was born.[92] Raphael died at the age of 37 in 1520, the year after Leonardo died, but Michelangelo went on creating for another 45 years.[93][94]
+Within Leonardo's lifetime, his extraordinary powers of invention, his "outstanding physical beauty," "infinite grace," "great strength and generosity," "regal spirit and tremendous breadth of mind," as described by Vasari,[98] as well as all other aspects of his life, attracted the curiosity of others. One such aspect was his love for animals, likely including vegetarianism and according to Vasari, a habit of purchasing caged birds and releasing them.[99][100]
+Leonardo had many friends who are now renowned either in their fields or for their historical significance. They included the mathematician Luca Pacioli,[101] with whom he collaborated on the book Divina proportione in the 1490s. Leonardo appears to have had no close relationships with women except for his friendship with Cecilia Gallerani and the two Este sisters, Beatrice and Isabella.[102] While on a journey that took him through Mantua, he drew a portrait of Isabella that appears to have been used to create a painted portrait, now lost.[27]
+Beyond friendship, Leonardo kept his private life secret. His sexuality has been the subject of satire, analysis, and speculation. This trend began in the mid-16th century and was revived in the 19th and 20th centuries, most notably by Sigmund Freud.[103] Leonardo's most intimate relationships were perhaps with his pupils Salaì and Melzi. Melzi, writing to inform Leonardo's brothers of his death, described Leonardo's feelings for his pupils as both loving and passionate. It has been claimed since the 16th century that these relationships were of a sexual or erotic nature. Court records of 1476, when he was aged twenty-four, show that Leonardo and three other young men were charged with sodomy in an incident involving a well-known male prostitute. The charges were dismissed for lack of evidence, and there is speculation that since one of the accused, Lionardo de Tornabuoni, was related to Lorenzo de' Medici, the family exerted its influence to secure the dismissal.[104] Since that date much has been written about his presumed homosexuality and its role in his art, particularly in the androgyny and eroticism manifested in Saint John the Baptist and Bacchus and more explicitly in a number of erotic drawings.[105]
+Gian Giacomo Caprotti da Oreno, nicknamed Salaì or Il Salaino ("The Little Unclean One," i.e., the devil), entered Leonardo's household in 1490. After only a year, Leonardo made a list of his misdemeanours, calling him "a thief, a liar, stubborn, and a glutton," after he had made off with money and valuables on at least five occasions and spent a fortune on clothes.[106] Nevertheless, Leonardo treated him with great indulgence, and he remained in Leonardo's household for the next thirty years.[107] Salaì executed a number of paintings under the name of Andrea Salaì, but although Vasari claims that Leonardo "taught him a great deal about painting,"[74] his work is generally considered to be of less artistic merit than others among Leonardo's pupils, such as Marco d'Oggiono and Boltraffio. In 1515, he painted a nude version of the Mona Lisa, known as Monna Vanna.[108] Salaì owned the Mona Lisa at the time of his death in 1524, and in his will it was assessed at 505 lire, an exceptionally high valuation for a small panel portrait.[109]
+In 1506, Leonardo took on another pupil, Count Francesco Melzi, the son of a Lombard aristocrat, who is considered to have been his favourite student. He travelled to France with Leonardo and remained with him until Leonardo's death.[27] Melzi inherited the artistic and scientific works, manuscripts, and collections of Leonardo and administered the estate.
+Despite the recent awareness and admiration of Leonardo as a scientist and inventor, for the better part of four hundred years his fame rested on his achievements as a painter. A handful of works that are either authenticated or attributed to him have been regarded as among the great masterpieces. These paintings are famous for a variety of qualities that have been much imitated by students and discussed at great length by connoisseurs and critics. By the 1490s Leonardo had already been described as a "Divine" painter.[110]
+Among the qualities that make Leonardo's work unique are his innovative techniques for laying on the paint; his detailed knowledge of anatomy, light, botany and geology; his interest in physiognomy and the way humans register emotion in expression and gesture; his innovative use of the human form in figurative composition; and his use of subtle gradation of tone. All these qualities come together in his most famous painted works, the Mona Lisa, the Last Supper, and the Virgin of the Rocks.[111]
+Leonardo first gained attention for his work on the Baptism of Christ, painted in conjunction with Verrocchio. Two other paintings appear to date from his time at Verrocchio's workshop, both of which are Annunciations. One is small, 59 centimetres (23 in) long and 14 centimetres (5.5 in) high. It is a "predella" to go at the base of a larger composition, a painting by Lorenzo di Credi from which it has become separated. The other is a much larger work, 217 centimetres (85 in) long.[112] In both Annunciations, Leonardo used a formal arrangement, like two well-known pictures by Fra Angelico of the same subject, of the Virgin Mary sitting or kneeling to the right of the picture, approached from the left by an angel in profile, with a rich flowing garment, raised wings and bearing a lily. Although previously attributed to Ghirlandaio, the larger work is now generally attributed to Leonardo.[113]
+In the smaller painting, Mary averts her eyes and folds her hands in a gesture that symbolised submission to God's will. Mary is not submissive, however, in the larger piece. The girl, interrupted in her reading by this unexpected messenger, puts a finger in her bible to mark the place and raises her hand in a formal gesture of greeting or surprise.[91] This calm young woman appears to accept her role as the Mother of God, not with resignation but with confidence. In this painting, the young Leonardo presents the humanist face of the Virgin Mary, recognising humanity's role in God's incarnation.
+In the 1480s, Leonardo received two very important commissions and commenced another work that was of ground-breaking importance in terms of composition. Two of the three were never finished, and the third took so long that it was subject to lengthy negotiations over completion and payment.
+One of these paintings was Saint Jerome in the Wilderness, which Bortolon associates with a difficult period of Leonardo's life, as evidenced in his diary: "I thought I was learning to live; I was only learning to die."[27] Although the painting is barely begun, the composition can be seen and is very unusual.[w] Jerome, as a penitent, occupies the middle of the picture, set on a slight diagonal and viewed somewhat from above. His kneeling form takes on a trapezoid shape, with one arm stretched to the outer edge of the painting and his gaze looking in the opposite direction. J. Wasserman points out the link between this painting and Leonardo's anatomical studies.[114] Across the foreground sprawls his symbol, a great lion whose body and tail make a double spiral across the base of the picture space. The other remarkable feature is the sketchy landscape of craggy rocks against which the figure is silhouetted.
+The daring display of figure composition, the landscape elements and personal drama also appear in the great unfinished masterpiece, the Adoration of the Magi, a commission from the Monks of San Donato a Scopeto. It is a complex composition, of about 250 x 250 centimetres. Leonardo did numerous drawings and preparatory studies, including a detailed one in linear perspective of the ruined classical architecture that forms part of the background. In 1482 Leonardo went to Milan at the behest of Lorenzo de' Medici in order to win favour with Ludovico il Moro, and the painting was abandoned.[20][113]
+The third important work of this period is the Virgin of the Rocks, commissioned in Milan for the Confraternity of the Immaculate Conception. The painting, to be done with the assistance of the de Predis brothers, was to fill a large complex altarpiece.[115] Leonardo chose to paint an apocryphal moment of the infancy of Christ when the infant John the Baptist, in protection of an angel, met the Holy Family on the road to Egypt. The painting demonstrates an eerie beauty as the graceful figures kneel in adoration around the infant Christ in a wild landscape of tumbling rock and whirling water.[116] While the painting is quite large, about 200×120 centimetres, it is not nearly as complex as the painting ordered by the monks of St Donato, having only four figures rather than about fifty and a rocky landscape rather than architectural details. The painting was eventually finished; in fact, two versions of the painting were finished: one remained at the chapel of the Confraternity, while Leonardo took the other to France. The Brothers did not get their painting, however, nor the de Predis their payment, until the next century.[38][54]
+Leonardo's most remarkable portrait of this period is the Lady with an Ermine, presumed to be  Cecilia Gallerani (c. 1483–1490), lover of Ludovico Sforza.[117][118] The painting is charcterised by the pose of the figure with the head turned at a very different angle to the torso, unusual at a date when many portraits were still rigidly in profile. The ermine plainly carries symbolic meaning, relating either to the sitter, or to Ludovico who belonged to the prestigious Order of the Ermine.[117]
+Leonardo's most famous painting of the 1490s is The Last Supper, commissioned for the refectory of the Convent of Santa Maria della Grazie in Milan. It represents the last meal shared by Jesus with his disciples before his capture and death, and shows the moment when Jesus has just said "one of you will betray me", and the consternation that this statement caused.[38]
+The novelist Matteo Bandello observed Leonardo at work and wrote that some days he would paint from dawn till dusk without stopping to eat and then not paint for three or four days at a time.[119] This was beyond the comprehension of the prior of the convent, who hounded him until Leonardo asked Ludovico to intervene. Vasari describes how Leonardo, troubled over his ability to adequately depict the faces of Christ and the traitor Judas, told the Duke that he might be obliged to use the prior as his model.[120]
+When finished, the painting was acclaimed as a masterpiece of design and characterization,[121] but it deteriorated rapidly, so that within a hundred years it was described by one viewer as "completely ruined."[122] Leonardo, instead of using the reliable technique of fresco, had used tempera over a ground that was mainly gesso, resulting in a surface subject to mould and to flaking.[123] Despite this, the painting remains one of the most reproduced works of art; countless copies have been made in various mediums.
+It is recorded that in 1492, Leonardo, with assistants painted the Sala delle Asse in the Sforza Castle in Milan,  with a trompe-l'œil depicting trees, with an intricate labyrinth of leaves and knots on the ceiling.[124]
+In 1505 Leonardo was commissioned to paint The Battle of Anghiari in the Salone dei Cinquecento (Hall of the Five Hundred) in the Palazzo Vecchio, Florence. Leonardo devised a dynamic composition depicting four men riding raging war horses engaged in a battle for possession of a standard, at the Battle of Anghiari in 1440.  Michelangelo was assigned the opposite wall to depict the Battle of Cascina. Leonardo's painting deteriorated rapidly and is now known from a copy by Rubens.[125]
+Among the works created by Leonardo in the 16th century is the small portrait known as the Mona Lisa or La Gioconda, the laughing one. In the present era, it is arguably the most famous painting in the world. Its fame rests, in particular, on the elusive smile on the woman's face, its mysterious quality perhaps due to the subtly shadowed corners of the mouth and eyes such that the exact nature of the smile cannot be determined. The shadowy quality for which the work is renowned came to be called "sfumato," or Leonardo's smoke. Vasari, who is generally thought to have known the painting only by repute, said that "the smile was so pleasing that it seemed divine rather than human; and those who saw it were amazed to find that it was as alive as the original."[126][x]
+Other characteristics of the painting are the unadorned dress, in which the eyes and hands have no competition from other details; the dramatic landscape background, in which the world seems to be in a state of flux; the subdued colouring; and the extremely smooth nature of the painterly technique, employing oils laid on much like tempera, and blended on the surface so that the brushstrokes are indistinguishable.[y] Vasari expressed the opinion that the manner of painting would make even "the most confident master...despair and lose heart."[129] The perfect state of preservation and the fact that there is no sign of repair or overpainting is rare in a panel painting of this date.[130]
+In the painting Virgin and Child with St. Anne, the composition again picks up the theme of figures in a landscape, which Wasserman describes as "breathtakingly beautiful"[131] and harkens back to the St Jerome picture with the figure set at an oblique angle. What makes this painting unusual is that there are two obliquely set figures superimposed. Mary is seated on the knee of her mother, St Anne. She leans forward to restrain the Christ Child as he plays roughly with a lamb, the sign of his own impending sacrifice.[38] This painting, which was copied many times, influenced Michelangelo, Raphael, and Andrea del Sarto,[132] and through them Pontormo and Correggio. The trends in composition were adopted in particular by the Venetian painters Tintoretto and Veronese.
+Leonardo was not a prolific painter, but he was a most prolific draftsman, keeping journals full of small sketches and detailed drawings recording all manner of things that took his attention. As well as the journals there exist many studies for paintings, some of which can be identified as preparatory to particular works such as The Adoration of the Magi, The Virgin of the Rocks and The Last Supper.[133]
+His earliest dated drawing is a Landscape of the Arno Valley, 1473, which shows the river, the mountains, Montelupo Castle and the farmlands beyond it in great detail.[27][133][z] According to art historian Ludwig Heydenreich, this is "The first true landscape in art."[134] Massimo Polidoro says that it was the first landscape "not to be the background of some religious scene or a portrait. It is the first [documented] time where a landscape was drawn just for the sake of it."[44]
+Among his famous drawings are the Vitruvian Man, a study of the proportions of the human body; the Head of an Angel, for The Virgin of the Rocks in the Louvre; a botanical study of Star of Bethlehem; and a large drawing (160×100 cm) in black chalk on coloured paper of The Virgin and Child with St. Anne and St. John the Baptist in the National Gallery, London.[133] This drawing employs the subtle sfumato technique of shading, in the manner of the Mona Lisa. It is thought that Leonardo never made a painting from it, the closest similarity being to The Virgin and Child with St. Anne in the Louvre.[135]
+Other drawings of interest include numerous studies generally referred to as "caricatures" because, although exaggerated, they appear to be based upon observation of live models. Vasari relates that if Leonardo saw a person with an interesting face he would follow them around all day observing them.[136] There are numerous studies of beautiful young men, often associated with Salaì, with the rare and much admired facial feature, the so-called "Grecian profile."[aa] These faces are often contrasted with that of a warrior.[133] Salaì is often depicted in fancy-dress costume. Leonardo is known to have designed sets for pageants with which these may be associated. Other, often meticulous, drawings show studies of drapery. A marked development in Leonardo's ability to draw drapery occurred in his early works. Another often-reproduced drawing is a macabre sketch that was done by Leonardo in Florence in 1479 showing the body of Bernardo Baroncelli, hanged in connection with the murder of Giuliano, brother of Lorenzo de' Medici, in the Pazzi conspiracy.[133] In his notes, Leonardo recorded the colours of the robes that Baroncelli was wearing when he died.
+Renaissance humanism recognised no mutually exclusive polarities between the sciences and the arts, and Leonardo's studies in science and engineering are sometimes considered as impressive and innovative as his artistic work.[38] These studies were recorded in 13,000 pages of notes and drawings, which fuse art and natural philosophy (the forerunner of modern science). They were made and maintained daily throughout Leonardo's life and travels, as he made continual observations of the world around him.[38]
+Leonardo's notes and drawings display an enormous range of interests and preoccupations, some as mundane as lists of groceries and people who owed him money and some as intriguing as designs for wings and shoes for walking on water. There are compositions for paintings, studies of details and drapery, studies of faces and emotions, of animals, babies, dissections, plant studies, rock formations, whirlpools, war machines, flying machines and architecture.[38]
+These notebooks—originally loose papers of different types and sizes, were largely entrusted to Leonardo's pupil and heir Francesco Melzi after the master's death.[137] These were to be published, a task of overwhelming difficulty because of its scope and Leonardo's idiosyncratic writing.[138] Some of Leonardo's drawings were copied by an anonymous Milanese artist for a planned treatise on art c. 1570.[139] After Melzi's death in 1570, the collection passed to his son, the lawyer Orazio, who initially took little interest in the journals.[137] In 1587, a Melzi household tutor named Lelio Gavardi took 13 of the manuscripts to Pisa; there, the architect Giovanni Magenta reproached Gavardi for having taken the manuscripts illicitly and returned them to Orazio. Having many more such works in his possession, Orazio gifted the volumes to Magenta. News spread of these lost works of Leonardo's, and Orazio retrieved seven of the 13 manuscripts, which he then gave to Pompeo Leoni for publication in two volumes; one of these was the Codex Atlanticus. The other six works had been distributed to a few others.[140] After Orazio's death, his heirs sold the rest of Leonardo's possessions, and thus began their dispersal.[141]
+Some works have found their way into major collections such as the Royal Library at Windsor Castle, the Louvre, the Biblioteca Nacional de España, the Victoria and Albert Museum, the Biblioteca Ambrosiana in Milan, which holds the 12-volume Codex Atlanticus, and the British Library in London, which has put a selection from the Codex Arundel (BL Arundel MS 263) online.[142] Works have also been at Holkham Hall, the Metropolitan Museum of Art, and in the private hands of John Nicholas Brown I and Robert Lehman.[137] The Codex Leicester is the only privately owned major scientific work of Leonardo; it is owned by Bill Gates and displayed once a year in different cities around the world.
+Most of Leonardo's writings are in mirror-image cursive.[143][44] Since Leonardo wrote with his left hand, it was probably easier for him to write from right to left.[144][ab] Leonardo used a variety of shorthand and symbols, and states in his notes that he intended to prepare them for publication.[143] In many cases a single topic is covered in detail in both words and pictures on a single sheet, together conveying information that would not be lost if the pages were published out of order.[147] Why they were not published during Leonardo's lifetime is unknown.[38]
+Leonardo's approach to science was observational: he tried to understand a phenomenon by describing and depicting it in utmost detail and did not emphasise experiments or theoretical explanation. Since he lacked formal education in Latin and mathematics, contemporary scholars mostly ignored Leonardo the scientist, although he did teach himself Latin. In the 1490s he studied mathematics under Luca Pacioli and prepared a series of drawings of regular solids in a skeletal form to be engraved as plates for Pacioli's book Divina proportione, published in 1509.[38] While living in Milan, he studied light from the summit of Monte Rosa.[63] Scientific writings in his notebook on fossils have been considered as influential on early palaeontology[148][149] and he has been called the father of ichnology.[150]
+The content of his journals suggest that he was planning a series of treatises on a variety of subjects. A coherent treatise on anatomy is said to have been observed during a visit by Cardinal Louis d'Aragon's secretary in 1517.[151] Aspects of his work on the studies of anatomy, light and the landscape were assembled for publication by Melzi and eventually published as A Treatise on Painting in France and Italy in 1651 and Germany in 1724,[152] with engravings based upon drawings by the Classical painter Nicolas Poussin.[153] According to Arasse, the treatise, which in France went into 62 editions in fifty years, caused Leonardo to be seen as "the precursor of French academic thought on art."[38]
+While Leonardo's experimentation followed scientific methods, a recent and exhaustive analysis of Leonardo as a scientist by Fritjof Capra argues that Leonardo was a fundamentally different kind of scientist from Galileo, Newton and other scientists who followed him in that, as a "Renaissance Man", his theorising and hypothesising integrated the arts and particularly painting.[154][page needed]
+Leonardo started his study in the anatomy of the human body under the apprenticeship of Andrea del Verrocchio, who demanded that his students develop a deep knowledge of the subject.[155] As an artist, he quickly became master of topographic anatomy, drawing many studies of muscles, tendons and other visible anatomical features.
+As a successful artist, Leonardo was given permission to dissect human corpses at the Hospital of Santa Maria Nuova in Florence and later at hospitals in Milan and Rome. From 1510 to 1511 he collaborated in his studies with the doctor Marcantonio della Torre. Leonardo made over 240 detailed drawings and wrote about 13,000 words towards a treatise on anatomy.[156] Only a small amount of the material on anatomy was published in Leonardo's Treatise on painting.[138] During the time that Melzi was ordering the material into chapters for publication, they were examined by a number of anatomists and artists, including Vasari, Cellini and Albrecht Dürer, who made a number of drawings from them.[138]
+Leonardo's anatomical drawings include many studies of the human skeleton and its parts, and of muscles and sinews. He studied the mechanical functions of the skeleton and the muscular forces that are applied to it in a manner that prefigured the modern science of biomechanics.[157] He drew the heart and vascular system, the sex organs and other internal organs, making one of the first scientific drawings of a fetus in utero.[133] The drawings and notation are far ahead of their time, and if published would undoubtedly have made a major contribution to medical science.[156]
+Leonardo also closely observed and recorded the effects of age and of human emotion on the physiology, studying in particular the effects of rage. He drew many figures who had significant facial deformities or signs of illness.[38][133] Leonardo also studied and drew the anatomy of many animals, dissecting cows, birds, monkeys, bears, and frogs, and comparing in his drawings their anatomical structure with that of humans. He also made a number of studies of horses.[133]
+Leonardo's dissections and documentation of muscles, nerves, and vessels helped to describe the physiology and mechanics of movement. He attempted to identify the source of 'emotions' and their expression. He found it difficult to incorporate the prevailing system and theories of bodily humours, but eventually he abandoned these physiological explanations of bodily functions. He made the observations that humours were not located in cerebral spaces or ventricles. He documented that the humours were not contained in the heart or the liver, and that it was the heart that defined the circulatory system. He was the first to define atherosclerosis and liver cirrhosis. He created models of the cerebral ventricles with the use of melted wax and constructed a glass aorta to observe the circulation of blood through the aortic valve by using water and grass seed to watch flow patterns. Vesalius published his work on anatomy and physiology in De humani corporis fabrica in 1543.[158]
+During his lifetime, Leonardo was also valued as an engineer. With the same rational and analytical approach that moved him to represent the human body and to investigate anatomy, Leonardo studied and designed a bewildering number of machines and devices. He drew their “anatomy” with unparalleled mastery, producing the first form of the modern technical drawing, including a perfected "exploded view" technique, to represent internal components. Those studies and projects collected in his codices fill more than 5,000 pages.[159] In a letter of 1482 to the lord of Milan Ludovico il Moro, he wrote that he could create all sorts of machines both for the protection of a city and for siege. When he fled from Milan to Venice in 1499, he found employment as an engineer and devised a system of moveable barricades to protect the city from attack. In 1502, he created a scheme for diverting the flow of the Arno river, a project on which Niccolò Machiavelli also worked.[160][161] He continued to contemplate the canalization of Lombardy's plains while in Louis XII's company[63] and of the Loire and its tributaries in the company of Francis I.[162] Leonardo's journals include a vast number of inventions, both practical and impractical. They include musical instruments, a mechanical knight, hydraulic pumps, reversible crank mechanisms, finned mortar shells, and a steam cannon.[27][38]
+Leonardo was fascinated by the phenomenon of flight for much of his life, producing many studies, including Codex on the Flight of Birds (c. 1505), as well as plans for several flying machines, such as a flapping ornithopter and a machine with a helical rotor.[38] A 2003 documentary by British television station Channel Four, titled Leonardo's Dream Machines, various designs by Leonardo, such as a parachute and a giant crossbow, were interpreted and constructed.[163][164] Some of those designs proved successful, whilst others fared less well when tested.
+Research performed by Marc van den Broek revealed older prototypes for more than 100 inventions that are ascribed to Leonardo. Similarities between Leonardo´s illustrations and drawings from the Middle Ages and from Ancient Greece and Rome, the Chinese and Persian Empires, and Egypt suggest that a large portion of Leonardo's inventions had been conceived before his lifetime. Leonardo´s innovation was to combine different functions from existing drafts and set them into scenes that illustrated their utility. By reconstituting technical inventions he created something new.[165]
+Leonardo's fame within his own lifetime was such that the King of France carried him away like a trophy, and was claimed to have supported him in his old age and held him in his arms as he died. Interest in Leonardo and his work has never diminished. Crowds still queue to see his best-known artworks, T-shirts still bear his most famous drawing, and writers continue to hail him as a genius while speculating about his private life, as well as about what one so intelligent actually believed in.[38]
+The continued admiration that Leonardo commanded from painters, critics and historians is reflected in many other written tributes. Baldassare Castiglione, author of Il Cortegiano (The Courtier), wrote in 1528: "...Another of the greatest painters in this world looks down on this art in which he is unequalled..."[166] while the biographer known as "Anonimo Gaddiano" wrote, c. 1540: "His genius was so rare and universal that it can be said that nature worked a miracle on his behalf..."[167]
+Giorgio Vasari, in the enlarged edition of Lives of the Artists (1568)[168] introduced his chapter on Leonardo with the following words:
+In the normal course of events many men and women are born with remarkable talents; but occasionally, in a way that transcends nature, a single person is marvellously endowed by Heaven with beauty, grace and talent in such abundance that he leaves other men far behind, all his actions seem inspired and indeed everything he does clearly comes from God rather than from human skill. Everyone acknowledged that this was true of Leonardo da Vinci, an artist of outstanding physical beauty, who displayed infinite grace in everything that he did and who cultivated his genius so brilliantly that all problems he studied he solved with ease.
+The 19th century brought a particular admiration for Leonardo's genius, causing Henry Fuseli to write in 1801: "Such was the dawn of modern art, when Leonardo da Vinci broke forth with a splendour that distanced former excellence: made up of all the elements that constitute the essence of genius..."[169] This is echoed by A.E. Rio who wrote in 1861: "He towered above all other artists through the strength and the nobility of his talents."[170]
+By the 19th century, the scope of Leonardo's notebooks was known, as well as his paintings. Hippolyte Taine wrote in 1866: "There may not be in the world an example of another genius so universal, so incapable of fulfilment, so full of yearning for the infinite, so naturally refined, so far ahead of his own century and the following centuries."[171]
+Art historian Bernard Berenson wrote in 1896: "Leonardo is the one artist of whom it may be said with perfect literalness: Nothing that he touched but turned into a thing of eternal beauty. Whether it be the cross section of a skull, the structure of a weed, or a study of muscles, he, with his feeling for line and for light and shade, forever transmuted it into life-communicating values."[172]
+The interest in Leonardo's genius has continued unabated; experts study and translate his writings, analyse his paintings using scientific techniques, argue over attributions and search for works which have been recorded but never found.[173] Liana Bortolon, writing in 1967, said: "Because of the multiplicity of interests that spurred him to pursue every field of knowledge...Leonardo can be considered, quite rightly, to have been the universal genius par excellence, and with all the disquieting overtones inherent in that term. Man is as uncomfortable today, faced with a genius, as he was in the 16th century. Five centuries have passed, yet we still view Leonardo with awe."[27]
+Twenty-first-century author Walter Isaacson based much of his biography of Leonardo[104] on thousands of notebook entries, studying the personal notes, sketches, budget notations, and musings of the man whom he considers the greatest of innovators. Isaacson was surprised to discover a "fun, joyous" side of Leonardo in addition to his limitless curiosity and creative genius.[174]
+On the 500th anniversary of Leonardo's death, the Louvre in Paris arranged for the largest ever single exhibit of his work, called Leonardo, between November 2019 and February 2020. The exhibit includes over 100 paintings, drawings and notebooks. Eleven of the paintings that Leonardo completed in his lifetime were included. Five of these are owned by the Louvre, but the Mona Lisa was not included because it is in such great demand among general visitors to the Louvre; it remains on display in its gallery. Vitruvian Man, however, is on display following a legal battle with its owner, the Gallerie dell'Accademia in Venice. Salvator Mundi was also not included because its Saudi owner did not agree to lease the work.[175][176]
+Much of the Collegiate Church of Saint Florentin at the Château d'Amboise, where Leonardo was buried, was damaged during the French Revolution, leading to the church's demolition in 1802.[177] Some of the graves were destroyed in the process, scattering the bones interred there and thereby leaving the whereabouts of Leonardo's remains subject to dispute. While excavating the site in 1863, fine-arts inspector general Arsène Houssaye found a partially complete skeleton with a bronze ring on one finger, some white hair, and stone fragments bearing the inscriptions "EO," "AR," "DUS," and "VINC"—interpreted as forming "Leonardus Vinci".[177][89][178][179] A silver shield found near the bones depicts a beardless Francis I, corresponding to the king's appearance during Leonardo's lifetime, and the skull's inclusion of eight teeth corresponds to someone of approximately the appropriate age.[178] The unusually large skull led Houssaye to believe that he had located Leonardo's remains,[179] but he thought the skeleton seemed too short.[178] Other art historians say that the 1.73 metres (5.7 ft) tall skeleton may well be Leonardo's.[180]
+The remains, except for the ring and a lock of hair which Houssaye kept,[89] were brought to Paris in a lead box, where the skull was allegedly presented to Napoleon III,[178] before being returned to the Château d'Amboise and re-interred in the Chapel of Saint Hubert in 1874.[177] A new memorial tombstone was added by sculptor Francesco La Monaca in the 1930s.[181] Reflecting doubts about the attribution, a plaque above the tomb states that the remains are only presumed to be those of Leonardo.[177] It has since been theorized that the folding of the skeleton's right arm over the head may correspond to the paralysis of Leonardo's right hand.[178][77][83]
+In 2016, it was announced that DNA tests would be conducted to determine whether the attribution is correct.[177] The DNA of the remains will be compared to that of samples collected from Leonardo's work and his half-brother Domenico's descendants;[177] it may also be sequenced.[182] The lock of hair and ring, now in a private US collection,[ac] were displayed in Vinci beginning on 2 May 2019, the 500th anniversary of the artist's death.[183][89]
+Salvator Mundi, a painting by Leonardo depicting Jesus holding an orb, sold for a world record US$450.3 million at a Christie's auction in New York, 15 November 2017.[184] The highest known sale price for any artwork was previously US$300 million, for Willem de Kooning's Interchange, which was sold privately in September 2015.[185] The highest price previously paid for a work of art at auction was for Pablo Picasso's Les Femmes d'Alger, which sold for US$179.4 million in May 2015 at Christie's New York.[185]
+Early sources
+Modern sources

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+Steel is an alloy of iron with typically a few percent of carbon to improve its strength and fracture resistance compared to iron. Many other additional elements may be present or added. Stainless steels that are corrosion and oxidation resistant need typically an additional 11% chromium. Because of its high tensile strength and low cost, steel is best used in buildings, infrastructure, tools, ships, trains, cars, machines, electrical appliances, and weapons. Iron is the base metal of steel and it can take on two crystalline forms (allotropic forms): body centred cubic and face-centred cubic. These forms depend on temperature. In the body-centred cubic arrangement, there is an iron atom in the centre and eight atoms at the vertices of each cubic unit cell; in the face-centred cubic, there is one atom at the centre of each of the six faces of the cubic unit cell and eight atoms at its vertices.  It is the interaction of the allotropes of iron with the alloying elements, primarily carbon, that gives steel and cast iron their range of unique properties.
+In pure iron, the crystal structure has relatively little resistance to the iron atoms slipping past one another, and so pure iron is quite ductile, or soft and easily formed.  In steel, small amounts of carbon, other elements, and inclusions within the iron act as hardening agents that prevent the movement of dislocations.
+The carbon in typical steel alloys may contribute up to 2.14% of its weight. Varying the amount of carbon and many other alloying elements, as well as controlling their chemical and physical makeup in the final steel (either as solute elements, or as precipitated phases), slows the movement of those dislocations that make pure iron ductile, and thus controls and enhances its qualities. These qualities include the hardness, quenching behaviour, need for annealing, tempering behaviour, yield strength, and tensile strength of the resulting steel. The increase in steel's strength compared to pure iron is possible only by reducing iron's ductility.
+Steel was produced in bloomery furnaces for thousands of years, but its large-scale, industrial use began only after more efficient production methods were devised in the 17th century, with the introduction of the blast furnace and production of crucible steel. This was followed by the open-hearth furnace and then the Bessemer process in England in the mid-19th
+century. With the invention of the Bessemer process, a new era of mass-produced steel began. Mild steel replaced wrought iron.
+Further refinements in the process, such as basic oxygen steelmaking (BOS), largely replaced earlier methods by further lowering the cost of production and increasing the quality of the final product. Today, steel is one of the most common manmade materials in the world, with more than 1.6 billion tons produced annually. Modern steel is generally identified by various grades defined by assorted standards organisations.
+The noun steel originates from the Proto-Germanic adjective stahliją or stakhlijan (made of steel), which is related to stahlaz or stahliją (standing firm).[1]
+The carbon content of steel is between 0.002% and 2.14% by weight for plain carbon steel (iron–carbon alloys).[citation needed] Too little carbon content leaves (pure) iron quite soft, ductile, and weak. Carbon contents higher than those of steel make a brittle alloy commonly called pig iron. Alloy steel is steel to which other alloying elements have been intentionally added to modify the characteristics of steel. Common alloying elements include: manganese, nickel, chromium, molybdenum, boron, titanium, vanadium, tungsten, cobalt, and niobium.[2] In contrast, cast iron does undergo eutectic reaction.  Additional elements, most frequently considered undesirable, are also important in steel: phosphorus, sulfur, silicon, and traces of oxygen, nitrogen, and copper.
+Plain carbon-iron alloys with a higher than 2.1% carbon content are known as cast iron. With modern steelmaking techniques such as powder metal forming, it is possible to make very high-carbon (and other alloy material) steels, but such are not common. Cast iron is not malleable even when hot, but it can be formed by casting as it has a lower melting point than steel and good castability properties.[2] Certain compositions of cast iron, while retaining the economies of melting and casting, can be heat treated after casting to make malleable iron or ductile iron objects. Steel is distinguishable from wrought iron (now largely obsolete), which may contain a small amount of carbon but large amounts of slag.
+Iron is commonly found in the Earth's crust in the form of an ore, usually an iron oxide, such as magnetite or hematite. Iron is extracted from iron ore by removing the oxygen through its combination with a preferred chemical partner such as carbon which is then lost to the atmosphere as carbon dioxide. This process, known as smelting, was first applied to metals with lower melting points, such as tin, which melts at about 250 °C (482 °F), and copper, which melts at about 1,100 °C (2,010 °F), and the combination, bronze, which has a melting point lower than 1,083 °C (1,981 °F). In comparison, cast iron melts at about 1,375 °C (2,507 °F).[3]  Small quantities of iron were smelted in ancient times, in the solid state, by heating the ore in a charcoal fire and then welding the clumps together with a hammer and in the process squeezing out the impurities.  With care, the carbon content could be controlled by moving it around in the fire. Unlike copper and tin, liquid or solid iron dissolves carbon quite readily.
+All of these temperatures could be reached with ancient methods used since the Bronze Age. Since the oxidation rate of iron increases rapidly beyond 800 °C (1,470 °F), it is important that smelting take place in a low-oxygen environment. Smelting, using carbon to reduce iron oxides, results in an alloy (pig iron) that retains too much carbon to be called steel.[3] The excess carbon and other impurities are removed in a subsequent step.
+Other materials are often added to the iron/carbon mixture to produce steel with desired properties. Nickel and manganese in steel add to its tensile strength and make the austenite form of the iron-carbon solution more stable, chromium increases hardness and melting temperature, and vanadium also increases hardness while making it less prone to metal fatigue.[4]
+To inhibit corrosion, at least 11% chromium is added to steel so that a hard oxide forms on the metal surface; this is known as stainless steel. Tungsten slows the formation of cementite, keeping carbon in the iron matrix and allowing martensite to preferentially form at slower quench rates, resulting in high speed steel. On the other hand, sulfur, nitrogen, and phosphorus are considered contaminants that make steel more brittle and are removed from the steel melt during processing.[4]
+The density of steel varies based on the alloying constituents but usually ranges between 7,750 and 8,050 kg/m3 (484 and 503 lb/cu ft), or 7.75 and 8.05 g/cm3 (4.48 and 4.65 oz/cu in).[5]
+Even in a narrow range of concentrations of mixtures of carbon and iron that make a steel, a number of different metallurgical structures, with very different properties can form. Understanding such properties is essential to making quality steel. At room temperature, the most stable form of pure iron is the body-centered cubic (BCC) structure called alpha iron or α-iron. It is a fairly soft metal that can dissolve only a small concentration of carbon, no more than 0.005% at 0 °C (32 °F) and 0.021 wt% at  723 °C (1,333 °F). The inclusion of carbon in alpha iron is called ferrite. At 910 °C, pure iron transforms into a face-centered cubic (FCC) structure, called gamma iron or γ-iron. The inclusion of carbon in gamma iron is called austenite. The more open FCC structure of austenite can dissolve considerably more carbon, as much as 2.1%[6] (38 times that of ferrite) carbon at 1,148 °C (2,098 °F), which reflects the upper carbon content of steel, beyond which is cast iron.[7] When carbon moves out of solution with iron, it forms a very hard, but brittle material called cementite (Fe3C).
+When steels with exactly 0.8% carbon (known as a eutectoid steel), are cooled, the austenitic phase (FCC) of the mixture attempts to revert to the ferrite phase (BCC). The carbon no longer fits within the FCC austenite structure, resulting in an excess of carbon. One way for carbon to leave the austenite is for it to precipitate out of solution as cementite, leaving behind a surrounding phase of BCC iron called ferrite with a small percentage of carbon in solution. The two, ferrite and cementite, precipitate simultaneously producing a layered structure called pearlite, named for its resemblance to mother of pearl. In a hypereutectoid composition (greater than 0.8% carbon), the carbon will first precipitate out as large inclusions of cementite at the austenite grain boundaries until the percentage of carbon in the grains has decreased to the eutectoid composition (0.8% carbon), at which point the pearlite structure forms. For steels that have less than 0.8% carbon (hypoeutectoid), ferrite will first form within the grains until the remaining composition rises to 0.8% of carbon, at which point the pearlite structure will form. No large inclusions of cementite will form at the boundaries in hypoeuctoid steel.[8] The above assumes that the cooling process is very slow, allowing enough time for the carbon to migrate.
+As the rate of cooling is increased the carbon will have less time to migrate to form carbide at the grain boundaries but will have increasingly large amounts of pearlite of a finer and finer structure within the grains; hence the carbide is more widely dispersed and acts to prevent slip of defects within those grains, resulting in hardening of the steel. At the very high cooling rates produced by quenching, the carbon has no time to migrate but is locked within the face-centered austenite and forms martensite. Martensite is a highly strained and stressed, supersaturated form of carbon and iron and is exceedingly hard but brittle. Depending on the carbon content, the martensitic phase takes different forms. Below 0.2% carbon, it takes on a ferrite BCC crystal form, but at higher carbon content it takes a body-centered tetragonal (BCT) structure. There is no thermal activation energy for the transformation from austenite to martensite.[clarification needed] Moreover, there is no compositional change so the atoms generally retain their same neighbors.[9]
+Martensite has a lower density (it expands during the cooling) than does austenite, so that the transformation between them results in a change of volume. In this case, expansion occurs. Internal stresses from this expansion generally take the form of compression on the crystals of martensite and tension on the remaining ferrite, with a fair amount of shear on both constituents. If quenching is done improperly, the internal stresses can cause a part to shatter as it cools. At the very least, they cause internal work hardening and other microscopic imperfections. It is common for quench cracks to form when steel is water quenched, although they may not always be visible.[10]
+There are many types of heat treating processes available to steel. The most common are annealing, quenching, and tempering. Heat treatment is effective on compositions above the eutectoid composition (hypereutectoid) of 0.8% carbon. Hypoeutectoid steel does not benefit from heat treatment.
+Annealing is the process of heating the steel to a sufficiently high temperature to relieve local internal stresses. It does not create a general softening of the product but only locally relieves strains and stresses locked up within the material. Annealing goes through three phases: recovery, recrystallization, and grain growth. The temperature required to anneal a particular steel depends on the type of annealing to be achieved and the alloying constituents.[11]
+Quenching involves heating the steel to create the austenite phase then quenching it in water or oil. This rapid cooling results in a hard but brittle martensitic structure.[9] The steel is then tempered, which is just a specialized type of annealing, to reduce brittleness. In this application the annealing (tempering) process transforms some of the martensite into cementite, or spheroidite and hence it reduces the internal stresses and defects. The result is a more ductile and fracture-resistant steel.[12]
+When iron is smelted from its ore, it contains more carbon than is desirable.  To become steel, it must be reprocessed to reduce the carbon to the correct amount, at which point other elements can be added.  In the past, steel facilities would cast the raw steel product into ingots which would be stored until use in further refinement processes that resulted in the finished product. In modern facilities, the initial product is close to the final composition and is continuously cast into long slabs, cut and shaped into bars and extrusions and heat-treated to produce a final product. Today, approximately 96% of steel is continuously cast, while only 4% is produced as ingots.[13]
+The ingots are then heated in a soaking pit and hot rolled into slabs, billets, or blooms. Slabs are hot or cold rolled into sheet metal or plates. Billets are hot or cold rolled into bars, rods, and wire.  Blooms are hot or cold rolled into structural steel, such as I-beams and rails.  In modern steel mills these processes often occur in one assembly line, with ore coming in and finished steel products coming out.[14] Sometimes after a steel's final rolling, it is heat treated for strength; however, this is relatively rare.[15]
+Steel was known in antiquity and was produced in bloomeries and crucibles.[16][17]
+The earliest known production of steel is seen in pieces of ironware excavated from an archaeological site in Anatolia (Kaman-Kalehöyük) and are nearly 4,000 years old, dating from 1800 BC.[18][19] Horace identifies steel weapons such as the falcata in the Iberian Peninsula, while Noric steel was used by the Roman military.[20]
+The reputation of Seric iron of South India (wootz steel) grew considerably in the rest of the world.[17] Metal production sites in Sri Lanka employed wind furnaces driven by the monsoon winds, capable of producing high-carbon steel. Large-scale Wootz steel production in Tamilakam using crucibles and carbon sources such as the plant Avāram occurred by the sixth century BC, the pioneering precursor to modern steel production and metallurgy.[16][17]
+The Chinese of the Warring States period (403–221 BC) had quench-hardened steel,[21] while Chinese of the Han dynasty (202 BC – 220 AD) created steel by melting together wrought iron with cast iron, gaining an ultimate product of a carbon-intermediate steel by the 1st century AD.[22][23]
+There is evidence that carbon steel was made in Western Tanzania by the ancestors of the Haya people as early as 2,000 years ago by a complex process of "pre-heating" allowing temperatures inside a furnace to reach 1300 to 1400 °C.[24][25][26][27][28][29]
+Evidence of the earliest production of high carbon steel in India are found in Kodumanal in Tamil Nadu, the Golconda area in Andhra Pradesh  and Karnataka, and in the Samanalawewa areas of Sri Lanka.[30] This came to be known as Wootz steel, produced in South India by about sixth century BC and exported globally.[31][32] The steel technology existed prior to 326 BC in the region as they are mentioned in literature of Sangam Tamil, Arabic and Latin as the finest steel in the world exported to the Romans, Egyptian, Chinese and Arab worlds at that time – what they called Seric Iron.[33] A 200 BC Tamil trade guild in Tissamaharama, in the South East of Sri Lanka, brought with them some of the oldest iron and steel artifacts and production processes to the island from the classical period.[34][35][36] The Chinese and locals in Anuradhapura, Sri Lanka had also adopted the production methods of creating Wootz steel from the Chera Dynasty Tamils of South India by the 5th century AD.[37][38] In Sri Lanka, this early steel-making method employed a unique wind furnace, driven by the monsoon winds, capable of producing high-carbon steel.[39][40] Since the technology was acquired from the Tamilians from South India,[citation needed] the origin of steel technology in India can be conservatively estimated at 400–500 BC.[31][40]
+The manufacture of what came to be called Wootz, or Damascus steel, famous for its durability and ability to hold an edge, may have been taken by the Arabs from Persia, who took it from India. It was originally created from a number of different materials including various trace elements, apparently ultimately from the writings of Zosimos of Panopolis. In 327 BC, Alexander the Great was rewarded by the defeated King Porus, not with gold or silver but with 30 pounds of steel.[41]  Recent studies have suggested that carbon nanotubes were included in its structure, which might explain some of its legendary qualities, though given the technology of that time, such qualities were produced by chance rather than by design.[42] Natural wind was used where the soil containing iron was heated by the use of wood. The ancient Sinhalese managed to extract a ton of steel for every 2 tons of soil,[39] a remarkable feat at the time. One such furnace was found in Samanalawewa and archaeologists were able to produce steel as the ancients did.[39][43]
+Crucible steel, formed by slowly heating and cooling pure iron and carbon (typically in the form of charcoal) in a crucible, was produced in Merv by the 9th to 10th century AD.[32] In the 11th century, there is evidence of the production of steel in Song China using two techniques: a "berganesque" method that produced inferior, inhomogeneous steel, and a precursor to the modern Bessemer process that used partial decarbonization via repeated forging under a cold blast.[44]
+Since the 17th century, the first step in European steel production has been the smelting of iron ore into pig iron in a blast furnace.[45] Originally employing charcoal, modern methods use coke, which has proven more economical.[46][47][48]
+In these processes pig iron was refined (fined) in a finery forge to produce bar iron, which was then used in steel-making.[45]
+The production of steel by the cementation process was described in a treatise published in Prague in 1574 and was in use in Nuremberg from 1601. A similar process for case hardening armor and files was described in a book published in Naples in 1589. The process was introduced to England in about 1614 and used to produce such steel by Sir Basil Brooke at Coalbrookdale during the 1610s.[49]
+The raw material for this process were bars of iron. During the 17th century it was realized that the best steel came from oregrounds iron of a region north of Stockholm, Sweden.  This was still the usual raw material source in the 19th century, almost as long as the process was used.[50][51]
+Crucible steel is steel that has been melted in a crucible rather than having been forged, with the result that it is more homogeneous. Most previous furnaces could not reach high enough temperatures to melt the steel. The early modern crucible steel industry resulted from the invention of Benjamin Huntsman in the 1740s. Blister steel (made as above) was melted in a crucible or in a furnace, and cast (usually) into ingots.[51][52]
+The modern era in steelmaking began with the introduction of Henry Bessemer's Bessemer process in 1855, the raw material for which was pig iron.[53] His method let him produce steel in large quantities cheaply, thus mild steel came to be used for most purposes for which wrought iron was formerly used.[54] The Gilchrist-Thomas process (or basic Bessemer process) was an improvement to the Bessemer process, made by lining the converter with a basic material to remove phosphorus.
+Another 19th-century steelmaking process was the Siemens-Martin process, which complemented the Bessemer process.[51]  It consisted of co-melting bar iron (or steel scrap) with pig iron.
+These methods of steel production were rendered obsolete by the Linz-Donawitz process of basic oxygen steelmaking (BOS), developed in 1952,[55] and other oxygen steel making methods. Basic oxygen steelmaking is superior to previous steelmaking methods because the oxygen pumped into the furnace limited impurities, primarily nitrogen, that previously had entered from the air used,[56] and because, with respect to the open-hearth process, the same quantity of steel from a BOS process is manufactured in one-twelfth the time.[55] Today, electric arc furnaces (EAF) are a common method of reprocessing scrap metal to create new steel. They can also be used for converting pig iron to steel, but they use a lot of electrical energy (about 440 kWh per metric ton), and are thus generally only economical when there is a plentiful supply of cheap electricity.[57]
+The steel industry is often considered an indicator of economic progress, because of the critical role played by steel in infrastructural and overall economic development.[58] In 1980, there were more than 500,000 U.S. steelworkers. By 2000, the number of steelworkers fell to 224,000.[59]
+The economic boom in China and India caused a massive increase in the demand for steel. Between 2000 and 2005, world steel demand increased by 6%. Since 2000, several Indian[60] and Chinese steel firms have risen to prominence,[according to whom?] such as Tata Steel (which bought Corus Group in 2007), Baosteel Group and Shagang Group. As of 2017[update], though, ArcelorMittal is the world's largest steel producer.[61] In 2005, the British Geological Survey stated China was the top steel producer with about one-third of the world share; Japan, Russia, and the US followed respectively.[62]
+In 2008, steel began trading as a commodity on the London Metal Exchange. At the end of 2008, the steel industry faced a sharp downturn that led to many cut-backs.[63]
+Steel is one of the world's most-recycled materials, with a recycling rate of over 60% globally;[64] in the United States alone, over 82,000,000 metric tons (81,000,000 long tons; 90,000,000 short tons) were recycled in the year 2008, for an overall recycling rate of 83%.[65]
+As more steel is produced than is scrapped, the amount of recycled raw materials is about 40% of the total of steel produced - in 2016, 1,628,000,000 tonnes (1.602×109 long tons; 1.795×109 short tons) of crude steel was produced globally, with 630,000,000 tonnes (620,000,000 long tons; 690,000,000 short tons) recycled.[66]
+Modern steels are made with varying combinations of alloy metals to fulfill many purposes.[4] Carbon steel, composed simply of iron and carbon, accounts for 90% of steel production.[2] Low alloy steel is alloyed with other elements, usually molybdenum, manganese, chromium, or nickel, in amounts of up to 10% by weight to improve the hardenability of thick sections.[2] High strength low alloy steel has small additions (usually < 2% by weight) of other elements, typically 1.5% manganese, to provide additional strength for a modest price increase.[67]
+Recent Corporate Average Fuel Economy (CAFE) regulations have given rise to a new variety of steel known as Advanced High Strength Steel (AHSS). This material is both strong and ductile so that vehicle structures can maintain their current safety levels while using less material. There are several commercially available grades of AHSS, such as dual-phase steel, which is heat-treated to contain both a ferritic and martensitic microstructure to produce formable, high strength steel.[68] Transformation Induced Plasticity (TRIP) steel involves special alloying and heat treatments to stabilize amounts of austenite at room temperature in normally austenite-free low-alloy ferritic steels. By applying strain, the austenite undergoes a phase transition to martensite without the addition of heat.[69] Twinning Induced Plasticity (TWIP) steel uses a specific type of strain to increase the effectiveness of work hardening on the alloy.[70]
+Carbon Steels are often galvanized, through hot-dip or electroplating in zinc for protection against rust.[71]
+Stainless steels contain a minimum of 11% chromium, often combined with nickel, to resist corrosion. Some stainless steels, such as the ferritic stainless steels are magnetic, while others, such as the austenitic, are nonmagnetic.[72] Corrosion-resistant steels are abbreviated as CRES.
+Some more modern steels include tool steels, which are alloyed with large amounts of tungsten and cobalt or other elements to maximize solution hardening. This also allows the use of precipitation hardening and improves the alloy's temperature resistance.[2] Tool steel is generally used in axes, drills, and other devices that need a sharp, long-lasting cutting edge. Other special-purpose alloys include weathering steels such as Cor-ten, which weather by acquiring a stable, rusted surface, and so can be used un-painted.[73] Maraging steel is alloyed with nickel and other elements, but unlike most steel contains little carbon (0.01%). This creates a very strong but still malleable steel.[74]
+Eglin steel uses a combination of over a dozen different elements in varying amounts to create a relatively low-cost steel for use in bunker buster weapons. Hadfield steel (after Sir Robert Hadfield) or manganese steel contains 12–14% manganese which when abraded strain-hardens to form a very hard skin which resists wearing. Examples include tank tracks, bulldozer blade edges and cutting blades on the jaws of life.[75]
+Most of the more commonly used steel alloys are categorized into various grades by standards organizations. For example, the Society of Automotive Engineers has a series of grades defining many types of steel.[76] The American Society for Testing and Materials has a separate set of standards, which define alloys such as A36 steel, the most commonly used structural steel in the United States.[77] The JIS also define series of steel grades that are being used extensively in Japan as well as in developing countries.
+Iron and steel are used widely in the construction of roads, railways, other infrastructure, appliances, and buildings. Most large modern structures, such as stadiums and skyscrapers, bridges, and airports, are supported by a steel skeleton. Even those with a concrete structure employ steel for reinforcing. In addition, it sees widespread use in major appliances and cars. Despite growth in usage of aluminium, it is still the main material for car bodies. Steel is used in a variety of other construction materials, such as bolts, nails and screws and other household products and cooking utensils.[78]
+Other common applications include shipbuilding, pipelines, mining, offshore construction, aerospace, white goods (e.g. washing machines), heavy equipment such as bulldozers, office furniture, steel wool, tool and armour in the form of personal vests or vehicle armour (better known as rolled homogeneous armour in this role).
+Before the introduction of the Bessemer process and other modern production techniques, steel was expensive and was only used where no cheaper alternative existed, particularly for the cutting edge of knives, razors, swords, and other items where a hard, sharp edge was needed. It was also used for springs, including those used in clocks and watches.[51]
+With the advent of speedier and thriftier production methods, steel has become easier to obtain and much cheaper. It has replaced wrought iron for a multitude of purposes. However, the availability of plastics in the latter part of the 20th century allowed these materials to replace steel in some applications due to their lower fabrication cost and weight.[79] Carbon fiber is replacing steel in some cost insensitive applications such as sports equipment and high end automobiles.
+Steel manufactured after World War II became contaminated with radionuclides by nuclear weapons testing. Low-background steel, steel manufactured prior to 1945, is used for certain radiation-sensitive applications such as Geiger counters and radiation shielding.

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+Mesothelae
+Opisthothelae
+ See Spider taxonomy.
+Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom,[2] and spinnerets that extrude silk.[3] They are the largest order of arachnids and rank seventh in total species diversity among all orders of organisms.[4] Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of July 2019[update], at least 48,200 spider species, and 120 families have been recorded by taxonomists.[1] However, there has been dissension within the scientific community as to how all these families should be classified, as evidenced by the over 20 different classifications that have been proposed since 1900.[5]
+Anatomically, spiders (as with all arachnids) differ from other arthropods in that the usual body segments are fused into two tagmata, the prosoma, or cephalothorax, and opisthosoma, or abdomen, and joined by a small, cylindrical pedicel (however, as there is currently neither paleontological nor embryological evidence that spiders ever had a separate thorax-like division, there exists an argument against the validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly, arguments can be formed against use of the term abdomen, as the opisthosoma of all spiders contains a heart and respiratory organs, organs atypical of an abdomen[6]). Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.
+Their abdomens bear appendages that have been modified into spinnerets that extrude silk from up to six types of glands. Spider webs vary widely in size, shape and the amount of sticky thread used. It now appears that the spiral orb web may be one of the earliest forms, and spiders that produce tangled cobwebs are more abundant and diverse than orb-web spiders. Spider-like arachnids with silk-producing spigots appeared in the Devonian period about 386 million years ago, but these animals apparently lacked spinnerets. True spiders have been found in Carboniferous rocks from 318 to 299 million years ago, and are very similar to the most primitive surviving suborder, the Mesothelae. The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, before 200 million years ago.
+The species Bagheera kiplingi was described as herbivorous in 2008,[7] but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. It is estimated that the world's 25 million tons of spiders kill 400–800 million tons of prey per year.[8] Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, so they liquefy their food by flooding it with digestive enzymes. They also grind food with the bases of their pedipalps, as arachnids do not have the mandibles that crustaceans and insects have.
+To avoid being eaten by the females, which are typically much larger, male spiders identify themselves to potential mates by a variety of complex courtship rituals. Males of most species survive a few matings, limited mainly by their short life spans. Females weave silk egg-cases, each of which may contain hundreds of eggs. Females of many species care for their young, for example by carrying them around or by sharing food with them. A minority of species are social, building communal webs that may house anywhere from a few to 50,000 individuals. Social behavior ranges from precarious toleration, as in the widow spiders, to co-operative hunting and food-sharing. Although most spiders live for at most two years, tarantulas and other mygalomorph spiders can live up to 25 years in captivity.
+While the venom of a few species is dangerous to humans, scientists are now researching the use of spider venom in medicine and as non-polluting pesticides. Spider silk provides a combination of lightness, strength and elasticity that is superior to that of synthetic materials, and spider silk genes have been inserted into mammals and plants to see if these can be used as silk factories. As a result of their wide range of behaviors, spiders have become common symbols in art and mythology symbolizing various combinations of patience, cruelty and creative powers. An abnormal fear of spiders is called arachnophobia.
+1: pedipalp
+ 2: trichobothria
+ 3: carapace of prosoma (cephalothorax)
+ 4: opisthosoma (abdomen)
+ 5: eyes – AL (anterior lateral)
+    AM (anterior median)
+    PL (posterior lateral)
+    PM (posterior median)
+Leg segments:
+ 6: costa
+ 7: trochanter
+ 8: patella
+ 9: tibia
+10: metatarsus
+11: tarsus
+13: claw
+14: chelicera
+15: sternum of prosoma
+16: pedicel (also called pedicle)
+17: book lung sac
+18: book lung stigma
+19: epigastric fold
+20: epigyne
+21: anterior spinneret
+22: posterior spinneret
+Spiders are chelicerates and therefore arthropods.[9] As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo.[10] Being chelicerates, their bodies consist of two tagmata, sets of segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete fusion of the segments that in an insect would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma.[9] In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel.[11] The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws".[10][12] The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.[9]
+Spiders and scorpions are members of one chelicerate group, the arachnids.[12] Scorpions' chelicerae have three sections and are used in feeding.[13] Spiders' chelicerae have two sections and terminate in fangs that are generally venomous, and fold away behind the upper sections while not in use. The upper sections generally have thick "beards" that filter solid lumps out of their food, as spiders can take only liquid food.[11] Scorpions' pedipalps generally form large claws for capturing prey,[13] while those of spiders are fairly small appendages whose bases also act as an extension of the mouth; in addition, those of male spiders have enlarged last sections used for sperm transfer.[11]
+In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex carapace, while the underside is covered by two rather flat plates. The abdomen is soft and egg-shaped. It shows no sign of segmentation, except that the primitive Mesothelae, whose living members are the Liphistiidae, have segmented plates on the upper surface.[11]
+Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end.[14] However, in spiders, it occupies only the upper part of the abdomen, and blood is discharged into the hemocoel by one artery that opens at the rear end of the abdomen and by branching arteries that pass through the pedicle and open into several parts of the cephalothorax. Hence spiders have open circulatory systems.[11] The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.[12]
+Spiders have developed several different respiratory anatomies, based on book lungs, a tracheal system, or both. Mygalomorph and Mesothelae spiders have two pairs of book lungs filled with haemolymph, where openings on the ventral surface of the abdomen allow air to enter and diffuse oxygen. This is also the case for some basal araneomorph spiders, like the family Hypochilidae, but the remaining members of this group have just the anterior pair of book lungs intact while the posterior pair of breathing organs are partly or fully modified into tracheae, through which oxygen is diffused into the haemolymph or directly to the tissue and organs.[11] The tracheal system has most likely evolved in small ancestors to help resist desiccation.[12] The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets.[11] Spiders that have tracheae generally have higher metabolic rates and better water conservation.[15] Spiders are ectotherms, so environmental temperatures affect their activity.[16]
+Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae.[11] The families Uloboridae and Holarchaeidae, and some Liphistiidae spiders, have lost their venom glands, and kill their prey with silk instead.[17] Like most arachnids, including scorpions,[12] spiders have a narrow gut that can only cope with liquid food and two sets of filters to keep solids out.[11] They use one of two different systems of external digestion. Some pump digestive enzymes from the midgut into the prey and then suck the liquified tissues of the prey into the gut, eventually leaving behind the empty husk of the prey. Others grind the prey to pulp using the chelicerae and the bases of the pedipalps, while flooding it with enzymes; in these species, the chelicerae and the bases of the pedipalps form a preoral cavity that holds the food they are processing.[11]
+The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The midgut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.[11]
+Most spiders convert nitrogenous waste products into uric acid, which can be excreted as a dry material. Malphigian tubules ("little tubes") extract these wastes from the blood in the hemocoel and dump them into the cloacal chamber, from which they are expelled through the anus.[11] Production of uric acid and its removal via Malphigian tubules are a water-conserving feature that has evolved independently in several arthropod lineages that can live far away from water,[18] for example the tubules of insects and arachnids develop from completely different parts of the embryo.[12] However, a few primitive spiders, the suborder Mesothelae and infraorder Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"),[11] which use large amounts of water to excrete nitrogenous waste products as ammonia.[18]
+The basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia.[19] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen;[11][12][19] in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.[15]
+Despite the relatively small central nervous system, some spiders (like Portia) exhibit complex behaviour, including the ability to use a trial-and-error approach.[20][21]
+Spiders have primarily four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another.[11] The principal pair at the front are of the type called pigment-cup ocelli ("little eyes"), which in most arthropods are only capable of detecting the direction from which light is coming, using the shadow cast by the walls of the cup. However, in spiders these eyes are capable of forming images.[22][23] The other pairs, called secondary eyes, are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the principal eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torchlight reflected from the tapeta. On the other hand, jumping spiders' secondary eyes have no tapeta.[11]
+Other differences between the principal and secondary eyes are that the latter have rhabdomeres that point away from incoming light, just like in vertebrates, while the arrangement is the opposite in the former. The principal eyes are also the only ones with eye muscles, allowing them to move the retina. Having no muscles, the secondary eyes are immobile.[24]
+Some jumping spiders' visual acuity exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects; in fact the human eye is only about five times sharper than a jumping spider's. They achieve this by a telephotographic series of lenses, a four-layer retina and the ability to swivel their eyes and integrate images from different stages in the scan. The downside is that the scanning and integrating processes are relatively slow.[20]
+There are spiders with a reduced number of eyes. Of these, those with six eyes (such as Periegops suterii) are the most numerous and are missing a pair of eyes on the anterior median line;[25] other species have four eyes and some just two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.
+As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae.[22] An adult Araneus may have up to 1,000 such chemosensitive setae, most on the tarsi of the first pair of legs. Males have more chemosensitive bristles on their pedipalps than females. They have been shown to be responsive to sex pheromones produced by females, both contact and air-borne.[26] The jumping spider Evarcha culicivora uses the scent of blood from mammals and other vertebrates, which is obtained by capturing blood-filled mosquitoes, to attract the opposite sex. Because they are able to tell the sexes apart, it is assumed the blood scent is mixed with pheromones.[27] Spiders also have in the joints of their limbs slit sensillae that detect force and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.[11]
+Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' proprioceptors, sensors that report the force exerted by muscles and the degree of bending in the body and joints, are well-understood. On the other hand, little is known about what other internal sensors spiders or other arthropods may have.[22]
+Each of the eight legs of a spider consists of seven distinct parts. The part closest to and attaching the leg to the cephalothorax is the coxa; the next segment is the short trochanter that works as a hinge for the following long segment, the femur; next is the spider's knee, the patella, which acts as the hinge for the tibia; the metatarsus is next, and it connects the tibia to the tarsus (which may be thought of as a foot of sorts); the tarsus ends in a claw made up of either two or three points, depending on the family to which the spider belongs. Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors.[28] The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter).[29] As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up.[11] Spiders can generate pressures up to eight times their resting level to extend their legs,[30] and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs.[11] Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.[29]
+Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine bristles between the paired claws at the tips of their legs. These tufts, known as scopulae, consist of bristles whose ends are split into as many as 1,000 branches, and enable spiders with scopulae to walk up vertical glass and upside down on ceilings. It appears that scopulae get their grip from contact with extremely thin layers of water on surfaces.[11] Spiders, like most other arachnids, keep at least four legs on the surface while walking or running.[31]
+The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk.[11]
+Silk is mainly composed of a protein very similar to that used in insect silk. It is initially a liquid, and hardens not by exposure to air but as a result of being drawn out, which changes the internal structure of the protein.[32] It is similar in tensile strength to nylon and biological materials such as chitin, collagen and cellulose, but is much more elastic. In other words, it can stretch much further before breaking or losing shape.[11]
+Some spiders have a cribellum, a modified spinneret with up to 40,000 spigots, each of which produces a single very fine fiber. The fibers are pulled out by the calamistrum, a comblike set of bristles on the jointed tip of the cribellum, and combined into a composite woolly thread that is very effective in snagging the bristles of insects. The earliest spiders had cribella, which produced the first silk capable of capturing insects, before spiders developed silk coated with sticky droplets. However, most modern groups of spiders have lost the cribellum.[11]
+Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a "safety rope"; for nest-building; and as "parachutes" by the young of some species.[11]
+Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods,[33] male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs onto which they ejaculate and then transfer the sperm to special syringe-styled structures, palpal bulbs or palpal organs, borne on the tips of the pedipalps of mature males. When a male detects signs of a female nearby he checks whether she is of the same species and whether she is ready to mate; for example in species that produce webs or "safety ropes", the male can identify the species and sex of these objects by "smell".[11]
+Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female via one or two openings on the underside of her abdomen.[11]
+Female spiders' reproductive tracts are arranged in one of two ways. The ancestral arrangement ("haplogyne" or "non-entelegyne") consists of a single genital opening, leading to two seminal receptacles (spermathecae) in which females store sperm. In the more advanced arrangement ("entelegyne" ), there are two further openings leading directly to the spermathecae, creating a "flow through" system rather than a "first-in first-out" one. Eggs are as a general rule only fertilized during oviposition when the stored sperm is released from its chamber, rather than in the ovarian cavity.[34] A few exceptions exist, such as Parasteatoda tepidariorum. In these species the female appears to be able to activate the dormant sperm before oviposition, allowing them to migrate to the ovarian cavity where fertilization occurs.[35][36][37] The only known example of direct fertilization between male and female is an Israeli spider, Harpactea sadistica, which has evolved traumatic insemination. In this species the male will penetrate its pedipalps through the female's body wall and inject his sperm directly into her ovaries, where the embryos inside the fertilized eggs will start to develop before being laid.[38]
+Males of the genus Tidarren amputate one of their palps before maturation and enter adult life with one palp only. The palps are 20% of the male's body mass in this species, and detaching one of the two improves mobility. In the Yemeni species Tidarren argo, the remaining palp is then torn off by the female. The separated palp remains attached to the female's epigynum for about four hours and apparently continues to function independently. In the meantime, the female feeds on the palpless male.[39] In over 60% of cases, the female of the Australian redback spider kills and eats the male after it inserts its second palp into the female's genital opening; in fact, the males co-operate by trying to impale themselves on the females' fangs. Observation shows that most male redbacks never get an opportunity to mate, and the "lucky" ones increase the likely number of offspring by ensuring that the females are well-fed.[40] However, males of most species survive a few matings, limited mainly by their short life spans. Some even live for a while in their mates' webs.[41]
+The tiny male of the Golden orb weaver (Trichonephila clavipes) (near the top of the leaf) is protected from the female by producing the right vibrations in the web, and may be too small to be worth eating.
+Orange spider egg sac hanging from ceiling
+Gasteracantha mammosa spiderlings next to their eggs capsule
+Wolf spider carrying its young on its abdomen
+Females lay up to 3,000 eggs in one or more silk egg sacs,[11] which maintain a fairly constant humidity level.[41] In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.[11]
+Baby spiders pass all their larval stages inside the egg and hatch as spiderlings, very small and sexually immature but similar in shape to adults. Some spiders care for their young, for example a wolf spider's brood clings to rough bristles on the mother's back,[11] and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.[41]
+Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch.[42] In some species males mate with newly-molted females, which are too weak to be dangerous to the males.[41] Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years,[11][43] and an Australian female trapdoor spider was documented to have lived in the wild for 43 years, dying of a parasitic wasp attack.[44]
+Spiders occur in a large range of sizes. The smallest, Patu digua from Colombia, are less than 0.37 mm (0.015 in) in body length. The largest and heaviest spiders occur among tarantulas, which can have body lengths up to 90 mm (3.5 in) and leg spans up to 250 mm (9.8 in).[45]
+Only three classes of pigment (ommochromes, bilins and guanine) have been identified in spiders, although other pigments have been detected but not yet characterized. Melanins, carotenoids and pterins, very common in other animals, are apparently absent. In some species, the exocuticle of the legs and prosoma is modified by a tanning process, resulting in a brown coloration.[46]
+Bilins are found, for example, in Micrommata virescens, resulting in its green color. Guanine is responsible for the white markings of the European garden spider Araneus diadematus. It is in many species accumulated in specialized cells called guanocytes. In genera such as Tetragnatha, Leucauge, Argyrodes or Theridiosoma, guanine creates their silvery appearance. While guanine is originally an end-product of protein metabolism, its excretion can be blocked in spiders, leading to an increase in its storage.[46] Structural colors occur in some species, which are the result of the diffraction, scattering or interference of light, for example by modified setae or scales. The white prosoma of Argiope results from bristles reflecting the light, Lycosa and Josa both have areas of modified cuticle that act as light reflectors.[46]
+While in many spiders color is fixed throughout their lifespan, in some groups, color may be variable in response to environmental and internal conditions.[46] Choice of prey may be able to alter the color of spiders. For example, the abdomen of Theridion grallator will become orange if the spider ingests certain species of Diptera and adult Lepidoptera, but if it consumes Homoptera or larval Lepidoptera, then the abdomen becomes green.[47] Environmentally induced color changes may be morphological (occurring over several days) or physiological (occurring near instantly). Morphological changes require pigment synthesis and degradation. In contrast to this, physiological changes occur by changing the position of pigment-containing cells.[46] An example of morphological color changes is background matching. Misumena vatia for instance can change its body color to match the substrate it lives on which makes it more difficult to be detected by prey.[48] An example of physiological color change is observed in Cyrtophora cicatrosa, which can change its body color from white to brown near instantly.[46]
+Although spiders are generally regarded as predatory, the jumping spider Bagheera kiplingi gets over 90% of its food from fairly solid plant material produced by acacias as part of a mutually beneficial relationship with a species of ant.[49]
+Juveniles of some spiders in the families Anyphaenidae, Corinnidae, Clubionidae, Thomisidae and Salticidae feed on plant nectar. Laboratory studies show that they do so deliberately and over extended periods, and periodically clean themselves while feeding. These spiders also prefer sugar solutions to plain water, which indicates that they are seeking nutrients. Since many spiders are nocturnal, the extent of nectar consumption by spiders may have been underestimated. Nectar contains amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.[50]
+Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.[50]
+The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.[11]
+Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it.[51] A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.[11]
+Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus Hyptiotes and the family Theridiosomatidae stretch their webs and then release them when prey strike them, but do not actively move their webs. Those of the family Deinopidae weave even smaller webs, hold them outstretched between their first two pairs of legs, and lunge and push the webs as much as twice their own body length to trap prey, and this move may increase the webs' area by a factor of up to ten. Experiments have shown that Deinopis spinosus has two different techniques for trapping prey: backwards strikes to catch flying insects, whose vibrations it detects; and forward strikes to catch ground-walking prey that it sees. These two techniques have also been observed in other deinopids. Walking insects form most of the prey of most deinopids, but one population of Deinopis subrufa appears to live mainly on tipulid flies that they catch with the backwards strike.[52]
+Mature female bolas spiders of the genus Mastophora build "webs" that consist of only a single "trapeze line", which they patrol. They also construct a bolas made of a single thread, tipped with a large ball of very wet sticky silk. They emit chemicals that resemble the pheromones of moths, and then swing the bolas at the moths. Although they miss on about 50% of strikes, they catch about the same weight of insects per night as web-weaving spiders of similar size. The spiders eat the bolas if they have not made a kill in about 30 minutes, rest for a while, and then make new bolas.[53][54] Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.[55]
+The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush predators that lurk in burrows, often closed by trapdoors and often surrounded by networks of silk threads that alert these spiders to the presence of prey.[15] Other ambush predators do without such aids, including many crab spiders,[11] and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking.[46] Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.[11]
+Some jumping spiders of the genus Portia hunt other spiders in ways that seem intelligent,[20] outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species.[56] However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.[20]
+Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective bristles to resemble the shiny bodies of ants. In some spider species, males and females mimic different ant species, as female spiders are usually much larger than males. Ant-mimicking spiders also modify their behavior to resemble that of the target species of ant; for example, many adopt a zig-zag pattern of movement, ant-mimicking jumping spiders avoid jumping, and spiders of the genus Synemosyna walk on the outer edges of leaves in the same way as Pseudomyrmex. Ant mimicry in many spiders and other arthropods may be for protection from predators that hunt by sight, including birds, lizards and spiders. However, several ant-mimicking spiders prey either on ants or on the ants' "livestock", such as aphids. When at rest, the ant-mimicking crab spider Amyciaea does not closely resemble Oecophylla, but while hunting it imitates the behavior of a dying ant to attract worker ants. After a kill, some ant-mimicking spiders hold their victims between themselves and large groups of ants to avoid being attacked.[57]
+There is strong evidence that spiders' coloration is camouflage that helps them to evade their major predators, birds and parasitic wasps, both of which have good color vision. Many spider species are colored so as to merge with their most common backgrounds, and some have disruptive coloration, stripes and blotches that break up their outlines. In a few species, such as the Hawaiian happy-face spider, Theridion grallator, several coloration schemes are present in a ratio that appears to remain constant, and this may make it more difficult for predators to recognize the species. Most spiders are insufficiently dangerous or unpleasant-tasting for warning coloration to offer much benefit. However, a few species with powerful venom, large jaws or irritant bristles have patches of warning colors, and some actively display these colors when threatened.[46][58]
+Many of the family Theraphosidae, which includes tarantulas and baboon spiders, have urticating hairs on their abdomens and use their legs to flick them at attackers. These bristles are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom.[59] A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles.[60] The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.[61]
+A few spider species that build webs live together in large colonies and show social behavior, although not as complex as in social insects. Anelosimus eximius (in the family Theridiidae) can form colonies of up to 50,000 individuals.[62] The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social.[63] Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food.[64] Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae).[65] Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this.[66] The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings.[49] Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.[67]
+There is no consistent relationship between the classification of spiders and the types of web they build: species in the same genus may build very similar or significantly different webs. Nor is there much correspondence between spiders' classification and the chemical composition of their silks. Convergent evolution in web construction, in other words use of similar techniques by remotely related species, is rampant. Orb web designs and the spinning behaviors that produce them are the best understood. The basic radial-then-spiral sequence visible in orb webs and the sense of direction required to build them may have been inherited from the common ancestors of most spider groups.[68] However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a subgroup that evolved from orb-web spiders, and non-orb spiders have over 40% more species and are four times as abundant as orb-web spiders. Their greater success may be because sphecid wasps, which are often the dominant predators of spiders, much prefer to attack spiders that have flat webs.[69]
+About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey.[68]
+The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.[68]
+Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the backlighting from the sky; enabling oscillations to catch insects in slow horizontal flight. However, there is no single explanation for the common use of horizontal orb webs.[68]
+Spiders often attach highly visible silk bands, called decorations or stabilimenta, to their webs. Field research suggests that webs with more decorative bands captured more prey per hour.[70] However, a laboratory study showed that spiders reduce the building of these decorations if they sense the presence of predators.[71]
+There are several unusual variants of orb web, many of them convergently evolved, including: attachment of lines to the surface of water, possibly to trap insects in or on the surface; webs with twigs through their centers, possibly to hide the spiders from predators; "ladderlike" webs that appear most effective in catching moths. However, the significance of many variations is unclear.[68]
+In 1973, Skylab 3 took two orb-web spiders into space to test their web-spinning capabilities in zero gravity. At first, both produced rather sloppy webs, but they adapted quickly.[72]
+Members of the family Theridiidae weave irregular, tangled, three-dimensional webs, popularly known as cobwebs. There seems to be an evolutionary trend towards a reduction in the amount of sticky silk used, leading to its total absence in some species. The construction of cobwebs is less stereotyped than that of orb-webs, and may take several days.[69]
+The Linyphiidae generally make horizontal but uneven sheets, with tangles of stopping threads above. Insects that hit the stopping threads fall onto the sheet or are shaken onto it by the spider, and are held by sticky threads on the sheet until the spider can attack from below.[73]
+Although the fossil record of spiders is considered poor,[74] almost 1000 species have been described from fossils.[75] Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber.[75] The oldest known amber that contains fossil arthropods dates from 130 million years ago in the Early Cretaceous period. In addition to preserving spiders' anatomy in very fine detail, pieces of amber show spiders mating, killing prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached;[76] the oldest fossil web found so far is 100 million years old.[77] Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.[76]
+The oldest known exclusively terrestrial arachnid is the trigonotarbid Palaeotarbus jerami, from about 420 million years ago in the Silurian period, and had a triangular cephalothorax and segmented abdomen, as well as eight legs and a pair of pedipalps.[78] Attercopus fimbriunguis, from 386 million years ago in the Devonian period, bears the earliest known silk-producing spigots, and was therefore hailed as a spider at the time of its discovery.[79] However, these spigots may have been mounted on the underside of the abdomen rather than on spinnerets, which are modified appendages and whose mobility is important in the building of webs. Hence Attercopus and the similar Permian arachnid Permarachne may not have been true spiders, and probably used silk for lining nests or producing egg cases rather than for building webs.[3] The largest known fossil spider as of 2011 is the araneid Nephila jurassica, from about 165 million years ago, recorded from Daohuogo, Inner Mongolia in China.[80] Its body length is almost 25 mm, (i.e., almost one inch).
+Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae.[79]
+The mesothelid Paleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets.[81] Although the Permian period 299 to 251 million years ago saw rapid diversification of flying insects, there are very few fossil spiders from this period.[79]
+The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appear in the Triassic well before 200 million years ago. Some Triassic mygalomorphs appear to be members of the family Hexathelidae, whose modern members include the notorious Sydney funnel-web spider, and their spinnerets appear adapted for building funnel-shaped webs to catch jumping insects. Araneomorphae account for the great majority of modern spiders, including those that weave the familiar orb-shaped webs. The Jurassic and Cretaceous periods provide a large number of fossil spiders, including representatives of many modern families.[79]
+Xiphosura (horseshoe crabs)
+Eurypterida†
+Chasmataspidida†
+Scorpiones
+Opiliones (harvestmen)
+Pseudoscorpiones
+Solifugae (sun spiders)
+Palpigradi (microwhip scorpions)
+Trigonotarbida†
+Araneae (spiders)
+Haptopoda†
+Amblypygi (whip spiders)
+Thelyphonida (whip scorpions)
+Schizomida
+Ricinulei (hooded tickspiders)
+Anactinotrichida
+Acariformes (mites)
+It is now agreed that spiders (Araneae) are monophyletic (i.e., members of a group of organisms that form a clade, consisting of a last common ancestor and all of its descendants).[83] There has been debate about what their closest evolutionary relatives are, and how all of these evolved from the ancestral chelicerates, which were marine animals. The cladogram on the right is based on J. W. Shultz' analysis (2007). Other views include proposals that: scorpions are more closely related to the extinct marine scorpioid eurypterids than to spiders; spiders and Amblypygi are a monophyletic group. The appearance of several multi-way branchings in the tree on the right shows that there are still uncertainties about relationships between the groups involved.[83]
+Arachnids lack some features of other chelicerates, including backward-pointing mouths and gnathobases ("jaw bases") at the bases of their legs;[83] both of these features are part of the ancestral arthropod feeding system.[84] Instead, they have mouths that point forwards and downwards, and all have some means of breathing air.[83] Spiders (Araneae) are distinguished from other arachnid groups by several characteristics, including spinnerets and, in males, pedipalps that are specially adapted for sperm transfer.[85]
+Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two infraorders, Mygalomorphae and Araneomorphae. Over 48,000 living species of spiders (order Araneae) have been identified and as of 2019 grouped into 120 families and about 4,100 genera by arachnologists.[1]
+The only living members of the primitive Mesothelae are the family Liphistiidae, found only in Southeast Asia, China, and Japan.[85] Most of the Liphistiidae construct silk-lined burrows with thin trapdoors, although some species of the genus Liphistius build camouflaged silk tubes with a second trapdoor as an emergency exit. Members of the genus Liphistius run silk "tripwires" outwards from their tunnels to help them detect approaching prey, while those of the genus Heptathela do not and instead rely on their built-in vibration sensors.[88] Spiders of the genus Heptathela have no venom glands, although they do have venom gland outlets on the fang tip.[89]
+The extinct families Arthrolycosidae, found in Carboniferous and Permian rocks, and Arthromygalidae, so far found only in Carboniferous rocks, have been classified as members of the Mesothelae.[90]
+The Mygalomorphae, which first appeared in the Triassic period,[79] are generally heavily built and ″hairy″, with large, robust chelicerae and fangs (technically, spiders do not have true hairs, but rather setae).[91][85] Well-known examples include tarantulas, ctenizid trapdoor spiders and the Australasian funnel-web spiders.[11] Most spend the majority of their time in burrows, and some run silk tripwires out from these, but a few build webs to capture prey. However, mygalomorphs cannot produce the pirifom silk that the Araneomorphae use as an instant adhesive to glue silk to surfaces or to other strands of silk, and this makes web construction more difficult for mygalomorphs. Since mygalomorphs rarely "balloon" by using air currents for transport, their populations often form clumps.[85] In addition to arthropods, some mygalomorphs are known to prey on frogs, small mammals, lizards, snakes, snails, and small birds.[92][93]
+In addition to accounting for over 90% of spider species, the Araneomorphae, also known as the "true spiders", include orb-web spiders, the cursorial wolf spiders, and jumping spiders,[85] as well as the only known herbivorous spider, Bagheera kiplingi.[49] They are distinguished by having fangs that oppose each other and cross in a pinching action, in contrast to the Mygalomorphae, which have fangs that are nearly parallel in alignment.[94]
+Although spiders are widely feared, only a few species are dangerous to people.[96] Spiders will only bite humans in self-defense, and few produce worse effects than a mosquito bite or bee sting.[97] Most of those with medically serious bites, such as recluse spiders (genus Loxosceles) and widow spiders (genus Latrodectus), would rather flee and bite only when trapped, although this can easily arise by accident.[98][99] The defensive tactics of Australian funnel-web spiders (family Atracidae) include fang display. Their venom, although they rarely inject much, has resulted in 13 attributed human deaths over 50 years.[100] They have been deemed to be the world's most dangerous spiders on clinical and venom toxicity grounds,[96] though this claim has also been attributed to the Brazilian wandering spider (genus Phoneutria).[101]
+There were about 100 reliably reported deaths from spider bites in the 20th century,[102] compared to about 1,500 from jellyfish stings.[103] Many alleged cases of spider bites may represent incorrect diagnoses,[104] which would make it more difficult to check the effectiveness of treatments for genuine bites.[105] A review published in 2016 agreed with this conclusion, showing that 78% of 134 published medical case studies of supposed spider bites did not meet the necessary criteria for a spider bite to be verified. In the case of the two genera with the highest reported number of bites, Loxosceles and Latrodectus, spider bites were not verified in over 90% of the reports. Even when verification had occurred, details of the treatment and its effects were often lacking.[106]
+Spider venoms may be a less polluting alternative to conventional pesticides, as they are deadly to insects but the great majority are harmless to vertebrates. Australian funnel web spiders are a promising source, as most of the world's insect pests have had no opportunity to develop any immunity to their venom, and funnel web spiders thrive in captivity and are easy to "milk". It may be possible to target specific pests by engineering genes for the production of spider toxins into viruses that infect species such as cotton bollworms.[107]
+The Ch'ol Maya use a beverage created from the tarantula species Brachypelma vagans for the treatment of a condition they term 'tarantula wind', the symptoms of which include chest pain, asthma and coughing.[108]
+Possible medical uses for spider venoms are being investigated, for the treatment of cardiac arrhythmia,[109] Alzheimer's disease,[110] strokes,[111] and erectile dysfunction.[112]
+The peptide GsMtx-4, found in the venom of Brachypelma vagans, is being researched to determine whether or not it could effectively be used for the treatment of cardiac arrhythmia, muscular dystrophy or glioma.[108]
+Because spider silk is both light and very strong, attempts are being made to produce it in goats' milk and in the leaves of plants, by means of genetic engineering.[113][114]
+Spiders can also be used as food. Cooked tarantulas are considered a delicacy in Cambodia,[115] and by the Piaroa Indians of southern Venezuela – provided the highly irritant bristles, the spiders' main defense system, are removed first.[116]
+Arachnophobia is a specific phobia—it is the abnormal fear of spiders or anything reminiscent of spiders, such as webs or spiderlike shapes. It is one of the most common specific phobias,[117][118] and some statistics show that 50% of women and 10% of men show symptoms.[119] It may be an exaggerated form of an instinctive response that helped early humans to survive,[120] or a cultural phenomenon that is most common in predominantly European societies.[121]
+Spiders have been the focus of stories and mythologies of various cultures for centuries.[122] Uttu, the ancient Sumerian goddess of weaving, was envisioned as a spider spinning her web.[123][124] According to her main myth, she resisted her father Enki's sexual advances by ensconcing herself in her web,[124] but let him in after he promised her fresh produce as a marriage gift,[124] thereby allowing him to intoxicate her with beer and rape her.[124] Enki's wife Ninhursag heard Uttu's screams and rescued her,[124] removing Enki's semen from her vagina and planting it in the ground to produce eight previously-nonexistent plants.[124] In a story told by the Roman poet Ovid in his Metamorphoses, Arachne was a Lydian girl who challenged the goddess Athena to a weaving contest.[125][126] Arachne won, but Athena destroyed her tapestry out of jealousy,[126][127] causing Arachne to hang herself.[126][127] In an act of mercy, Athena brought Arachne back to life as the first spider.[126][127] Stories about the trickster-spider Anansi are prominent in the folktales of West Africa and the Caribbean.[128]
+In some cultures, spiders have symbolized patience due to their hunting technique of setting webs and waiting for prey, as well as mischief and malice due to their venomous bites.[129] The Italian tarantella is a dance to rid the young woman of the lustful effects of a spider bite. Web-spinning also caused the association of the spider with creation myths, as they seem to have the ability to produce their own worlds.[130] Dreamcatchers are depictions of spiderwebs. The Moche people of ancient Peru worshipped nature.[131] They placed emphasis on animals and often depicted spiders in their art.[132]

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+Leonardo da Vinci (English: /ˌliːəˈnɑːrdoʊ də ˈvɪntʃi, ˌliːoʊˈ-, ˌleɪoʊˈ-/;[a] 14/15 April 1452[b] – 2 May 1519),[4] was an Italian polymath of the High Renaissance who is widely considered one of the greatest painters of all time (despite less than 25 of his paintings having survived).[c] The Mona Lisa is the most famous of Leonardo's works and the most famous portrait ever made.[5] The Last Supper is the most reproduced religious painting of all time[6] and his Vitruvian Man drawing is also regarded as a cultural icon.[7] He is also known for his his notebooks, in which he made drawings and notes on science and invention; these involve a variety of subjects including anatomy, cartography, painting, and paleontology. Leonardo's collective works compose a contribution to later generations of artists rivalled only by that of his contemporary Michelangelo.[8]
+Properly named Leonardo di ser Piero da Vinci,[d] Leonardo was born out of wedlock to a notary, Piero da Vinci, and a peasant woman, Caterina, in Vinci, in the region of Florence, Italy. Leonardo was educated in the studio of the renowned Italian painter Andrea del Verrocchio. Much of his earlier working life was spent in the service of Ludovico il Moro in Milan, and he later worked in Rome, Bologna and Venice. He spent his last three years in France, where he died in 1519.
+Although he had no formal academic training,[9] many historians and scholars regard Leonardo as the prime exemplar of the "Universal Genius" or "Renaissance Man", an individual of "unquenchable curiosity" and "feverishly inventive imagination."[6] He is widely considered one of the most diversely talented individuals ever to have lived.[10] According to art historian Helen Gardner, the scope and depth of his interests were without precedent in recorded history, and "his mind and personality seem to us superhuman, while the man himself mysterious and remote."[6] Scholars interpret his view of the world as being based in logic, though the empirical methods he used were unorthodox for his time.[11]
+Leonardo is revered for his technological ingenuity. He conceptualized flying machines, a type of armoured fighting vehicle, concentrated solar power, an adding machine,[12] and the double hull. Relatively few of his designs were constructed or even feasible during his lifetime, as the modern scientific approaches to metallurgy and engineering were only in their infancy during the Renaissance. Some of his smaller inventions, however, entered the world of manufacturing unheralded, such as an automated bobbin winder and a machine for testing the tensile strength of wire. He is also sometimes credited with the inventions of the parachute, helicopter, and tank.[13][14] He made substantial discoveries in anatomy, civil engineering, geology, optics, and hydrodynamics, but he did not publish his findings and they had little to no direct influence on subsequent science.[15]
+Leonardo was born on 14/15 April 1452[b] in the Tuscan hill town of Vinci, in the lower valley of the Arno river in the territory of the Medici-ruled Republic of Florence.[18] He was the out-of-wedlock son of Messer Piero Fruosino di Antonio da Vinci, a wealthy Florentine legal notary, and a peasant named Caterina,[e] identified as Caterina Buti del Vacca and more recently as Caterina di Meo Lippi by historian Martin Kemp. There have been many theories regarding Leonardo's mother's identity, including that she was a slave of foreign origin or an impoverished local youth.[17][20][21][22][f] Leonardo had no surname in the modern sense—da Vinci simply meaning "of Vinci"; his full birth name was Lionardo di ser Piero da Vinci,[2][25] meaning "Leonardo, (son) of ser Piero (from) Vinci."[18][d]
+Leonardo spent his first years in the hamlet of Anchiano in the home of his mother, and from at least 1457 lived in the household of his father, grandparents and uncle in the small town of Vinci.[26] His father had married a 16-year-old girl named Albiera Amadori, who loved Leonardo but died young[27] in 1465 without children. In 1468, when Leonardo was 16, his father married again to 20-year-old Francesca Lanfredini, who also died without children. Piero's legitimate heirs were born from his third wife Margherita di Guglielmo, who gave birth to six children, and his fourth and final wife, Lucrezia Cortigiani, who bore him another six heirs.[28][29]
+In all, Leonardo had 12 half-siblings, who were much younger than he was (the last was born when Leonardo was 40 years old) and with whom he had very little contact.[g]
+Leonardo received an informal education in Latin, geometry and mathematics. In later life, Leonardo recorded few distinct childhood incidents. One was of a kite coming to his cradle and opening his mouth with its tail; he regarded this as an omen of his writing on the subject.[31][32] The second occurred while he was exploring in the mountains: he discovered a cave and was both terrified that some great monster might lurk there and driven by curiosity to find out what was inside.[27] He also seems to have remembered some of his childhood observations of water, writing and crossing out the name of his hometown in one of his notebooks on the formation of rivers.[26]
+Leonardo's early life has been the subject of historical conjecture.[33] Vasari, the 16th-century biographer of Renaissance painters, tells a story of Leonardo as a very young man: A local peasant made himself a round shield and requested that Ser Piero have it painted for him. Leonardo, inspired by the story of Medusa, responded with a painting of a monster spitting fire that was so terrifying that his father bought a different shield to give to the peasant and sold Leonardo's to a Florentine art dealer for 100 ducats, who in turn sold it to the Duke of Milan.[34]
+In the mid-1460s, Leonardo's family moved to Florence, and around the age of 14,[30] he became a garzone (studio boy) in the workshop of Verrocchio, who was the leading Florentine painter and sculptor of his time.[35] Leonardo became an apprentice by the age of 17 and remained in training for seven years.[36][37] Other famous painters apprenticed in the workshop or associated with it include Ghirlandaio, Perugino, Botticelli, and Lorenzo di Credi.[27][38] Leonardo was exposed to both theoretical training and a wide range of technical skills,[39] including drafting, chemistry, metallurgy, metal working, plaster casting, leather working, mechanics, and wood-work, as well as the artistic skills of drawing, painting, sculpting, and modelling.[40][h]
+Much of the painting in Verrocchio's workshop was done by his employees. According to Vasari, Leonardo collaborated with Verrocchio on his The Baptism of Christ, painting the young angel holding Jesus' robe in a manner that was so far superior to his master's that Verrocchio put down his brush and never painted again, although this is believed to be an apocryphal story.[41] Close examination reveals areas of the work that have been painted or touched-up over the tempera, using the new technique of oil paint, including the landscape, the rocks seen through the brown mountain stream, and much of the figure of Jesus, bearing witness to the hand of Leonardo.[42] Leonardo may have been the model for two works by Verrocchio: the bronze statue of David in the Bargello, and the Archangel Raphael in Tobias and the Angel.[20]
+By 1472, at the age of 20, Leonardo qualified as a master in the Guild of Saint Luke, the guild of artists and doctors of medicine,[i] but even after his father set him up in his own workshop, his attachment to Verrocchio was such that he continued to collaborate and live with him.[27][43] Leonardo's earliest known dated work is a 1473 pen-and-ink drawing of the Arno valley,[38] which has been cited as the first "pure" landscape in the Occident.[j][44] According to Vasari, the young Leonardo was the first to suggest making the Arno river a navigable channel between Florence and Pisa.[45]
+In January 1478, Leonardo received an independent commission to paint an altarpiece for the Chapel of St. Bernard in the Palazzo Vecchio,[46] an indication of his independence from Verrocchio's studio. One anonymous writer claims that in 1480, Leonardo was living with the Medici and often worked in the garden of the Piazza San Marco, Florence, where a Neoplatonic academy of artists, poets and philosophers organized by the Medici met.[20] In March 1481, he received a commission from the monks of San Donato in Scopeto for The Adoration of the Magi.[47]
+Neither of these initial commissions were completed, being abandoned when Leonardo went to offer his services to Duke of Milan Ludovico Sforza. In 1482, he casted a silver stringed instrument[k] from a horse's skull and ram horns to bring to Sforza,[48] whom he wrote a letter describing the diverse things that he could achieve in the fields of engineering and weapon design, and mentioning that he could paint.[38][48][49]
+Leonardo worked in Milan from 1482 until 1499. He was commissioned to paint the Virgin of the Rocks for the Confraternity of the Immaculate Conception and The Last Supper for the monastery of Santa Maria delle Grazie.[51] In the spring of 1485, Leonardo travelled to Hungary on behalf of Sforza to meet king Matthias Corvinus, and was commissioned by him to paint a Madonna.[52]
+Leonardo was employed on many other projects for Sforza, including the preparation of floats and pageants for special occasions, a drawing and wooden model for a competition to design the cupola for Milan Cathedral (which he withdrew),[53] and a model for a huge equestrian monument to Ludovico's predecessor Francesco Sforza. This would have surpassed in size the only two large equestrian statues of the Renaissance, Donatello's Gattamelata in Padua and Verrocchio's Bartolomeo Colleoni in Venice, and became known as the Gran Cavallo.[38] Leonardo completed a model for the horse and made detailed plans for its casting,[38] but in November 1494, Ludovico gave the bronze to his brother-in-law to be used for a cannon to defend the city from Charles VIII.[38]
+With Ludovico Sforza overthrown at the dawn of the Second Italian War, Leonardo, with his assistant Salaì and friend, the mathematician Luca Pacioli, fled Milan for Venice.[54] There, he was employed as a military architect and engineer, devising methods to defend the city from naval attack.[27] On his return to Florence in 1500, he and his household were guests of the Servite monks at the monastery of Santissima Annunziata and were provided with a workshop where, according to Vasari, Leonardo created the cartoon of The Virgin and Child with St Anne and St John the Baptist, a work that won such admiration that "men and women, young and old" flocked to see it "as if they were attending a great festival."[55][l]
+In Cesena in 1502, Leonardo entered the service of Cesare Borgia, the son of Pope Alexander VI, acting as a military architect and engineer and travelling throughout Italy with his patron.[54] Leonardo created a map of Cesare Borgia's stronghold, a town plan of Imola in order to win his patronage. Maps were extremely rare at the time and it would have seemed like a new concept. Upon seeing it, Cesare hired Leonardo as his chief military engineer and architect. Later in the year, Leonardo produced another map for his patron, one of Chiana Valley, Tuscany, so as to give his patron a better overlay of the land and greater strategic position. He created this map in conjunction with his other project of constructing a dam from the sea to Florence, in order to allow a supply of water to sustain the canal during all seasons.
+Leonardo had left Borgia's service and returned to Florence by early 1503,[57] where he rejoined the Guild of Saint Luke on 18 October of that year. By this same month, Leonardo had begun working on a portrait of Lisa del Giocondo, the model for the Mona Lisa,[58][59] which he would continue working on until his twilight years. In January 1504, he was part of a committee formed to recommend where Michelangelo's statue of David should be placed.[60] He then spent two years in Florence designing and painting a mural of The Battle of Anghiari for the Signoria,[54] with Michelangelo designing its companion piece, The Battle of Cascina.[m]
+In 1506, Leonardo was summoned to Milan by Charles II d'Amboise, the acting French governor of the city. The Council of Florence wished Leonardo to return promptly to finish The Battle of Anghiari, but he was given leave at the behest of Louis XII, who considered commissioning the artist to make some portraits.[63] Leonardo may have commenced a project for an equestrian figure of d'Amboise;[64] a wax model survives and, if genuine, is the only extant example of Leonardo's sculpture. Leonardo was otherwise free to pursue his scientific interests.[63] Many of Leonardo's most prominent pupils either knew or worked with him in Milan,[27] including Bernardino Luini, Giovanni Antonio Boltraffio, and Marco d'Oggiono.
+In 1507, Leonardo was in Florence sorting out a dispute with his brothers over the estate of his father, who had died in 1504. By 1508, Leonardo was back in Milan, living in his own house in Porta Orientale in the parish of Santa Babila.[65]
+In 1512, Leonardo was working on plans for an equestrian monument for Gian Giacomo Trivulzio, but this was prevented by an invasion of a confederation of Swiss, Spanish and Venetian forces, which drove the French from Milan. Leonardo stayed in the city, spending several months in 1513 at the Medici's Vaprio d'Adda villa.[67] In March of that year, Lorenzo de' Medici's son Giovanni assumed the papacy (as Leo X); Leonardo went to Rome that September, where he was received by the pope's brother Giuliano.[67] From September 1513 to 1516, Leonardo spent much of his time living in the Belvedere Courtyard in the Apostolic Palace, where Michelangelo and Raphael were both active.[65] Leonardo was given an allowance of 33 ducats a month, and according to Vasari, decorated a lizard with scales dipped in quicksilver.[68] The pope gave him a painting commission of unknown subject matter, but cancelled it when the artist set about developing a new kind of varnish.[68][n] Leonardo became ill, in what may have been the first of multiple strokes leading to his death.[68] He practiced botany in the Gardens of Vatican City, and was commissioned to make plans for the pope's proposed draining of the Pontine Marshes.[69] He also dissected cadavers, making notes for a treatise on vocal cords;[70] these he gave to an official in hopes of regaining the pope's favor, but was unsuccessful.[68]
+In October 1515, King Francis I of France recaptured Milan.[47] Leonardo was present at the 19 December meeting of Francis I and Leo X, which took place in Bologna.[27][71][72] In 1516, Leonardo entered Francis' service, being given the use of the manor house Clos Lucé, near the king's residence at the royal Château d'Amboise. Being frequently visited by Francis, he drew plans for an immense castle town the king intended to erect at Romorantin, and made a mechanical lion, which during a pageant walked toward the king and—upon being struck by a wand—opened its chest to reveal a cluster of lilies.[73][74][o] Leonardo was accompanied during this time by his friend and apprentice, Francesco Melzi, and supported by a pension totalling 10,000 scudi.[65] At some point, Melzi drew a portrait of Leonardo; the only others known from his lifetime were a sketch by an unknown assistant on the back of one of Leonardo's studies (c. 1517)[76] and a drawing by Giovanni Ambrogio Figino depicting an elderly Leonardo with his right arm assuaged by cloth.[77][p][q] The latter, in addition to the record of an October 1517 visit by Louis d'Aragon,[r] confirms an account of Leonardo's right hand being paralytic at the age of 65,[80] which may indicate why he left works such as the Mona Lisa unfinished.[78][81][82] He continued to work at some capacity until eventually becoming ill and bedridden for several months.[80]
+Leonardo died at Clos Lucé on 2 May 1519 at the age of 67, possibly of a stroke.[83][82] Francis I had become a close friend. Vasari describes Leonardo as lamenting on his deathbed, full of repentance, that "he had offended against God and men by failing to practice his art as he should have done."[84] Vasari states that in his last days, Leonardo sent for a priest to make his confession and to receive the Holy Sacrament.[85] Vasari also records that the king held Leonardo's head in his arms as he died, although this story may be legend rather than fact.[s][t] In accordance with his will, sixty beggars carrying tapers followed Leonardo's casket.[87][u] Melzi was the principal heir and executor, receiving, as well as money, Leonardo's paintings, tools, library and personal effects. Leonardo also remembered his other long-time pupil and companion, Salaì, and his servant Baptista de Vilanis, who each received half of Leonardo's vineyards.[88] His brothers received land, and his serving woman received a fur-lined cloak. On 12 August 1519, Leonardo's remains were interred in the Collegiate Church of Saint Florentin at the Château d'Amboise.[89]
+Some 20 years after Leonardo's death, Francis was reported by the goldsmith and sculptor Benvenuto Cellini as saying: "There had never been another man born in the world who knew as much as Leonardo, not so much about painting, sculpture and architecture, as that he was a very great philosopher."[90]
+Florence at the time of Leonardo's youth was the centre of Christian Humanist thought and culture.[35] Leonardo commenced his apprenticeship with Verrocchio in 1466, the year that Verrocchio's master, the great sculptor Donatello, died.[v] The painter Uccello, whose early experiments with perspective were to influence the development of landscape painting, was a very old man. The painters Piero della Francesca and Filippo Lippi, sculptor Luca della Robbia, and architect and writer Leon Battista Alberti were in their sixties. The successful artists of the next generation were Leonardo's teacher Verrocchio, Antonio del Pollaiuolo, and the portrait sculptor Mino da Fiesole.[91][92][93][94]
+Leonardo's youth was spent in a Florence that was ornamented by the works of these artists and by Donatello's contemporaries, Masaccio, whose figurative frescoes were imbued with realism and emotion; and Ghiberti, whose Gates of Paradise, gleaming with gold leaf, displayed the art of combining complex figure compositions with detailed architectural backgrounds. Piero della Francesca had made a detailed study of perspective,[95] and was the first painter to make a scientific study of light. These studies and Alberti's treatise De pictura[96] were to have a profound effect on younger artists and in particular on Leonardo's own observations and artworks.[91][93][94]
+A prevalent tradition in Florence was the small altarpiece of the Virgin and Child. Many of these were created in tempera or glazed terracotta by the workshops of Filippo Lippi, Verrocchio and the prolific della Robbia family.[91] Leonardo's early Madonnas such as The Madonna with a carnation and the Benois Madonna followed this tradition while showing idiosyncratic departures, particularly in the latter in which the Virgin is set at an oblique angle to the picture space with the Christ Child at the opposite angle. This compositional theme was to emerge in Leonardo's later paintings such as The Virgin and Child with St. Anne.[27]
+Leonardo was a contemporary of Botticelli, Domenico Ghirlandaio and Perugino, who were all slightly older than he was.[92] He would have met them at the workshop of Verrocchio, with whom they had associations, and at the Academy of the Medici.[27] Botticelli was a particular favourite of the Medici family, and thus his success as a painter was assured. Ghirlandaio and Perugino were both prolific and ran large workshops. They competently delivered commissions to well-satisfied patrons who appreciated Ghirlandaio's ability to portray the wealthy citizens of Florence within large religious frescoes, and Perugino's ability to deliver a multitude of saints and angels of unfailing sweetness and innocence.[91]
+These three were among those commissioned to paint the walls of the Sistine Chapel, the work commencing with Perugino's employment in 1479. Leonardo was not part of this prestigious commission. His first significant commission, The Adoration of the Magi for the Monks of Scopeto, was never completed.[27]
+In 1476, during the time of Leonardo's association with Verrocchio's workshop, the Portinari Altarpiece by Hugo van der Goes arrived in Florence, bringing from Northern Europe new painterly techniques that were to profoundly affect Leonardo, Ghirlandaio, Perugino and others.[92] In 1479, the Sicilian painter Antonello da Messina, who worked exclusively in oils, travelled north on his way to Venice, where the leading painter Giovanni Bellini adopted the technique of oil painting, quickly making it the preferred method in Venice. Leonardo was also later to visit Venice.[92][94]
+Like the two contemporary architects Donato Bramante (who designed the Belvedere Courtyard) and Antonio da Sangallo the Elder, Leonardo experimented with designs for centrally planned churches, a number of which appear in his journals, as both plans and views, although none was ever realised.[92][97]
+Leonardo's political contemporaries were Lorenzo de' Medici (il Magnifico), who was three years older, and his younger brother Giuliano, who was slain in the Pazzi conspiracy in 1478. Leonardo was sent as an ambassador by the Medici court to Ludovico il Moro, who ruled Milan between 1479 and 1499.[92]
+With Alberti, Leonardo visited the home of the Medici and through them came to know the older Humanist philosophers of whom Marsiglio Ficino, proponent of Neo Platonism; Cristoforo Landino, writer of commentaries on Classical writings, and John Argyropoulos, teacher of Greek and translator of Aristotle were the foremost. Also associated with the Academy of the Medici was Leonardo's contemporary, the brilliant young poet and philosopher Pico della Mirandola.[92][94][87] Leonardo later wrote in the margin of a journal, "The Medici made me and the Medici destroyed me." While it was through the action of Lorenzo that Leonardo received his employment at the court of Milan, it is not known exactly what Leonardo meant by this cryptic comment.[27]
+Although usually named together as the three giants of the High Renaissance, Leonardo, Michelangelo and Raphael were not of the same generation. Leonardo was 23 when Michelangelo was born and 31 when Raphael was born.[92] Raphael died at the age of 37 in 1520, the year after Leonardo died, but Michelangelo went on creating for another 45 years.[93][94]
+Within Leonardo's lifetime, his extraordinary powers of invention, his "outstanding physical beauty," "infinite grace," "great strength and generosity," "regal spirit and tremendous breadth of mind," as described by Vasari,[98] as well as all other aspects of his life, attracted the curiosity of others. One such aspect was his love for animals, likely including vegetarianism and according to Vasari, a habit of purchasing caged birds and releasing them.[99][100]
+Leonardo had many friends who are now renowned either in their fields or for their historical significance. They included the mathematician Luca Pacioli,[101] with whom he collaborated on the book Divina proportione in the 1490s. Leonardo appears to have had no close relationships with women except for his friendship with Cecilia Gallerani and the two Este sisters, Beatrice and Isabella.[102] While on a journey that took him through Mantua, he drew a portrait of Isabella that appears to have been used to create a painted portrait, now lost.[27]
+Beyond friendship, Leonardo kept his private life secret. His sexuality has been the subject of satire, analysis, and speculation. This trend began in the mid-16th century and was revived in the 19th and 20th centuries, most notably by Sigmund Freud.[103] Leonardo's most intimate relationships were perhaps with his pupils Salaì and Melzi. Melzi, writing to inform Leonardo's brothers of his death, described Leonardo's feelings for his pupils as both loving and passionate. It has been claimed since the 16th century that these relationships were of a sexual or erotic nature. Court records of 1476, when he was aged twenty-four, show that Leonardo and three other young men were charged with sodomy in an incident involving a well-known male prostitute. The charges were dismissed for lack of evidence, and there is speculation that since one of the accused, Lionardo de Tornabuoni, was related to Lorenzo de' Medici, the family exerted its influence to secure the dismissal.[104] Since that date much has been written about his presumed homosexuality and its role in his art, particularly in the androgyny and eroticism manifested in Saint John the Baptist and Bacchus and more explicitly in a number of erotic drawings.[105]
+Gian Giacomo Caprotti da Oreno, nicknamed Salaì or Il Salaino ("The Little Unclean One," i.e., the devil), entered Leonardo's household in 1490. After only a year, Leonardo made a list of his misdemeanours, calling him "a thief, a liar, stubborn, and a glutton," after he had made off with money and valuables on at least five occasions and spent a fortune on clothes.[106] Nevertheless, Leonardo treated him with great indulgence, and he remained in Leonardo's household for the next thirty years.[107] Salaì executed a number of paintings under the name of Andrea Salaì, but although Vasari claims that Leonardo "taught him a great deal about painting,"[74] his work is generally considered to be of less artistic merit than others among Leonardo's pupils, such as Marco d'Oggiono and Boltraffio. In 1515, he painted a nude version of the Mona Lisa, known as Monna Vanna.[108] Salaì owned the Mona Lisa at the time of his death in 1524, and in his will it was assessed at 505 lire, an exceptionally high valuation for a small panel portrait.[109]
+In 1506, Leonardo took on another pupil, Count Francesco Melzi, the son of a Lombard aristocrat, who is considered to have been his favourite student. He travelled to France with Leonardo and remained with him until Leonardo's death.[27] Melzi inherited the artistic and scientific works, manuscripts, and collections of Leonardo and administered the estate.
+Despite the recent awareness and admiration of Leonardo as a scientist and inventor, for the better part of four hundred years his fame rested on his achievements as a painter. A handful of works that are either authenticated or attributed to him have been regarded as among the great masterpieces. These paintings are famous for a variety of qualities that have been much imitated by students and discussed at great length by connoisseurs and critics. By the 1490s Leonardo had already been described as a "Divine" painter.[110]
+Among the qualities that make Leonardo's work unique are his innovative techniques for laying on the paint; his detailed knowledge of anatomy, light, botany and geology; his interest in physiognomy and the way humans register emotion in expression and gesture; his innovative use of the human form in figurative composition; and his use of subtle gradation of tone. All these qualities come together in his most famous painted works, the Mona Lisa, the Last Supper, and the Virgin of the Rocks.[111]
+Leonardo first gained attention for his work on the Baptism of Christ, painted in conjunction with Verrocchio. Two other paintings appear to date from his time at Verrocchio's workshop, both of which are Annunciations. One is small, 59 centimetres (23 in) long and 14 centimetres (5.5 in) high. It is a "predella" to go at the base of a larger composition, a painting by Lorenzo di Credi from which it has become separated. The other is a much larger work, 217 centimetres (85 in) long.[112] In both Annunciations, Leonardo used a formal arrangement, like two well-known pictures by Fra Angelico of the same subject, of the Virgin Mary sitting or kneeling to the right of the picture, approached from the left by an angel in profile, with a rich flowing garment, raised wings and bearing a lily. Although previously attributed to Ghirlandaio, the larger work is now generally attributed to Leonardo.[113]
+In the smaller painting, Mary averts her eyes and folds her hands in a gesture that symbolised submission to God's will. Mary is not submissive, however, in the larger piece. The girl, interrupted in her reading by this unexpected messenger, puts a finger in her bible to mark the place and raises her hand in a formal gesture of greeting or surprise.[91] This calm young woman appears to accept her role as the Mother of God, not with resignation but with confidence. In this painting, the young Leonardo presents the humanist face of the Virgin Mary, recognising humanity's role in God's incarnation.
+In the 1480s, Leonardo received two very important commissions and commenced another work that was of ground-breaking importance in terms of composition. Two of the three were never finished, and the third took so long that it was subject to lengthy negotiations over completion and payment.
+One of these paintings was Saint Jerome in the Wilderness, which Bortolon associates with a difficult period of Leonardo's life, as evidenced in his diary: "I thought I was learning to live; I was only learning to die."[27] Although the painting is barely begun, the composition can be seen and is very unusual.[w] Jerome, as a penitent, occupies the middle of the picture, set on a slight diagonal and viewed somewhat from above. His kneeling form takes on a trapezoid shape, with one arm stretched to the outer edge of the painting and his gaze looking in the opposite direction. J. Wasserman points out the link between this painting and Leonardo's anatomical studies.[114] Across the foreground sprawls his symbol, a great lion whose body and tail make a double spiral across the base of the picture space. The other remarkable feature is the sketchy landscape of craggy rocks against which the figure is silhouetted.
+The daring display of figure composition, the landscape elements and personal drama also appear in the great unfinished masterpiece, the Adoration of the Magi, a commission from the Monks of San Donato a Scopeto. It is a complex composition, of about 250 x 250 centimetres. Leonardo did numerous drawings and preparatory studies, including a detailed one in linear perspective of the ruined classical architecture that forms part of the background. In 1482 Leonardo went to Milan at the behest of Lorenzo de' Medici in order to win favour with Ludovico il Moro, and the painting was abandoned.[20][113]
+The third important work of this period is the Virgin of the Rocks, commissioned in Milan for the Confraternity of the Immaculate Conception. The painting, to be done with the assistance of the de Predis brothers, was to fill a large complex altarpiece.[115] Leonardo chose to paint an apocryphal moment of the infancy of Christ when the infant John the Baptist, in protection of an angel, met the Holy Family on the road to Egypt. The painting demonstrates an eerie beauty as the graceful figures kneel in adoration around the infant Christ in a wild landscape of tumbling rock and whirling water.[116] While the painting is quite large, about 200×120 centimetres, it is not nearly as complex as the painting ordered by the monks of St Donato, having only four figures rather than about fifty and a rocky landscape rather than architectural details. The painting was eventually finished; in fact, two versions of the painting were finished: one remained at the chapel of the Confraternity, while Leonardo took the other to France. The Brothers did not get their painting, however, nor the de Predis their payment, until the next century.[38][54]
+Leonardo's most remarkable portrait of this period is the Lady with an Ermine, presumed to be  Cecilia Gallerani (c. 1483–1490), lover of Ludovico Sforza.[117][118] The painting is charcterised by the pose of the figure with the head turned at a very different angle to the torso, unusual at a date when many portraits were still rigidly in profile. The ermine plainly carries symbolic meaning, relating either to the sitter, or to Ludovico who belonged to the prestigious Order of the Ermine.[117]
+Leonardo's most famous painting of the 1490s is The Last Supper, commissioned for the refectory of the Convent of Santa Maria della Grazie in Milan. It represents the last meal shared by Jesus with his disciples before his capture and death, and shows the moment when Jesus has just said "one of you will betray me", and the consternation that this statement caused.[38]
+The novelist Matteo Bandello observed Leonardo at work and wrote that some days he would paint from dawn till dusk without stopping to eat and then not paint for three or four days at a time.[119] This was beyond the comprehension of the prior of the convent, who hounded him until Leonardo asked Ludovico to intervene. Vasari describes how Leonardo, troubled over his ability to adequately depict the faces of Christ and the traitor Judas, told the Duke that he might be obliged to use the prior as his model.[120]
+When finished, the painting was acclaimed as a masterpiece of design and characterization,[121] but it deteriorated rapidly, so that within a hundred years it was described by one viewer as "completely ruined."[122] Leonardo, instead of using the reliable technique of fresco, had used tempera over a ground that was mainly gesso, resulting in a surface subject to mould and to flaking.[123] Despite this, the painting remains one of the most reproduced works of art; countless copies have been made in various mediums.
+It is recorded that in 1492, Leonardo, with assistants painted the Sala delle Asse in the Sforza Castle in Milan,  with a trompe-l'œil depicting trees, with an intricate labyrinth of leaves and knots on the ceiling.[124]
+In 1505 Leonardo was commissioned to paint The Battle of Anghiari in the Salone dei Cinquecento (Hall of the Five Hundred) in the Palazzo Vecchio, Florence. Leonardo devised a dynamic composition depicting four men riding raging war horses engaged in a battle for possession of a standard, at the Battle of Anghiari in 1440.  Michelangelo was assigned the opposite wall to depict the Battle of Cascina. Leonardo's painting deteriorated rapidly and is now known from a copy by Rubens.[125]
+Among the works created by Leonardo in the 16th century is the small portrait known as the Mona Lisa or La Gioconda, the laughing one. In the present era, it is arguably the most famous painting in the world. Its fame rests, in particular, on the elusive smile on the woman's face, its mysterious quality perhaps due to the subtly shadowed corners of the mouth and eyes such that the exact nature of the smile cannot be determined. The shadowy quality for which the work is renowned came to be called "sfumato," or Leonardo's smoke. Vasari, who is generally thought to have known the painting only by repute, said that "the smile was so pleasing that it seemed divine rather than human; and those who saw it were amazed to find that it was as alive as the original."[126][x]
+Other characteristics of the painting are the unadorned dress, in which the eyes and hands have no competition from other details; the dramatic landscape background, in which the world seems to be in a state of flux; the subdued colouring; and the extremely smooth nature of the painterly technique, employing oils laid on much like tempera, and blended on the surface so that the brushstrokes are indistinguishable.[y] Vasari expressed the opinion that the manner of painting would make even "the most confident master...despair and lose heart."[129] The perfect state of preservation and the fact that there is no sign of repair or overpainting is rare in a panel painting of this date.[130]
+In the painting Virgin and Child with St. Anne, the composition again picks up the theme of figures in a landscape, which Wasserman describes as "breathtakingly beautiful"[131] and harkens back to the St Jerome picture with the figure set at an oblique angle. What makes this painting unusual is that there are two obliquely set figures superimposed. Mary is seated on the knee of her mother, St Anne. She leans forward to restrain the Christ Child as he plays roughly with a lamb, the sign of his own impending sacrifice.[38] This painting, which was copied many times, influenced Michelangelo, Raphael, and Andrea del Sarto,[132] and through them Pontormo and Correggio. The trends in composition were adopted in particular by the Venetian painters Tintoretto and Veronese.
+Leonardo was not a prolific painter, but he was a most prolific draftsman, keeping journals full of small sketches and detailed drawings recording all manner of things that took his attention. As well as the journals there exist many studies for paintings, some of which can be identified as preparatory to particular works such as The Adoration of the Magi, The Virgin of the Rocks and The Last Supper.[133]
+His earliest dated drawing is a Landscape of the Arno Valley, 1473, which shows the river, the mountains, Montelupo Castle and the farmlands beyond it in great detail.[27][133][z] According to art historian Ludwig Heydenreich, this is "The first true landscape in art."[134] Massimo Polidoro says that it was the first landscape "not to be the background of some religious scene or a portrait. It is the first [documented] time where a landscape was drawn just for the sake of it."[44]
+Among his famous drawings are the Vitruvian Man, a study of the proportions of the human body; the Head of an Angel, for The Virgin of the Rocks in the Louvre; a botanical study of Star of Bethlehem; and a large drawing (160×100 cm) in black chalk on coloured paper of The Virgin and Child with St. Anne and St. John the Baptist in the National Gallery, London.[133] This drawing employs the subtle sfumato technique of shading, in the manner of the Mona Lisa. It is thought that Leonardo never made a painting from it, the closest similarity being to The Virgin and Child with St. Anne in the Louvre.[135]
+Other drawings of interest include numerous studies generally referred to as "caricatures" because, although exaggerated, they appear to be based upon observation of live models. Vasari relates that if Leonardo saw a person with an interesting face he would follow them around all day observing them.[136] There are numerous studies of beautiful young men, often associated with Salaì, with the rare and much admired facial feature, the so-called "Grecian profile."[aa] These faces are often contrasted with that of a warrior.[133] Salaì is often depicted in fancy-dress costume. Leonardo is known to have designed sets for pageants with which these may be associated. Other, often meticulous, drawings show studies of drapery. A marked development in Leonardo's ability to draw drapery occurred in his early works. Another often-reproduced drawing is a macabre sketch that was done by Leonardo in Florence in 1479 showing the body of Bernardo Baroncelli, hanged in connection with the murder of Giuliano, brother of Lorenzo de' Medici, in the Pazzi conspiracy.[133] In his notes, Leonardo recorded the colours of the robes that Baroncelli was wearing when he died.
+Renaissance humanism recognised no mutually exclusive polarities between the sciences and the arts, and Leonardo's studies in science and engineering are sometimes considered as impressive and innovative as his artistic work.[38] These studies were recorded in 13,000 pages of notes and drawings, which fuse art and natural philosophy (the forerunner of modern science). They were made and maintained daily throughout Leonardo's life and travels, as he made continual observations of the world around him.[38]
+Leonardo's notes and drawings display an enormous range of interests and preoccupations, some as mundane as lists of groceries and people who owed him money and some as intriguing as designs for wings and shoes for walking on water. There are compositions for paintings, studies of details and drapery, studies of faces and emotions, of animals, babies, dissections, plant studies, rock formations, whirlpools, war machines, flying machines and architecture.[38]
+These notebooks—originally loose papers of different types and sizes, were largely entrusted to Leonardo's pupil and heir Francesco Melzi after the master's death.[137] These were to be published, a task of overwhelming difficulty because of its scope and Leonardo's idiosyncratic writing.[138] Some of Leonardo's drawings were copied by an anonymous Milanese artist for a planned treatise on art c. 1570.[139] After Melzi's death in 1570, the collection passed to his son, the lawyer Orazio, who initially took little interest in the journals.[137] In 1587, a Melzi household tutor named Lelio Gavardi took 13 of the manuscripts to Pisa; there, the architect Giovanni Magenta reproached Gavardi for having taken the manuscripts illicitly and returned them to Orazio. Having many more such works in his possession, Orazio gifted the volumes to Magenta. News spread of these lost works of Leonardo's, and Orazio retrieved seven of the 13 manuscripts, which he then gave to Pompeo Leoni for publication in two volumes; one of these was the Codex Atlanticus. The other six works had been distributed to a few others.[140] After Orazio's death, his heirs sold the rest of Leonardo's possessions, and thus began their dispersal.[141]
+Some works have found their way into major collections such as the Royal Library at Windsor Castle, the Louvre, the Biblioteca Nacional de España, the Victoria and Albert Museum, the Biblioteca Ambrosiana in Milan, which holds the 12-volume Codex Atlanticus, and the British Library in London, which has put a selection from the Codex Arundel (BL Arundel MS 263) online.[142] Works have also been at Holkham Hall, the Metropolitan Museum of Art, and in the private hands of John Nicholas Brown I and Robert Lehman.[137] The Codex Leicester is the only privately owned major scientific work of Leonardo; it is owned by Bill Gates and displayed once a year in different cities around the world.
+Most of Leonardo's writings are in mirror-image cursive.[143][44] Since Leonardo wrote with his left hand, it was probably easier for him to write from right to left.[144][ab] Leonardo used a variety of shorthand and symbols, and states in his notes that he intended to prepare them for publication.[143] In many cases a single topic is covered in detail in both words and pictures on a single sheet, together conveying information that would not be lost if the pages were published out of order.[147] Why they were not published during Leonardo's lifetime is unknown.[38]
+Leonardo's approach to science was observational: he tried to understand a phenomenon by describing and depicting it in utmost detail and did not emphasise experiments or theoretical explanation. Since he lacked formal education in Latin and mathematics, contemporary scholars mostly ignored Leonardo the scientist, although he did teach himself Latin. In the 1490s he studied mathematics under Luca Pacioli and prepared a series of drawings of regular solids in a skeletal form to be engraved as plates for Pacioli's book Divina proportione, published in 1509.[38] While living in Milan, he studied light from the summit of Monte Rosa.[63] Scientific writings in his notebook on fossils have been considered as influential on early palaeontology[148][149] and he has been called the father of ichnology.[150]
+The content of his journals suggest that he was planning a series of treatises on a variety of subjects. A coherent treatise on anatomy is said to have been observed during a visit by Cardinal Louis d'Aragon's secretary in 1517.[151] Aspects of his work on the studies of anatomy, light and the landscape were assembled for publication by Melzi and eventually published as A Treatise on Painting in France and Italy in 1651 and Germany in 1724,[152] with engravings based upon drawings by the Classical painter Nicolas Poussin.[153] According to Arasse, the treatise, which in France went into 62 editions in fifty years, caused Leonardo to be seen as "the precursor of French academic thought on art."[38]
+While Leonardo's experimentation followed scientific methods, a recent and exhaustive analysis of Leonardo as a scientist by Fritjof Capra argues that Leonardo was a fundamentally different kind of scientist from Galileo, Newton and other scientists who followed him in that, as a "Renaissance Man", his theorising and hypothesising integrated the arts and particularly painting.[154][page needed]
+Leonardo started his study in the anatomy of the human body under the apprenticeship of Andrea del Verrocchio, who demanded that his students develop a deep knowledge of the subject.[155] As an artist, he quickly became master of topographic anatomy, drawing many studies of muscles, tendons and other visible anatomical features.
+As a successful artist, Leonardo was given permission to dissect human corpses at the Hospital of Santa Maria Nuova in Florence and later at hospitals in Milan and Rome. From 1510 to 1511 he collaborated in his studies with the doctor Marcantonio della Torre. Leonardo made over 240 detailed drawings and wrote about 13,000 words towards a treatise on anatomy.[156] Only a small amount of the material on anatomy was published in Leonardo's Treatise on painting.[138] During the time that Melzi was ordering the material into chapters for publication, they were examined by a number of anatomists and artists, including Vasari, Cellini and Albrecht Dürer, who made a number of drawings from them.[138]
+Leonardo's anatomical drawings include many studies of the human skeleton and its parts, and of muscles and sinews. He studied the mechanical functions of the skeleton and the muscular forces that are applied to it in a manner that prefigured the modern science of biomechanics.[157] He drew the heart and vascular system, the sex organs and other internal organs, making one of the first scientific drawings of a fetus in utero.[133] The drawings and notation are far ahead of their time, and if published would undoubtedly have made a major contribution to medical science.[156]
+Leonardo also closely observed and recorded the effects of age and of human emotion on the physiology, studying in particular the effects of rage. He drew many figures who had significant facial deformities or signs of illness.[38][133] Leonardo also studied and drew the anatomy of many animals, dissecting cows, birds, monkeys, bears, and frogs, and comparing in his drawings their anatomical structure with that of humans. He also made a number of studies of horses.[133]
+Leonardo's dissections and documentation of muscles, nerves, and vessels helped to describe the physiology and mechanics of movement. He attempted to identify the source of 'emotions' and their expression. He found it difficult to incorporate the prevailing system and theories of bodily humours, but eventually he abandoned these physiological explanations of bodily functions. He made the observations that humours were not located in cerebral spaces or ventricles. He documented that the humours were not contained in the heart or the liver, and that it was the heart that defined the circulatory system. He was the first to define atherosclerosis and liver cirrhosis. He created models of the cerebral ventricles with the use of melted wax and constructed a glass aorta to observe the circulation of blood through the aortic valve by using water and grass seed to watch flow patterns. Vesalius published his work on anatomy and physiology in De humani corporis fabrica in 1543.[158]
+During his lifetime, Leonardo was also valued as an engineer. With the same rational and analytical approach that moved him to represent the human body and to investigate anatomy, Leonardo studied and designed a bewildering number of machines and devices. He drew their “anatomy” with unparalleled mastery, producing the first form of the modern technical drawing, including a perfected "exploded view" technique, to represent internal components. Those studies and projects collected in his codices fill more than 5,000 pages.[159] In a letter of 1482 to the lord of Milan Ludovico il Moro, he wrote that he could create all sorts of machines both for the protection of a city and for siege. When he fled from Milan to Venice in 1499, he found employment as an engineer and devised a system of moveable barricades to protect the city from attack. In 1502, he created a scheme for diverting the flow of the Arno river, a project on which Niccolò Machiavelli also worked.[160][161] He continued to contemplate the canalization of Lombardy's plains while in Louis XII's company[63] and of the Loire and its tributaries in the company of Francis I.[162] Leonardo's journals include a vast number of inventions, both practical and impractical. They include musical instruments, a mechanical knight, hydraulic pumps, reversible crank mechanisms, finned mortar shells, and a steam cannon.[27][38]
+Leonardo was fascinated by the phenomenon of flight for much of his life, producing many studies, including Codex on the Flight of Birds (c. 1505), as well as plans for several flying machines, such as a flapping ornithopter and a machine with a helical rotor.[38] A 2003 documentary by British television station Channel Four, titled Leonardo's Dream Machines, various designs by Leonardo, such as a parachute and a giant crossbow, were interpreted and constructed.[163][164] Some of those designs proved successful, whilst others fared less well when tested.
+Research performed by Marc van den Broek revealed older prototypes for more than 100 inventions that are ascribed to Leonardo. Similarities between Leonardo´s illustrations and drawings from the Middle Ages and from Ancient Greece and Rome, the Chinese and Persian Empires, and Egypt suggest that a large portion of Leonardo's inventions had been conceived before his lifetime. Leonardo´s innovation was to combine different functions from existing drafts and set them into scenes that illustrated their utility. By reconstituting technical inventions he created something new.[165]
+Leonardo's fame within his own lifetime was such that the King of France carried him away like a trophy, and was claimed to have supported him in his old age and held him in his arms as he died. Interest in Leonardo and his work has never diminished. Crowds still queue to see his best-known artworks, T-shirts still bear his most famous drawing, and writers continue to hail him as a genius while speculating about his private life, as well as about what one so intelligent actually believed in.[38]
+The continued admiration that Leonardo commanded from painters, critics and historians is reflected in many other written tributes. Baldassare Castiglione, author of Il Cortegiano (The Courtier), wrote in 1528: "...Another of the greatest painters in this world looks down on this art in which he is unequalled..."[166] while the biographer known as "Anonimo Gaddiano" wrote, c. 1540: "His genius was so rare and universal that it can be said that nature worked a miracle on his behalf..."[167]
+Giorgio Vasari, in the enlarged edition of Lives of the Artists (1568)[168] introduced his chapter on Leonardo with the following words:
+In the normal course of events many men and women are born with remarkable talents; but occasionally, in a way that transcends nature, a single person is marvellously endowed by Heaven with beauty, grace and talent in such abundance that he leaves other men far behind, all his actions seem inspired and indeed everything he does clearly comes from God rather than from human skill. Everyone acknowledged that this was true of Leonardo da Vinci, an artist of outstanding physical beauty, who displayed infinite grace in everything that he did and who cultivated his genius so brilliantly that all problems he studied he solved with ease.
+The 19th century brought a particular admiration for Leonardo's genius, causing Henry Fuseli to write in 1801: "Such was the dawn of modern art, when Leonardo da Vinci broke forth with a splendour that distanced former excellence: made up of all the elements that constitute the essence of genius..."[169] This is echoed by A.E. Rio who wrote in 1861: "He towered above all other artists through the strength and the nobility of his talents."[170]
+By the 19th century, the scope of Leonardo's notebooks was known, as well as his paintings. Hippolyte Taine wrote in 1866: "There may not be in the world an example of another genius so universal, so incapable of fulfilment, so full of yearning for the infinite, so naturally refined, so far ahead of his own century and the following centuries."[171]
+Art historian Bernard Berenson wrote in 1896: "Leonardo is the one artist of whom it may be said with perfect literalness: Nothing that he touched but turned into a thing of eternal beauty. Whether it be the cross section of a skull, the structure of a weed, or a study of muscles, he, with his feeling for line and for light and shade, forever transmuted it into life-communicating values."[172]
+The interest in Leonardo's genius has continued unabated; experts study and translate his writings, analyse his paintings using scientific techniques, argue over attributions and search for works which have been recorded but never found.[173] Liana Bortolon, writing in 1967, said: "Because of the multiplicity of interests that spurred him to pursue every field of knowledge...Leonardo can be considered, quite rightly, to have been the universal genius par excellence, and with all the disquieting overtones inherent in that term. Man is as uncomfortable today, faced with a genius, as he was in the 16th century. Five centuries have passed, yet we still view Leonardo with awe."[27]
+Twenty-first-century author Walter Isaacson based much of his biography of Leonardo[104] on thousands of notebook entries, studying the personal notes, sketches, budget notations, and musings of the man whom he considers the greatest of innovators. Isaacson was surprised to discover a "fun, joyous" side of Leonardo in addition to his limitless curiosity and creative genius.[174]
+On the 500th anniversary of Leonardo's death, the Louvre in Paris arranged for the largest ever single exhibit of his work, called Leonardo, between November 2019 and February 2020. The exhibit includes over 100 paintings, drawings and notebooks. Eleven of the paintings that Leonardo completed in his lifetime were included. Five of these are owned by the Louvre, but the Mona Lisa was not included because it is in such great demand among general visitors to the Louvre; it remains on display in its gallery. Vitruvian Man, however, is on display following a legal battle with its owner, the Gallerie dell'Accademia in Venice. Salvator Mundi was also not included because its Saudi owner did not agree to lease the work.[175][176]
+Much of the Collegiate Church of Saint Florentin at the Château d'Amboise, where Leonardo was buried, was damaged during the French Revolution, leading to the church's demolition in 1802.[177] Some of the graves were destroyed in the process, scattering the bones interred there and thereby leaving the whereabouts of Leonardo's remains subject to dispute. While excavating the site in 1863, fine-arts inspector general Arsène Houssaye found a partially complete skeleton with a bronze ring on one finger, some white hair, and stone fragments bearing the inscriptions "EO," "AR," "DUS," and "VINC"—interpreted as forming "Leonardus Vinci".[177][89][178][179] A silver shield found near the bones depicts a beardless Francis I, corresponding to the king's appearance during Leonardo's lifetime, and the skull's inclusion of eight teeth corresponds to someone of approximately the appropriate age.[178] The unusually large skull led Houssaye to believe that he had located Leonardo's remains,[179] but he thought the skeleton seemed too short.[178] Other art historians say that the 1.73 metres (5.7 ft) tall skeleton may well be Leonardo's.[180]
+The remains, except for the ring and a lock of hair which Houssaye kept,[89] were brought to Paris in a lead box, where the skull was allegedly presented to Napoleon III,[178] before being returned to the Château d'Amboise and re-interred in the Chapel of Saint Hubert in 1874.[177] A new memorial tombstone was added by sculptor Francesco La Monaca in the 1930s.[181] Reflecting doubts about the attribution, a plaque above the tomb states that the remains are only presumed to be those of Leonardo.[177] It has since been theorized that the folding of the skeleton's right arm over the head may correspond to the paralysis of Leonardo's right hand.[178][77][83]
+In 2016, it was announced that DNA tests would be conducted to determine whether the attribution is correct.[177] The DNA of the remains will be compared to that of samples collected from Leonardo's work and his half-brother Domenico's descendants;[177] it may also be sequenced.[182] The lock of hair and ring, now in a private US collection,[ac] were displayed in Vinci beginning on 2 May 2019, the 500th anniversary of the artist's death.[183][89]
+Salvator Mundi, a painting by Leonardo depicting Jesus holding an orb, sold for a world record US$450.3 million at a Christie's auction in New York, 15 November 2017.[184] The highest known sale price for any artwork was previously US$300 million, for Willem de Kooning's Interchange, which was sold privately in September 2015.[185] The highest price previously paid for a work of art at auction was for Pablo Picasso's Les Femmes d'Alger, which sold for US$179.4 million in May 2015 at Christie's New York.[185]
+Early sources
+Modern sources

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+Leonhard Euler (/ˈɔɪlər/ OY-lər;[2] German: [ˈɔʏlɐ] (listen)[3]; 15 April 1707 – 18 September 1783) was a Swiss mathematician, physicist, astronomer, geographer, logician and engineer who made important and influential discoveries in many branches of mathematics, such as infinitesimal calculus and graph theory, while also making pioneering contributions to several branches such as topology and analytic number theory. He also introduced much of the modern mathematical terminology and notation, particularly for mathematical analysis, such as the notion of a mathematical function.[4] He is also known for his work in mechanics, fluid dynamics, optics, astronomy and music theory.[5]
+Euler was one of the most eminent mathematicians of the 18th century and is held to be one of the greatest in history. He is also widely considered to be the most prolific, as his collected works fill 92 volumes,[6] more than anyone else in the field. He spent most of his adult life in Saint Petersburg, Russia, and in Berlin, then the capital of Prussia.
+A statement attributed to Pierre-Simon Laplace expresses Euler's influence on mathematics: "Read Euler, read Euler, he is the master of us all."[7][8]
+Leonhard Euler was born on 15 April 1707, in Basel, Switzerland, to Paul III Euler, a pastor of the Reformed Church, and Marguerite née Brucker, another pastor's daughter. He had two younger sisters, Anna Maria and Maria Magdalena, and a younger brother, Johann Heinrich.[9] Soon after the birth of Leonhard, the Eulers moved from Basel to the town of Riehen, Switzerland, where Leonhard spent most of his childhood. Paul was a friend of the Bernoulli family; Johann Bernoulli, then regarded as Europe's foremost mathematician, would eventually be the most important influence on young Leonhard.
+Euler's formal education started in Basel, where he was sent to live with his maternal grandmother. In 1720, at age thirteen, he enrolled at the University of Basel. In 1723, he received a Master of Philosophy with a dissertation that compared the philosophies of Descartes and Newton. During that time, he was receiving Saturday afternoon lessons from Johann Bernoulli, who quickly discovered his new pupil's incredible talent for mathematics.[10] At that time Euler's main studies included theology, Greek and Hebrew at his father's urging to become a pastor, but Bernoulli convinced his father that Leonhard was destined to become a great mathematician.
+In 1726, Euler completed a dissertation on the propagation of sound with the title De Sono.[11] At that time, he was unsuccessfully attempting to obtain a position at the University of Basel. In 1727, he first entered the Paris Academy Prize Problem competition; the problem that year was to find the best way to place the masts on a ship. Pierre Bouguer, who became known as "the father of naval architecture", won and Euler took second place. Euler later won this annual prize twelve times.[12]
+Around this time Johann Bernoulli's two sons, Daniel and Nicolaus, were working at the Imperial Russian Academy of Sciences in Saint Petersburg. On 31 July 1726, Nicolaus died of appendicitis after spending less than a year in Russia.[13][14] When Daniel assumed his brother's position in the mathematics/physics division, he recommended that the post in physiology that he had vacated be filled by his friend Euler. In November 1726 Euler eagerly accepted the offer, but delayed making the trip to Saint Petersburg while he unsuccessfully applied for a physics professorship at the University of Basel.[15]
+Euler arrived in Saint Petersburg on 17 May 1727. He was promoted from his junior post in the medical department of the academy to a position in the mathematics department. He lodged with Daniel Bernoulli with whom he often worked in close collaboration. Euler mastered Russian and settled into life in Saint Petersburg. He also took on an additional job as a medic in the Russian Navy.[16]
+The Academy at Saint Petersburg, established by Peter the Great, was intended to improve education in Russia and to close the scientific gap with Western Europe. As a result, it was made especially attractive to foreign scholars like Euler. The academy possessed ample financial resources and a comprehensive library drawn from the private libraries of Peter himself and of the nobility. Very few students were enrolled in the academy to lessen the faculty's teaching burden. The academy emphasized research and offered to its faculty both the time and the freedom to pursue scientific questions.[12]
+The Academy's benefactress, Catherine I, who had continued the progressive policies of her late husband, died on the day of Euler's arrival. The Russian nobility then gained power upon the ascension of the twelve-year-old Peter II. The nobility, suspicious of the academy's foreign scientists, cut funding and caused other difficulties for Euler and his colleagues.
+Conditions improved slightly after the death of Peter II, and Euler swiftly rose through the ranks in the academy and was made a professor of physics in 1731. Two years later, Daniel Bernoulli, who was fed up with the censorship and hostility he faced at Saint Petersburg, left for Basel. Euler succeeded him as the head of the mathematics department.[17]
+On 7 January 1734, he married Katharina Gsell (1707–1773), a daughter of Georg Gsell, a painter from the Academy Gymnasium.[18] The young couple bought a house by the Neva River. Of their thirteen children, only five survived childhood.[19]
+Concerned about the continuing turmoil in Russia, Euler left St. Petersburg on 19 June 1741 to take up a post at the Berlin Academy, which he had been offered by Frederick the Great of Prussia. He lived for 25 years in Berlin, where he wrote over 380 articles. In Berlin, he published the two works for which he would become most renowned: the Introductio in analysin infinitorum, a text on functions published in 1748, and the Institutiones calculi differentialis,[20] published in 1755 on differential calculus.[21] In 1755, he was elected a foreign member of the Royal Swedish Academy of Sciences.
+In addition, Euler was asked to tutor Friederike Charlotte of Brandenburg-Schwedt, the Princess of Anhalt-Dessau and Frederick's niece. Euler wrote over 200 letters to her in the early 1760s, which were later compiled into a best-selling volume entitled Letters of Euler on different Subjects in Natural Philosophy Addressed to a German Princess.[22] This work contained Euler's exposition on various subjects pertaining to physics and mathematics, as well as offering valuable insights into Euler's personality and religious beliefs. This book became more widely read than any of his mathematical works and was published across Europe and in the United States. The popularity of the "Letters" testifies to Euler's ability to communicate scientific matters effectively to a lay audience, a rare ability for a dedicated research scientist.[21]
+Despite Euler's immense contribution to the Academy's prestige, he eventually incurred the ire of Frederick and ended up having to leave Berlin. The Prussian king had a large circle of intellectuals in his court, and he found the mathematician unsophisticated and ill-informed on matters beyond numbers and figures. Euler was a simple, devoutly religious man who never questioned the existing social order or conventional beliefs, in many ways the polar opposite of Voltaire, who enjoyed a high place of prestige at Frederick's court. Euler was not a skilled debater and often made it a point to argue subjects that he knew little about, making him the frequent target of Voltaire's wit.[21] Frederick also expressed disappointment with Euler's practical engineering abilities:
+I wanted to have a water jet in my garden: Euler calculated the force of the wheels necessary to raise the water to a reservoir, from where it should fall back through channels, finally spurting out in Sanssouci. My mill was carried out geometrically and could not raise a mouthful of water closer than fifty paces to the reservoir. Vanity of vanities! Vanity of geometry![23]
+Euler's eyesight worsened throughout his mathematical career. In 1738, three years after nearly expiring from fever, he became almost blind in his right eye, but Euler rather blamed the painstaking work on cartography he performed for the St. Petersburg Academy for his condition. Euler's vision in that eye worsened throughout his stay in Germany, to the extent that Frederick referred to him as "Cyclops". Euler remarked on his loss of vision, "Now I will have fewer distractions."[24] He later developed a cataract in his left eye, which was discovered in 1766. Just a few weeks after its discovery, a failed surgical restoration rendered him almost totally blind. He was 59 years old then. However, his condition appeared to have little effect on his productivity, as he compensated for it with his mental calculation skills and exceptional memory. For example, Euler could repeat the Aeneid of Virgil from beginning to end without hesitation, and for every page in the edition he could indicate which line was the first and which the last. With the aid of his scribes, Euler's productivity on many areas of study actually increased. He produced, on average, one mathematical paper every week in the year 1775.[25] The Eulers bore a double name, Euler-Schölpi, the latter of which derives from schelb and schief, signifying squint-eyed, cross-eyed, or crooked. This suggests that the Eulers had a susceptibility to eye problems.[26]
+In 1760, with the Seven Years' War raging, Euler's farm in Charlottenburg was sacked by advancing Russian troops. Upon learning of this event, General Ivan Petrovich Saltykov paid compensation for the damage caused to Euler's estate, with Empress Elizabeth of Russia later adding a further payment of 4000 roubles—an exorbitant amount at the time.[27] The political situation in Russia stabilized after Catherine the Great's accession to the throne, so in 1766 Euler accepted an invitation to return to the St. Petersburg Academy. His conditions were quite exorbitant—a 3000 ruble annual salary, a pension for his wife, and the promise of high-ranking appointments for his sons. All of these requests were granted. He spent the rest of his life in Russia. However, his second stay in the country was marred by tragedy. A fire in St. Petersburg in 1771 cost him his home, and almost his life. In 1773, he lost his wife Katharina after 40 years of marriage.
+Three years after his wife's death, Euler married her half-sister, Salome Abigail Gsell (1723–1794).[28] This marriage lasted until his death. In 1782 he was elected a Foreign Honorary Member of the American Academy of Arts and Sciences.[29]
+In St. Petersburg on 18 September 1783, after a lunch with his family, Euler was discussing the newly discovered planet Uranus and its orbit with a fellow academician Anders Johan Lexell, when he collapsed from a brain hemorrhage. He died a few hours later.[30] Jacob von Staehlin-Storcksburg wrote a short obituary for the Russian Academy of Sciences and Russian mathematician Nicolas Fuss, one of Euler's disciples, wrote a more detailed eulogy,[31] which he delivered at a memorial meeting. In his eulogy for the French Academy, French mathematician and philosopher Marquis de Condorcet, wrote:
+il cessa de calculer et de vivre— ... he ceased to calculate and to live.[32]
+Euler was buried next to Katharina at the Smolensk Lutheran Cemetery on Goloday Island. In 1785, the Russian Academy of Sciences put a marble bust of Leonhard Euler on a pedestal next to the Director's seat and, in 1837, placed a headstone on Euler's grave. To commemorate the 250th anniversary of Euler's birth, the headstone was moved in 1956, together with his remains, to the 18th-century necropolis at the Alexander Nevsky Monastery.
+Euler worked in almost all areas of mathematics, such as geometry, infinitesimal calculus, trigonometry, algebra, and number theory, as well as continuum physics, lunar theory and other areas of physics. He is a seminal figure in the history of mathematics; if printed, his works, many of which are of fundamental interest, would occupy between 60 and 80 quarto volumes.[25] Euler's name is associated with a large number of topics.
+Euler is the only mathematician to have two numbers named after him: the important Euler's number in calculus, e, approximately equal to 2.71828, and the Euler–Mascheroni constant γ (gamma) sometimes referred to as just "Euler's constant", approximately equal to 0.57721. It is not known whether γ is rational or irrational.[33]
+Euler introduced and popularized several notational conventions through his numerous and widely circulated textbooks. Most notably, he introduced the concept of a function[4] and was the first to write f(x) to denote the function f applied to the argument x. He also introduced the modern notation for the trigonometric functions, the letter e for the base of the natural logarithm (now also known as Euler's number), the Greek letter Σ for summations and the letter i to denote the imaginary unit.[34] The use of the Greek letter π to denote the ratio of a circle's circumference to its diameter was also popularized by Euler, although it originated with Welsh mathematician William Jones.[35]
+The development of infinitesimal calculus was at the forefront of 18th-century mathematical research, and the Bernoullis—family friends of Euler—were responsible for much of the early progress in the field. Thanks to their influence, studying calculus became the major focus of Euler's work. While some of Euler's proofs are not acceptable by modern standards of mathematical rigour[36] (in particular his reliance on the principle of the generality of algebra), his ideas led to many great advances.
+Euler is well known in analysis for his frequent use and development of power series, the expression of functions as sums of infinitely many terms, such as
+Euler directly proved the power series expansions for e and the inverse tangent function. (Indirect proof via the inverse power series technique was given by Newton and Leibniz between 1670 and 1680.) His daring use of power series enabled him to solve the famous Basel problem in 1735 (he provided a more elaborate argument in 1741):[36]
+Euler introduced the use of the exponential function and logarithms in analytic proofs. He discovered ways to express various logarithmic functions using power series, and he successfully defined logarithms for negative and complex numbers, thus greatly expanding the scope of mathematical applications of logarithms.[34] He also defined the exponential function for complex numbers, and discovered its relation to the trigonometric functions. For any real number φ (taken to be radians), Euler's formula states that the complex exponential function satisfies
+A special case of the above formula is known as Euler's identity,
+called "the most remarkable formula in mathematics" by Richard P. Feynman, for its single uses of the notions of addition, multiplication, exponentiation, and equality, and the single uses of the important constants 0, 1, e, i and π.[37] In 1988, readers of the Mathematical Intelligencer voted it "the Most Beautiful Mathematical Formula Ever".[38] In total, Euler was responsible for three of the top five formulae in that poll.[38]
+De Moivre's formula is a direct consequence of Euler's formula.
+Euler elaborated the theory of higher transcendental functions by introducing the gamma function and introduced a new method for solving quartic equations. He found a way to calculate integrals with complex limits, foreshadowing the development of modern complex analysis. He invented the calculus of variations including its best-known result, the Euler–Lagrange equation.
+Euler pioneered the use of analytic methods to solve number theory problems. In doing so, he united two disparate branches of mathematics and introduced a new field of study, analytic number theory. In breaking ground for this new field, Euler created the theory of hypergeometric series, q-series, hyperbolic trigonometric functions and the analytic theory of continued fractions. For example, he proved the infinitude of primes using the divergence of the harmonic series, and he used analytic methods to gain some understanding of the way prime numbers are distributed. Euler's work in this area led to the development of the prime number theorem.[39]
+Euler's interest in number theory can be traced to the influence of Christian Goldbach, his friend in the St. Petersburg Academy. A lot of Euler's early work on number theory was based on the works of Pierre de Fermat. Euler developed some of Fermat's ideas and disproved some of his conjectures.
+Euler linked the nature of prime distribution with ideas in analysis. He proved that the sum of the reciprocals of the primes diverges. In doing so, he discovered the connection between the Riemann zeta function and the prime numbers; this is known as the Euler product formula for the Riemann zeta function.
+Euler proved Newton's identities, Fermat's little theorem, Fermat's theorem on sums of two squares, and he made distinct contributions to Lagrange's four-square theorem. He also invented the totient function φ(n), the number of positive integers less than or equal to the integer n that are coprime to n. Using properties of this function, he generalized Fermat's little theorem to what is now known as Euler's theorem. He contributed significantly to the theory of perfect numbers, which had fascinated mathematicians since Euclid. He proved that the relationship shown between even perfect numbers and Mersenne primes earlier proved by Euclid was one-to-one, a result otherwise known as the Euclid–Euler theorem. Euler also conjectured the law of quadratic reciprocity. The concept is regarded as a fundamental theorem of number theory, and his ideas paved the way for the work of Carl Friedrich Gauss.[40]
+By 1772 Euler had proved that 231 − 1 = 2,147,483,647 is a Mersenne prime. It may have remained the largest known prime until 1867.[41]
+In 1735, Euler presented a solution to the problem known as the Seven Bridges of Königsberg.[42] The city of Königsberg, Prussia was set on the Pregel River, and included two large islands that were connected to each other and the mainland by seven bridges. The problem is to decide whether it is possible to follow a path that crosses each bridge exactly once and returns to the starting point. It is not possible: there is no Eulerian circuit. This solution is considered to be the first theorem of graph theory, specifically of planar graph theory.[42]
+Euler also discovered the formula
+V
+−
+E
++
+F
+=
+2
+{\displaystyle V-E+F=2}
+ relating the number of vertices, edges and faces of a convex polyhedron,[43] and hence of a planar graph. The constant in this formula is now known as the Euler characteristic for the graph (or other mathematical object), and is related to the genus of the object.[44] The study and generalization of this formula, specifically by Cauchy[45] and L'Huilier,[46] is at the origin of topology.
+Some of Euler's greatest successes were in solving real-world problems analytically, and in describing numerous applications of the Bernoulli numbers, Fourier series, Euler numbers, the constants e and π, continued fractions and integrals. He integrated Leibniz's differential calculus with Newton's Method of Fluxions, and developed tools that made it easier to apply calculus to physical problems. He made great strides in improving the numerical approximation of integrals, inventing what are now known as the Euler approximations. The most notable of these approximations are Euler's method and the Euler–Maclaurin formula. He also facilitated the use of differential equations, in particular introducing the Euler–Mascheroni constant:
+One of Euler's more unusual interests was the application of mathematical ideas in music. In 1739 he wrote the Tentamen novae theoriae musicae, hoping to eventually incorporate musical theory as part of mathematics. This part of his work, however, did not receive wide attention and was once described as too mathematical for musicians and too musical for mathematicians.[47]
+In 1911, almost 130 years after Euler's death, Alfred J. Lotka used Euler's work to derive the Euler–Lotka equation for calculating rates of population growth for age-structured populations, a fundamental method that is commonly used in population biology and ecology.
+Euler helped develop the Euler–Bernoulli beam equation, which became a cornerstone of engineering. Besides successfully applying his analytic tools to problems in classical mechanics, Euler applied these techniques to celestial problems. His work in astronomy was recognized by multiple Paris Academy Prizes over the course of his career. His accomplishments include determining with great accuracy the orbits of comets and other celestial bodies, understanding the nature of comets, and calculating the parallax of the Sun. His calculations contributed to the development of accurate longitude tables.[48]
+Euler made important contributions in optics. He disagreed with Newton's corpuscular theory of light in the Opticks, which was then the prevailing theory. His 1740s papers on optics helped ensure that the wave theory of light proposed by Christiaan Huygens would become the dominant mode of thought, at least until the development of the quantum theory of light.[49]
+In 1757 he published an important set of equations for inviscid flow, that are now known as the Euler equations.[50] In differential form, the equations are:
+where
+Euler is well known in structural engineering for his formula giving the critical buckling load of an ideal strut, which depends only on its length and flexural stiffness:[51]
+where
+Euler is credited with using closed curves to illustrate syllogistic reasoning (1768). These diagrams have become known as Euler diagrams.[52]
+An Euler diagram is a diagrammatic means of representing sets and their relationships. Euler diagrams consist of simple closed curves (usually circles) in the plane that depict sets.  Each Euler curve divides the plane into two regions or "zones": the interior, which symbolically represents the elements of the set, and the exterior, which represents all elements that are not members of the set.  The sizes or shapes of the curves are not important; the significance of the diagram is in how they overlap. The spatial relationships between the regions bounded by each curve (overlap, containment or neither) corresponds to set-theoretic relationships (intersection, subset and disjointness).   Curves whose interior zones do not intersect represent disjoint sets.  Two curves whose interior zones intersect represent sets that have common elements; the zone inside both curves represents the set of elements common to both sets (the intersection of the sets).  A curve that is contained completely within the interior zone of another represents a subset of it. Euler diagrams (and their generalization in Venn diagrams) were incorporated as part of instruction in set theory as part of the new math movement in the 1960s. Since then, they have also been adopted by other curriculum fields such as reading.[53]
+Even when dealing with music, Euler's approach is mainly mathematical. His writings on music are not particularly numerous (a few hundred pages, in his total production of about thirty thousand pages), but they reflect an early preoccupation and one that did not leave him throughout his life.[54]
+A first point of Euler's musical theory is the definition of "genres", i.e. of possible divisions of the octave using the prime numbers 3 and 5. Euler describes 18 such genres, with the general definition 2mA, where A is the "exponent" of the genre (i.e. the sum of the exponents of 3 and 5) and 2m (where "m is an indefinite number, small or large, so long as the sounds are perceptible"[55]), expresses that the relation holds independently of the number of octaves concerned. The first genre, with A = 1, is the octave itself (or its duplicates); the second genre, 2m.3, is the octave divided by the fifth (fifth + fourth, C–G–C); the third genre is 2m.5, major third + minor sixth (C–E–C); the fourth is 2m.32, two-fourths and a tone (C–F–B♭–C); the fifth is 2m.3.5 (C–E–G–B–C); etc. Genres 12 (2m.33.5), 13 (2m.32.52) and 14 (2m.3.53) are corrected versions of the diatonic, chromatic and enharmonic, respectively, of the Ancients. Genre 18 (2m.33.52) is the "diatonico-chromatic", "used generally in all compositions",[56] and which turns out to be identical with the system described by Johann Mattheson.[57] Euler later envisaged the possibility of describing genres including the prime number 7.[58]
+Euler devised a specific graph, the Speculum musicum,[59] to illustrate the diatonico-chromatic genre, and discussed paths in this graph for specific intervals, recalling his interest in the Seven Bridges of Königsberg (see above). The device drew renewed interest as the Tonnetz in neo-Riemannian theory (see also Lattice (music)).[60]
+Euler further used the principle of the "exponent" to propose a derivation of the gradus suavitatis (degree of suavity, of agreeableness) of intervals and chords from their prime factors – one must keep in mind that he considered just intonation, i.e. 1 and the prime numbers 3 and 5 only.[61] Formulas have been proposed extending this system to any number of prime numbers, e.g. in the form
+where pi are prime numbers and ki their exponents.[62]
+Euler and his friend Daniel Bernoulli were opponents of Leibniz's monadism and the philosophy of Christian Wolff. Euler insisted that knowledge is founded in part on the basis of precise quantitative laws, something that monadism and Wolffian science were unable to provide. Euler's religious leanings might also have had a bearing on his dislike of the doctrine; he went so far as to label Wolff's ideas as "heathen and atheistic".[63]
+Much of what is known of Euler's religious beliefs can be deduced from his Letters to a German Princess and an earlier work, Rettung der Göttlichen Offenbahrung Gegen die Einwürfe der Freygeister (Defense of the Divine Revelation against the Objections of the Freethinkers). These works show that Euler was a devout Christian who believed the Bible to be inspired; the Rettung was primarily an argument for the divine inspiration of scripture.[64]
+There is a famous legend[65] inspired by Euler's arguments with secular philosophers over religion, which is set during Euler's second stint at the St. Petersburg Academy. The French philosopher Denis Diderot was visiting Russia on Catherine the Great's invitation. However, the Empress was alarmed that the philosopher's arguments for atheism were influencing members of her court, and so Euler was asked to confront the Frenchman. Diderot was informed that a learned mathematician had produced a proof of the existence of God: he agreed to view the proof as it was presented in court. Euler appeared, advanced toward Diderot, and in a tone of perfect conviction announced this non-sequitur: "Sir, a+bn/n=x, hence God exists—reply!"
+Diderot, to whom (says the story) all mathematics was gibberish, stood dumbstruck as peals of laughter erupted from the court.  Embarrassed, he asked to leave Russia, a request that was graciously granted by the Empress. However amusing the anecdote may be, it is apocryphal, given that Diderot himself did research in mathematics.[66]
+The legend was apparently first told by
+Dieudonné Thiébault[67] with embellishment by Augustus De Morgan.[68][69]
+Euler was featured on the sixth series of the Swiss 10-franc banknote and on numerous Swiss, German, and Russian postage stamps. The asteroid 2002 Euler was named in his honor. He is also commemorated by the Lutheran Church on their Calendar of Saints on 24 May—he was a devout Christian (and believer in biblical inerrancy) who wrote apologetics and argued forcefully against the prominent atheists of his time.[64]
+Euler has an extensive bibliography. His best-known books include:
+The first collection of Euler's work was made by Paul Heinrich von Fuss in 1862.[71] A definitive collection of Euler's works, entitled Opera Omnia, has been published since 1911 by the Euler Commission of the Swiss Academy of Sciences. A complete chronological list of Euler's works is available at The Eneström Index.[72] Full text, open access versions of many of Euler's papers are available in the original language and English translations at the Euler Archive, hosted by University of the Pacific. The Euler Archive was started at Dartmouth College[73] before moving to the Mathematical Association of America[74] and, most recently, to University of the Pacific in 2017.
+Illustration from Solutio problematis... a. 1743 propositi published in Acta Eruditorum, 1744
+The title page of Euler's Methodus inveniendi lineas curvas.

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+Leonhard Euler (/ˈɔɪlər/ OY-lər;[2] German: [ˈɔʏlɐ] (listen)[3]; 15 April 1707 – 18 September 1783) was a Swiss mathematician, physicist, astronomer, geographer, logician and engineer who made important and influential discoveries in many branches of mathematics, such as infinitesimal calculus and graph theory, while also making pioneering contributions to several branches such as topology and analytic number theory. He also introduced much of the modern mathematical terminology and notation, particularly for mathematical analysis, such as the notion of a mathematical function.[4] He is also known for his work in mechanics, fluid dynamics, optics, astronomy and music theory.[5]
+Euler was one of the most eminent mathematicians of the 18th century and is held to be one of the greatest in history. He is also widely considered to be the most prolific, as his collected works fill 92 volumes,[6] more than anyone else in the field. He spent most of his adult life in Saint Petersburg, Russia, and in Berlin, then the capital of Prussia.
+A statement attributed to Pierre-Simon Laplace expresses Euler's influence on mathematics: "Read Euler, read Euler, he is the master of us all."[7][8]
+Leonhard Euler was born on 15 April 1707, in Basel, Switzerland, to Paul III Euler, a pastor of the Reformed Church, and Marguerite née Brucker, another pastor's daughter. He had two younger sisters, Anna Maria and Maria Magdalena, and a younger brother, Johann Heinrich.[9] Soon after the birth of Leonhard, the Eulers moved from Basel to the town of Riehen, Switzerland, where Leonhard spent most of his childhood. Paul was a friend of the Bernoulli family; Johann Bernoulli, then regarded as Europe's foremost mathematician, would eventually be the most important influence on young Leonhard.
+Euler's formal education started in Basel, where he was sent to live with his maternal grandmother. In 1720, at age thirteen, he enrolled at the University of Basel. In 1723, he received a Master of Philosophy with a dissertation that compared the philosophies of Descartes and Newton. During that time, he was receiving Saturday afternoon lessons from Johann Bernoulli, who quickly discovered his new pupil's incredible talent for mathematics.[10] At that time Euler's main studies included theology, Greek and Hebrew at his father's urging to become a pastor, but Bernoulli convinced his father that Leonhard was destined to become a great mathematician.
+In 1726, Euler completed a dissertation on the propagation of sound with the title De Sono.[11] At that time, he was unsuccessfully attempting to obtain a position at the University of Basel. In 1727, he first entered the Paris Academy Prize Problem competition; the problem that year was to find the best way to place the masts on a ship. Pierre Bouguer, who became known as "the father of naval architecture", won and Euler took second place. Euler later won this annual prize twelve times.[12]
+Around this time Johann Bernoulli's two sons, Daniel and Nicolaus, were working at the Imperial Russian Academy of Sciences in Saint Petersburg. On 31 July 1726, Nicolaus died of appendicitis after spending less than a year in Russia.[13][14] When Daniel assumed his brother's position in the mathematics/physics division, he recommended that the post in physiology that he had vacated be filled by his friend Euler. In November 1726 Euler eagerly accepted the offer, but delayed making the trip to Saint Petersburg while he unsuccessfully applied for a physics professorship at the University of Basel.[15]
+Euler arrived in Saint Petersburg on 17 May 1727. He was promoted from his junior post in the medical department of the academy to a position in the mathematics department. He lodged with Daniel Bernoulli with whom he often worked in close collaboration. Euler mastered Russian and settled into life in Saint Petersburg. He also took on an additional job as a medic in the Russian Navy.[16]
+The Academy at Saint Petersburg, established by Peter the Great, was intended to improve education in Russia and to close the scientific gap with Western Europe. As a result, it was made especially attractive to foreign scholars like Euler. The academy possessed ample financial resources and a comprehensive library drawn from the private libraries of Peter himself and of the nobility. Very few students were enrolled in the academy to lessen the faculty's teaching burden. The academy emphasized research and offered to its faculty both the time and the freedom to pursue scientific questions.[12]
+The Academy's benefactress, Catherine I, who had continued the progressive policies of her late husband, died on the day of Euler's arrival. The Russian nobility then gained power upon the ascension of the twelve-year-old Peter II. The nobility, suspicious of the academy's foreign scientists, cut funding and caused other difficulties for Euler and his colleagues.
+Conditions improved slightly after the death of Peter II, and Euler swiftly rose through the ranks in the academy and was made a professor of physics in 1731. Two years later, Daniel Bernoulli, who was fed up with the censorship and hostility he faced at Saint Petersburg, left for Basel. Euler succeeded him as the head of the mathematics department.[17]
+On 7 January 1734, he married Katharina Gsell (1707–1773), a daughter of Georg Gsell, a painter from the Academy Gymnasium.[18] The young couple bought a house by the Neva River. Of their thirteen children, only five survived childhood.[19]
+Concerned about the continuing turmoil in Russia, Euler left St. Petersburg on 19 June 1741 to take up a post at the Berlin Academy, which he had been offered by Frederick the Great of Prussia. He lived for 25 years in Berlin, where he wrote over 380 articles. In Berlin, he published the two works for which he would become most renowned: the Introductio in analysin infinitorum, a text on functions published in 1748, and the Institutiones calculi differentialis,[20] published in 1755 on differential calculus.[21] In 1755, he was elected a foreign member of the Royal Swedish Academy of Sciences.
+In addition, Euler was asked to tutor Friederike Charlotte of Brandenburg-Schwedt, the Princess of Anhalt-Dessau and Frederick's niece. Euler wrote over 200 letters to her in the early 1760s, which were later compiled into a best-selling volume entitled Letters of Euler on different Subjects in Natural Philosophy Addressed to a German Princess.[22] This work contained Euler's exposition on various subjects pertaining to physics and mathematics, as well as offering valuable insights into Euler's personality and religious beliefs. This book became more widely read than any of his mathematical works and was published across Europe and in the United States. The popularity of the "Letters" testifies to Euler's ability to communicate scientific matters effectively to a lay audience, a rare ability for a dedicated research scientist.[21]
+Despite Euler's immense contribution to the Academy's prestige, he eventually incurred the ire of Frederick and ended up having to leave Berlin. The Prussian king had a large circle of intellectuals in his court, and he found the mathematician unsophisticated and ill-informed on matters beyond numbers and figures. Euler was a simple, devoutly religious man who never questioned the existing social order or conventional beliefs, in many ways the polar opposite of Voltaire, who enjoyed a high place of prestige at Frederick's court. Euler was not a skilled debater and often made it a point to argue subjects that he knew little about, making him the frequent target of Voltaire's wit.[21] Frederick also expressed disappointment with Euler's practical engineering abilities:
+I wanted to have a water jet in my garden: Euler calculated the force of the wheels necessary to raise the water to a reservoir, from where it should fall back through channels, finally spurting out in Sanssouci. My mill was carried out geometrically and could not raise a mouthful of water closer than fifty paces to the reservoir. Vanity of vanities! Vanity of geometry![23]
+Euler's eyesight worsened throughout his mathematical career. In 1738, three years after nearly expiring from fever, he became almost blind in his right eye, but Euler rather blamed the painstaking work on cartography he performed for the St. Petersburg Academy for his condition. Euler's vision in that eye worsened throughout his stay in Germany, to the extent that Frederick referred to him as "Cyclops". Euler remarked on his loss of vision, "Now I will have fewer distractions."[24] He later developed a cataract in his left eye, which was discovered in 1766. Just a few weeks after its discovery, a failed surgical restoration rendered him almost totally blind. He was 59 years old then. However, his condition appeared to have little effect on his productivity, as he compensated for it with his mental calculation skills and exceptional memory. For example, Euler could repeat the Aeneid of Virgil from beginning to end without hesitation, and for every page in the edition he could indicate which line was the first and which the last. With the aid of his scribes, Euler's productivity on many areas of study actually increased. He produced, on average, one mathematical paper every week in the year 1775.[25] The Eulers bore a double name, Euler-Schölpi, the latter of which derives from schelb and schief, signifying squint-eyed, cross-eyed, or crooked. This suggests that the Eulers had a susceptibility to eye problems.[26]
+In 1760, with the Seven Years' War raging, Euler's farm in Charlottenburg was sacked by advancing Russian troops. Upon learning of this event, General Ivan Petrovich Saltykov paid compensation for the damage caused to Euler's estate, with Empress Elizabeth of Russia later adding a further payment of 4000 roubles—an exorbitant amount at the time.[27] The political situation in Russia stabilized after Catherine the Great's accession to the throne, so in 1766 Euler accepted an invitation to return to the St. Petersburg Academy. His conditions were quite exorbitant—a 3000 ruble annual salary, a pension for his wife, and the promise of high-ranking appointments for his sons. All of these requests were granted. He spent the rest of his life in Russia. However, his second stay in the country was marred by tragedy. A fire in St. Petersburg in 1771 cost him his home, and almost his life. In 1773, he lost his wife Katharina after 40 years of marriage.
+Three years after his wife's death, Euler married her half-sister, Salome Abigail Gsell (1723–1794).[28] This marriage lasted until his death. In 1782 he was elected a Foreign Honorary Member of the American Academy of Arts and Sciences.[29]
+In St. Petersburg on 18 September 1783, after a lunch with his family, Euler was discussing the newly discovered planet Uranus and its orbit with a fellow academician Anders Johan Lexell, when he collapsed from a brain hemorrhage. He died a few hours later.[30] Jacob von Staehlin-Storcksburg wrote a short obituary for the Russian Academy of Sciences and Russian mathematician Nicolas Fuss, one of Euler's disciples, wrote a more detailed eulogy,[31] which he delivered at a memorial meeting. In his eulogy for the French Academy, French mathematician and philosopher Marquis de Condorcet, wrote:
+il cessa de calculer et de vivre— ... he ceased to calculate and to live.[32]
+Euler was buried next to Katharina at the Smolensk Lutheran Cemetery on Goloday Island. In 1785, the Russian Academy of Sciences put a marble bust of Leonhard Euler on a pedestal next to the Director's seat and, in 1837, placed a headstone on Euler's grave. To commemorate the 250th anniversary of Euler's birth, the headstone was moved in 1956, together with his remains, to the 18th-century necropolis at the Alexander Nevsky Monastery.
+Euler worked in almost all areas of mathematics, such as geometry, infinitesimal calculus, trigonometry, algebra, and number theory, as well as continuum physics, lunar theory and other areas of physics. He is a seminal figure in the history of mathematics; if printed, his works, many of which are of fundamental interest, would occupy between 60 and 80 quarto volumes.[25] Euler's name is associated with a large number of topics.
+Euler is the only mathematician to have two numbers named after him: the important Euler's number in calculus, e, approximately equal to 2.71828, and the Euler–Mascheroni constant γ (gamma) sometimes referred to as just "Euler's constant", approximately equal to 0.57721. It is not known whether γ is rational or irrational.[33]
+Euler introduced and popularized several notational conventions through his numerous and widely circulated textbooks. Most notably, he introduced the concept of a function[4] and was the first to write f(x) to denote the function f applied to the argument x. He also introduced the modern notation for the trigonometric functions, the letter e for the base of the natural logarithm (now also known as Euler's number), the Greek letter Σ for summations and the letter i to denote the imaginary unit.[34] The use of the Greek letter π to denote the ratio of a circle's circumference to its diameter was also popularized by Euler, although it originated with Welsh mathematician William Jones.[35]
+The development of infinitesimal calculus was at the forefront of 18th-century mathematical research, and the Bernoullis—family friends of Euler—were responsible for much of the early progress in the field. Thanks to their influence, studying calculus became the major focus of Euler's work. While some of Euler's proofs are not acceptable by modern standards of mathematical rigour[36] (in particular his reliance on the principle of the generality of algebra), his ideas led to many great advances.
+Euler is well known in analysis for his frequent use and development of power series, the expression of functions as sums of infinitely many terms, such as
+Euler directly proved the power series expansions for e and the inverse tangent function. (Indirect proof via the inverse power series technique was given by Newton and Leibniz between 1670 and 1680.) His daring use of power series enabled him to solve the famous Basel problem in 1735 (he provided a more elaborate argument in 1741):[36]
+Euler introduced the use of the exponential function and logarithms in analytic proofs. He discovered ways to express various logarithmic functions using power series, and he successfully defined logarithms for negative and complex numbers, thus greatly expanding the scope of mathematical applications of logarithms.[34] He also defined the exponential function for complex numbers, and discovered its relation to the trigonometric functions. For any real number φ (taken to be radians), Euler's formula states that the complex exponential function satisfies
+A special case of the above formula is known as Euler's identity,
+called "the most remarkable formula in mathematics" by Richard P. Feynman, for its single uses of the notions of addition, multiplication, exponentiation, and equality, and the single uses of the important constants 0, 1, e, i and π.[37] In 1988, readers of the Mathematical Intelligencer voted it "the Most Beautiful Mathematical Formula Ever".[38] In total, Euler was responsible for three of the top five formulae in that poll.[38]
+De Moivre's formula is a direct consequence of Euler's formula.
+Euler elaborated the theory of higher transcendental functions by introducing the gamma function and introduced a new method for solving quartic equations. He found a way to calculate integrals with complex limits, foreshadowing the development of modern complex analysis. He invented the calculus of variations including its best-known result, the Euler–Lagrange equation.
+Euler pioneered the use of analytic methods to solve number theory problems. In doing so, he united two disparate branches of mathematics and introduced a new field of study, analytic number theory. In breaking ground for this new field, Euler created the theory of hypergeometric series, q-series, hyperbolic trigonometric functions and the analytic theory of continued fractions. For example, he proved the infinitude of primes using the divergence of the harmonic series, and he used analytic methods to gain some understanding of the way prime numbers are distributed. Euler's work in this area led to the development of the prime number theorem.[39]
+Euler's interest in number theory can be traced to the influence of Christian Goldbach, his friend in the St. Petersburg Academy. A lot of Euler's early work on number theory was based on the works of Pierre de Fermat. Euler developed some of Fermat's ideas and disproved some of his conjectures.
+Euler linked the nature of prime distribution with ideas in analysis. He proved that the sum of the reciprocals of the primes diverges. In doing so, he discovered the connection between the Riemann zeta function and the prime numbers; this is known as the Euler product formula for the Riemann zeta function.
+Euler proved Newton's identities, Fermat's little theorem, Fermat's theorem on sums of two squares, and he made distinct contributions to Lagrange's four-square theorem. He also invented the totient function φ(n), the number of positive integers less than or equal to the integer n that are coprime to n. Using properties of this function, he generalized Fermat's little theorem to what is now known as Euler's theorem. He contributed significantly to the theory of perfect numbers, which had fascinated mathematicians since Euclid. He proved that the relationship shown between even perfect numbers and Mersenne primes earlier proved by Euclid was one-to-one, a result otherwise known as the Euclid–Euler theorem. Euler also conjectured the law of quadratic reciprocity. The concept is regarded as a fundamental theorem of number theory, and his ideas paved the way for the work of Carl Friedrich Gauss.[40]
+By 1772 Euler had proved that 231 − 1 = 2,147,483,647 is a Mersenne prime. It may have remained the largest known prime until 1867.[41]
+In 1735, Euler presented a solution to the problem known as the Seven Bridges of Königsberg.[42] The city of Königsberg, Prussia was set on the Pregel River, and included two large islands that were connected to each other and the mainland by seven bridges. The problem is to decide whether it is possible to follow a path that crosses each bridge exactly once and returns to the starting point. It is not possible: there is no Eulerian circuit. This solution is considered to be the first theorem of graph theory, specifically of planar graph theory.[42]
+Euler also discovered the formula
+V
+−
+E
++
+F
+=
+2
+{\displaystyle V-E+F=2}
+ relating the number of vertices, edges and faces of a convex polyhedron,[43] and hence of a planar graph. The constant in this formula is now known as the Euler characteristic for the graph (or other mathematical object), and is related to the genus of the object.[44] The study and generalization of this formula, specifically by Cauchy[45] and L'Huilier,[46] is at the origin of topology.
+Some of Euler's greatest successes were in solving real-world problems analytically, and in describing numerous applications of the Bernoulli numbers, Fourier series, Euler numbers, the constants e and π, continued fractions and integrals. He integrated Leibniz's differential calculus with Newton's Method of Fluxions, and developed tools that made it easier to apply calculus to physical problems. He made great strides in improving the numerical approximation of integrals, inventing what are now known as the Euler approximations. The most notable of these approximations are Euler's method and the Euler–Maclaurin formula. He also facilitated the use of differential equations, in particular introducing the Euler–Mascheroni constant:
+One of Euler's more unusual interests was the application of mathematical ideas in music. In 1739 he wrote the Tentamen novae theoriae musicae, hoping to eventually incorporate musical theory as part of mathematics. This part of his work, however, did not receive wide attention and was once described as too mathematical for musicians and too musical for mathematicians.[47]
+In 1911, almost 130 years after Euler's death, Alfred J. Lotka used Euler's work to derive the Euler–Lotka equation for calculating rates of population growth for age-structured populations, a fundamental method that is commonly used in population biology and ecology.
+Euler helped develop the Euler–Bernoulli beam equation, which became a cornerstone of engineering. Besides successfully applying his analytic tools to problems in classical mechanics, Euler applied these techniques to celestial problems. His work in astronomy was recognized by multiple Paris Academy Prizes over the course of his career. His accomplishments include determining with great accuracy the orbits of comets and other celestial bodies, understanding the nature of comets, and calculating the parallax of the Sun. His calculations contributed to the development of accurate longitude tables.[48]
+Euler made important contributions in optics. He disagreed with Newton's corpuscular theory of light in the Opticks, which was then the prevailing theory. His 1740s papers on optics helped ensure that the wave theory of light proposed by Christiaan Huygens would become the dominant mode of thought, at least until the development of the quantum theory of light.[49]
+In 1757 he published an important set of equations for inviscid flow, that are now known as the Euler equations.[50] In differential form, the equations are:
+where
+Euler is well known in structural engineering for his formula giving the critical buckling load of an ideal strut, which depends only on its length and flexural stiffness:[51]
+where
+Euler is credited with using closed curves to illustrate syllogistic reasoning (1768). These diagrams have become known as Euler diagrams.[52]
+An Euler diagram is a diagrammatic means of representing sets and their relationships. Euler diagrams consist of simple closed curves (usually circles) in the plane that depict sets.  Each Euler curve divides the plane into two regions or "zones": the interior, which symbolically represents the elements of the set, and the exterior, which represents all elements that are not members of the set.  The sizes or shapes of the curves are not important; the significance of the diagram is in how they overlap. The spatial relationships between the regions bounded by each curve (overlap, containment or neither) corresponds to set-theoretic relationships (intersection, subset and disjointness).   Curves whose interior zones do not intersect represent disjoint sets.  Two curves whose interior zones intersect represent sets that have common elements; the zone inside both curves represents the set of elements common to both sets (the intersection of the sets).  A curve that is contained completely within the interior zone of another represents a subset of it. Euler diagrams (and their generalization in Venn diagrams) were incorporated as part of instruction in set theory as part of the new math movement in the 1960s. Since then, they have also been adopted by other curriculum fields such as reading.[53]
+Even when dealing with music, Euler's approach is mainly mathematical. His writings on music are not particularly numerous (a few hundred pages, in his total production of about thirty thousand pages), but they reflect an early preoccupation and one that did not leave him throughout his life.[54]
+A first point of Euler's musical theory is the definition of "genres", i.e. of possible divisions of the octave using the prime numbers 3 and 5. Euler describes 18 such genres, with the general definition 2mA, where A is the "exponent" of the genre (i.e. the sum of the exponents of 3 and 5) and 2m (where "m is an indefinite number, small or large, so long as the sounds are perceptible"[55]), expresses that the relation holds independently of the number of octaves concerned. The first genre, with A = 1, is the octave itself (or its duplicates); the second genre, 2m.3, is the octave divided by the fifth (fifth + fourth, C–G–C); the third genre is 2m.5, major third + minor sixth (C–E–C); the fourth is 2m.32, two-fourths and a tone (C–F–B♭–C); the fifth is 2m.3.5 (C–E–G–B–C); etc. Genres 12 (2m.33.5), 13 (2m.32.52) and 14 (2m.3.53) are corrected versions of the diatonic, chromatic and enharmonic, respectively, of the Ancients. Genre 18 (2m.33.52) is the "diatonico-chromatic", "used generally in all compositions",[56] and which turns out to be identical with the system described by Johann Mattheson.[57] Euler later envisaged the possibility of describing genres including the prime number 7.[58]
+Euler devised a specific graph, the Speculum musicum,[59] to illustrate the diatonico-chromatic genre, and discussed paths in this graph for specific intervals, recalling his interest in the Seven Bridges of Königsberg (see above). The device drew renewed interest as the Tonnetz in neo-Riemannian theory (see also Lattice (music)).[60]
+Euler further used the principle of the "exponent" to propose a derivation of the gradus suavitatis (degree of suavity, of agreeableness) of intervals and chords from their prime factors – one must keep in mind that he considered just intonation, i.e. 1 and the prime numbers 3 and 5 only.[61] Formulas have been proposed extending this system to any number of prime numbers, e.g. in the form
+where pi are prime numbers and ki their exponents.[62]
+Euler and his friend Daniel Bernoulli were opponents of Leibniz's monadism and the philosophy of Christian Wolff. Euler insisted that knowledge is founded in part on the basis of precise quantitative laws, something that monadism and Wolffian science were unable to provide. Euler's religious leanings might also have had a bearing on his dislike of the doctrine; he went so far as to label Wolff's ideas as "heathen and atheistic".[63]
+Much of what is known of Euler's religious beliefs can be deduced from his Letters to a German Princess and an earlier work, Rettung der Göttlichen Offenbahrung Gegen die Einwürfe der Freygeister (Defense of the Divine Revelation against the Objections of the Freethinkers). These works show that Euler was a devout Christian who believed the Bible to be inspired; the Rettung was primarily an argument for the divine inspiration of scripture.[64]
+There is a famous legend[65] inspired by Euler's arguments with secular philosophers over religion, which is set during Euler's second stint at the St. Petersburg Academy. The French philosopher Denis Diderot was visiting Russia on Catherine the Great's invitation. However, the Empress was alarmed that the philosopher's arguments for atheism were influencing members of her court, and so Euler was asked to confront the Frenchman. Diderot was informed that a learned mathematician had produced a proof of the existence of God: he agreed to view the proof as it was presented in court. Euler appeared, advanced toward Diderot, and in a tone of perfect conviction announced this non-sequitur: "Sir, a+bn/n=x, hence God exists—reply!"
+Diderot, to whom (says the story) all mathematics was gibberish, stood dumbstruck as peals of laughter erupted from the court.  Embarrassed, he asked to leave Russia, a request that was graciously granted by the Empress. However amusing the anecdote may be, it is apocryphal, given that Diderot himself did research in mathematics.[66]
+The legend was apparently first told by
+Dieudonné Thiébault[67] with embellishment by Augustus De Morgan.[68][69]
+Euler was featured on the sixth series of the Swiss 10-franc banknote and on numerous Swiss, German, and Russian postage stamps. The asteroid 2002 Euler was named in his honor. He is also commemorated by the Lutheran Church on their Calendar of Saints on 24 May—he was a devout Christian (and believer in biblical inerrancy) who wrote apologetics and argued forcefully against the prominent atheists of his time.[64]
+Euler has an extensive bibliography. His best-known books include:
+The first collection of Euler's work was made by Paul Heinrich von Fuss in 1862.[71] A definitive collection of Euler's works, entitled Opera Omnia, has been published since 1911 by the Euler Commission of the Swiss Academy of Sciences. A complete chronological list of Euler's works is available at The Eneström Index.[72] Full text, open access versions of many of Euler's papers are available in the original language and English translations at the Euler Archive, hosted by University of the Pacific. The Euler Archive was started at Dartmouth College[73] before moving to the Mathematical Association of America[74] and, most recently, to University of the Pacific in 2017.
+Illustration from Solutio problematis... a. 1743 propositi published in Acta Eruditorum, 1744
+The title page of Euler's Methodus inveniendi lineas curvas.

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+P. p. japonensis (Gray, 1862)
+The Amur leopard (Panthera pardus orientalis) is a leopard subspecies native to the Primorye region of southeastern Russia and northern China. It is listed as Critically Endangered on the IUCN Red List. In 2007, only 19–26 wild leopards were estimated to survive in southeastern Russia and northeastern China.[1] It was considered as one of the rarest cats on Earth.[3]
+As of 2015[update], fewer than 60 individuals were estimated to survive in Russia and China.[4] Camera-trapping surveys conducted between 2014 and 2015 revealed 92 individuals in an 8,398 km2 (3,242 sq mi) large transboundary area along the Russian-Chinese border.[5]
+In 2019, it was reported that the population is close to 90 leopards.[6]
+Results of genetic research indicate that the Amur leopard is genetically close to leopards in northern China and Korea, suggesting that the leopard population in this region became fragmented in the early 20th century.[7] The North Chinese leopard was formerly recognised as a distinct subspecies P. p. japonensis, but was subsumed under the Amur leopard in 2017.[2]
+In 1857, Hermann Schlegel described a leopard skin from Korea under the scientific name Felis orientalis.[8]
+Since Schlegel's description, several naturalists and curators of natural history museums described zoological specimens of leopards from the Russian Far East and China:
+In 2017, the Cat Classification Task Force of the Cat Specialist Group subsumed P. p. japonensis to P. p. orientalis. The remaining are not considered valid subspecies.[2]
+Phylogenetic analysis of leopard samples from Primorsky Krai and North Korea revealed that they cannot be distinguished. It is considered probable that the population became fragmented less than a century ago.[7]
+Phylogenetic analysis of an old leopard skin from South Korea indicates that it belongs to the Amur leopard subspecies.[17]
+The complete mitochondrial genome of a wild male leopard specimen from Shaanxi Province in central China has been amplified and is 16,966 base pairs long.[18]
+The Amur leopard differs from other leopard subspecies by its thick fur that is pale cream-colored, particularly in winter. Rosettes on the flanks are 5 cm × 5 cm (2.0 in × 2.0 in) and widely spaced, up to 2.5 cm (0.98 in), with thick, unbroken rings and darkened centers.[16] Its fur is fairly soft with long and dense hair. The length of hair on the back is 20–25 mm (0.79–0.98 in) in summer and up to 70 mm (2.8 in) in winter. The winter coat varies from fairly light yellow to dense yellowish-red with a golden tinge or rusty-reddish-yellow. In summer, the fur is brighter with more vivid coloration pattern. It is rather small in body size, with males larger than females. Males measure from 107–136 cm (42–54 in) with an 82–90 cm (32–35 in) long tail, a shoulder height of 64–78 cm (25–31 in), and a weight of 32.2–48 kg (71–106 lb). Females weigh from 25–42.5 kg (55–94 lb).[19]
+The North Chinese leopard was first described on the basis of a single tanned skin which was fulvous above, and pale beneath, with large, roundish, oblong black spots on the back and limbs, and small black spots on the head. The spots on the back, shoulders and sides formed a ring around a central fulvous spot. The black spots on the nape were elongated, and large ones on the chest formed a necklace. The tail was spotted and had four black rings at the tip.[9]
+In the Russian Far East, the Amur leopard inhabits an area of about 7,000 km2 (2,700 sq mi) today.[20]
+It is well adapted to a cold climate and snow.[7]
+Leopards cross between Russia, China, and North Korea across the Tumen River despite a high and long wire fence marking the boundary.[21]
+The first camera trap image of an Amur leopard in northeastern China was taken in 2010 in Hunchun National Nature Reserve located in the Changbai Mountains in Jilin and Heilongjiang Provinces.[22]
+This habitat consists of broadleaved conifer and Korean pine forests at altitudes of 600–1,200 m (2,000–3,900 ft), where annual average temperature is about 1.5 °C (34.7 °F).[23]
+In this area, leopards were repeatedly photographed by camera traps set up between January 2013 and July 2014 covering up to 4,858 km2 (1,876 sq mi).[24][25][26]
+Elsewhere in China, leopard range is fragmented with small populations occurring foremost in isolated reserves. In Shanxi Province, leopards were recorded in 16 protected areas during camera trapping surveys between 2007 and 2014. In Shaanxi Province, leopards were recorded in six nature reserves, including Foping National Nature Reserve.[27]
+Leopard fossils from the Pleistocene have been excavated in Japan, but the species has not been identified with certainty.[28]
+Historic records from before 1930 indicate that the Amur leopard occurred in Eastern Siberia, in Northern China near Beijing and in the mountains to the northwest, and in Korea.[10][29]
+Its range once extended throughout Manchuria in northeastern China, including Jilin and Heilongjiang provinces, and throughout the Korean Peninsula. In Russia, its range was dramatically reduced during the 1970s to about 20% of its former range. The northern boundary of its occurrence commenced on the coast of the Sea of Japan at 44°N and ran south at a distance of 15–30 km (9.3–18.6 mi) from the coast to 43°10'N. There its range turned steeply westward, north of the Suchan River basin, then north to encompass the source of the Ussuri River and two right bank tributaries in the upper reaches of the Ussuri, and westward toward the bank of Khanka Lake. In the 1950s, leopards were observed 50 km (31 mi) north of Vladivostok and in Kedrovaya Pad Nature Reserve. The association of the leopard with mountains is fairly definite, and to snow-free south-facing rocky slopes in winter. The species is confined more to places where wild sika deer live or where deer husbandry is practiced.[19]
+Leopards were extirpated on the Korean Peninsula during the period of Korea under Japanese rule.[30] At least 624 leopards were killed during the Japanese occupation between 1910 and 1945. In South Korea, the last known leopard was captured in 1970.[17] The Amur leopard is considered locally extinct.[1]
+In China, Amur leopards occurred in the Lesser Khingan, Changbai Mountains and Wanda Mountains until the 1970s. In the following decades, the range decreased to a few areas in Jilin and Heilongjiang Provinces.[31]
+Today, only small and isolated populations remain in China.[18]
+Amur leopards are solitary, unless females have offspring.[19] Records from camera-traps indicate that they are more active during the day than at night and during  twilight, both in the summer and winter seasons. This activity pattern coincides with activity of prey species such as Siberian roe deer, sika deer and wild boar.[26]
+They are extremely conservative in their choice of territory. An individual's territory is usually located in a river basin which generally extends to the natural topographical borders of the area. The territory of two individuals overlaps sometimes, but only slightly. Depending on sex, age and family size, the size of an individual's territory varies from 5,000–30,000 ha (19–116 sq mi). They use the same hunting trails, migration routes and even rest places over the course of many years.[32]
+Leopards are resident at places where wild animals are abundant, and follow herds of ungulates. In the Ussuri region the main prey of leopards are Siberian roe deer (Capreolus pygargus) and Manchurian sika deer (Cervus nippon mantchuricus), Manchurian wapiti (Cervus canadensis xanthopygus), Siberian musk deer (Moschus moschiferus), Amur moose (Alces alces cameloides) and Ussuri wild boar (Sus scrofa ussuricus). They also catch hare (Lepus), Eurasian badger (Meles meles), fowl, and mice. In Kedrovaya Pad Nature Reserve roe deer is their main prey year-round, but they also prey on young Asian black bear if they are less than two years old.[19]
+When density of ungulates is low, leopards have large home ranges of up to 100 km2 (39 sq mi).[33]
+During a study of radio-collared Amur leopards in the early 1990s, a territorial dispute between two males at a deer farm was documented, suggesting that Amur leopards favour such farms for hunting.[34] Female leopards with cubs are often found in the proximity of deer farms. The large number of domesticated deer is a reliable food source in difficult times.[35]
+In 2011, an adult Amur leopard female was radio-collared in the vicinity of the Land of the Leopard National Park in the Khasansky region of Primorskii Krai. During three years of tracking, she used a home range of 161.7 km2 (62.4 sq mi) with a core area of 23.3 km2 (9.0 sq mi). During estrus, she moved in a core area of 52.9 km2 (20.4 sq mi). After giving birth in late June, she reduced her movements to an area of about 3 km2 (1.2 sq mi) for a month, in which she shifted her cubs three times. From autumn onwards, she gradually increased her home range. When the cubs were more than one year old, the family moved together in the initial home range of 161.7 km2 (62.4 sq mi).[36]
+Amur leopards share habitat with Siberian tigers, but in the Changbai Mountains have been recorded more often in higher altitudes and far from settlements and roads than tigers.[26]
+Amur leopards become sexually mature at the age of 2–3 years. They are able to reproduce up to 10–15 years of age. Estrus lasts 12–18 days, and in exceptional cases up to 25 days. Gestation lasts 90–105 days, and usually between 92 and 95 days. A newborn cub weighs 500–700 g (1.1–1.5 lb). The young open their eyes on the 7th–10th day and begin to crawl on the 12th–15th day. By the second month they emerge from their dens and also begin to eat meat. Cubs are weaned when three months old,  and then learn to hunt. Lactation continues for five or six months. Cubs reach independence at the approximate age of two to three years. They stay with their mother until they are around 18 months to two years old.[36] Juveniles sometimes stay with their mother until she comes into estrus again. Until the 1970s, cubs were seen in Kedrovaya Pad Nature Reserve and in northeastern China most often between the end of March and May. Litters comprised two to three cubs. In captivity some individuals have lived for 21 years.[19]
+During a population census in 1997, four females found with young had only one cub each. Results of radio telemetry studies confirmed that young stay with their mother for two years. In Kedrovaya Pad Nature Reserve, the young of two different litters were observed with their mothers at the same time.[32]
+The Amur leopard is threatened by poaching, poaching of prey species, habitat loss and deforestation or exploitation of forests. Its natural habitat is threatened by forest fires and construction of new roads.[1]
+Due to the small number of reproducing Amur leopards in the wild, the gene pool is so reduced that the population is at risk from inbreeding depression.[37]
+In 2015, a wild Amur leopard was found with canine distemper virus in Primorskii Krai. The small population is possibly exposed to domestic or wild disease carriers and transmitters.[38]
+Tigers can eliminate leopards if densities of large and medium-sized prey species are low. Competition between these predators supposedly decreases in summer, when small prey species are more available. In winter, conditions are less favorable for tigers and the extent of trophic niche overlap with that of leopards probably reaches its peak.[33]
+Poaching of leopards is a main threat to their survival. There are rumours but no evidence that Chinese traders buy leopard skins; no skins were confiscated at borders to China. In 14 months from February 2002 to April 2003, seven skins or part of skins were confiscated, six in Russia and one in China. Leopards are most often killed by local Russians from small villages in and around the leopard habitat. Most of these villagers hunt entirely illegally; they have no licenses for hunting nor for their guns, and they are not members of one of the local hunting leases.[35]
+In 1999, skins of poached leopards were offered for $500–1,000 near a protected area in Russia.[39]
+Human induced fires are a main threat to the survival of the leopard. Setting fire to fields is a habit of rural farmers who start them for a particular purpose such as improving fertility for livestock grazing, killing ticks and other insects, making scrap metals visible so that they can be easily collected, culling vegetation along train tracks, and stimulating fern growth. Young ferns are sold in shops, served in restaurants and also exported to China as a popular dish. Surveys using satellite images and GIS techniques revealed that on average 19% of south-west Primorye burns annually, and a total of 46% burned at least once in six years. Due to a long and frequent fire history, much of the land in south-west Primorye has been converted to permanent grasslands. These frequent fires cause degradation of suitable leopard habitat into unsuitable habitat. Repeated fires have created open "savannah" landscapes with grass, oak bushes and isolated trees that leopards seem to avoid, again probably because of low ungulate densities.[35]
+Large deer farms extended over thousands of hectares in leopard habitat. Deer farming used to be a large-scale business; the velvet of deer antlers was sold to Asian pharmacies.[34] The number of deer farms decreased considerably since the late 1990s.[35]
+A number of plans for economic activities in south-west Primorye were developed that posed a serious threat to the leopard's survival. A plan to build an oil pipeline from central Siberia through Primorye to the coast of the Sea of Japan has been shelved. A plan for an open pit coal mine in the heart of the leopard range was not carried out following pressure from environmentalists and the Ministry of Natural Resources. The strategic location of south-west Primorye, close to the main population centres of Primorsky Krai, the Japanese Sea and the borders of Korea and China, makes it more attractive for economic activities including transport, industries, tourism and development of infrastructure. Logging is not a major threat; the use of the road network established for the transport of logs from forests increases anthropogenic pressures in unprotected leopard habitat.[35]
+An acute problem is potential inbreeding. The remaining population could disappear as a result of genetic degeneration, even without direct human influence. The levels of diversity are remarkably low, indicative of a history of inbreeding in the population for several generations. Such levels of genetic reduction have been associated with severe reproductive and congenital abnormalities that impede the health, survival and reproduction of some but not all genetically diminished small populations. Cub survival has been declining from 1.9 cubs per one female in 1973 to 1.7 in 1984 and 1.0 in 1991. Besides a decline in natural replacement, there is a high probability of mortality for all age groups as a result of certain diseases or direct human impact.[40]
+Results of genetic analyses imply that the Amur leopard population lost genetic diversity over a short period of time.[7]
+The Amur leopard is listed in CITES Appendix I. It is stated to need better protection from illegal trade in skins and bones.[1]
+In 2001, a meeting was held in Vladivostok with the aim of devising and planning management recommendations and activities needed to ensure the recovery and continued survival of the wild Amur leopard population in range countries. Chinese participants announced the creation of a new protected area in Jilin Province, the Hunchun Nature Reserve.[41]
+Since 2014, Russian and Chinese biologists collaborate in transboundary monitoring of the Amur leopard population.[5]
+The Amur Leopard and Tiger Alliance (ALTA) is an initiative of Russian and western conservation organisations to conserve the Amur leopard and tiger, and secure a future for both species in the Russian Far East and Northeast China. ALTA operates across Northeast Asia under the guiding principle that only co-operative, co-ordinated conservation actions from all interested parties can save these endangered species from extinction. ALTA works in close co-operation with local, regional, and federal government and non-government organisations to protect the region's biological wealth through conservation, sustainable development, and local community involvement. The Phoenix Fund and the Wildlife Conservation Society provide a local framework for implementing ALTA projects, working closely with many Russian and Chinese agencies. With regard to conservation of leopards, ALTA aims at retaining a leopard population of 35 adult females (100 total) in south-west Primorye and the Jilin-Heilongjiang border region; and creating a second population of 20 adult females (60 total) in the former range of the leopard. Conservation projects for the leopard include:[35]
+An oil pipeline planned to be built through leopard habitat was rerouted, following a campaign by conservationists.[42]
+Since 1996, the idea of reintroducing leopards in the south of Sikhote-Alin has been discussed by ALTA members.[43] During a workshop in 2001, the outlines and principles of a plan for the development of a second population of the leopard in the Russian Far East was prepared. For reintroduction to be successful, one fundamental question needs to be answered: Why did leopards disappear from the southern Sikhote-Alin in the middle of the 20th century? It was recommended to assess reasons for localized extinctions, obtain support of local people, increase prey in areas proposed for reintroduction, ensure that conditions exist conducive for reintroduction in the selected area, and ensure survival of the existing population. There are two sources of leopards for reintroduction: leopards born and raised in zoos and leopards raised in a special reintroduction center passed through a rehabilitation program for life in the wild.[40]
+If this reintroduction is to succeed, it is clear that the design of the breeding and release centre, and the management of the leopards in it, must focus strongly on overcoming the difficulties imposed by the captive origin of the cats. Three necessary behaviours should be acquired prior to release: hunting and killing of live natural prey; avoidance of humans and avoidance of tigers.[44]
+In March 2009, the Minister of Natural Resources of Russia during his meeting with Vladimir Putin reassured that the ministry is planning to introduce new "imported" leopards into the area and creating suitable and safe habitat for them. The government already allocated all required funds for the project.[45]
+Contiguous patches of potential leopard habitat as potential reintroduction sites were identified in the southern Sikhote-Alin. Three coastal potential habitat patches could harbour approximately 72 adult leopards.[20]
+A captive breeding programme for the Amur leopard was established in 1961 from nine wild-born founders.[46] A molecular genetic survey revealed that at least two founders of the captive pedigree had genetic information that is not consistent with any wildborn Amur leopard.[37]
+Both the American and European zoo populations include contribution of genes from a male founder that was not an Amur leopard. It has been the strategy of the European Endangered Species Programme to minimize his contribution and maintain genetic diversity of the captive population.[44]
+As of December 2011[update], 173 captive Amur leopards were held in zoos worldwide. Within the EESP, 54 male, 40 female and 7 unsexed individuals are kept. In American and Canadian zoos, another 31 males and 41 females are kept within the Population Management Program.[47]
+The names 'Amurland leopard' and 'Amur leopard' were coined by Pocock in 1930, when he compared leopard specimens in the collection of the Natural History Museum, London. In particular, he referred to a leopard skin from the Amur Bay as 'Amur leopard'.[16] Since at least 1985, this name has been used for the leopard population in Eastern Siberia and for the captive population in zoos worldwide.[48][46]
+The Amur leopard is also known as the "Siberian leopard",[49] "Far Eastern leopard",[50][37][43] and "Korean leopard".[17]
+The Amur leopard were depicted in many Korean folk paintings in Joseon Dynasty.
+The Animal Planet documentary The Last Leopard (2008) is about the plight of Amur leopards in Russia. The television series "Wild Russia" showed a glimpse into the life of leopards. A female leopard and her cub were featured on Planet Earth episodes "Seasonal Forests".[3]
+Leopard subspecies: African leopard  · Arabian leopard  · Anatolian leopard  · Persian leopard  · Indian leopard  · Indochinese leopard  · Javan leopard  · Sri Lankan leopard  · Panthera pardus spelaea

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+P. p. japonensis (Gray, 1862)
+The Amur leopard (Panthera pardus orientalis) is a leopard subspecies native to the Primorye region of southeastern Russia and northern China. It is listed as Critically Endangered on the IUCN Red List. In 2007, only 19–26 wild leopards were estimated to survive in southeastern Russia and northeastern China.[1] It was considered as one of the rarest cats on Earth.[3]
+As of 2015[update], fewer than 60 individuals were estimated to survive in Russia and China.[4] Camera-trapping surveys conducted between 2014 and 2015 revealed 92 individuals in an 8,398 km2 (3,242 sq mi) large transboundary area along the Russian-Chinese border.[5]
+In 2019, it was reported that the population is close to 90 leopards.[6]
+Results of genetic research indicate that the Amur leopard is genetically close to leopards in northern China and Korea, suggesting that the leopard population in this region became fragmented in the early 20th century.[7] The North Chinese leopard was formerly recognised as a distinct subspecies P. p. japonensis, but was subsumed under the Amur leopard in 2017.[2]
+In 1857, Hermann Schlegel described a leopard skin from Korea under the scientific name Felis orientalis.[8]
+Since Schlegel's description, several naturalists and curators of natural history museums described zoological specimens of leopards from the Russian Far East and China:
+In 2017, the Cat Classification Task Force of the Cat Specialist Group subsumed P. p. japonensis to P. p. orientalis. The remaining are not considered valid subspecies.[2]
+Phylogenetic analysis of leopard samples from Primorsky Krai and North Korea revealed that they cannot be distinguished. It is considered probable that the population became fragmented less than a century ago.[7]
+Phylogenetic analysis of an old leopard skin from South Korea indicates that it belongs to the Amur leopard subspecies.[17]
+The complete mitochondrial genome of a wild male leopard specimen from Shaanxi Province in central China has been amplified and is 16,966 base pairs long.[18]
+The Amur leopard differs from other leopard subspecies by its thick fur that is pale cream-colored, particularly in winter. Rosettes on the flanks are 5 cm × 5 cm (2.0 in × 2.0 in) and widely spaced, up to 2.5 cm (0.98 in), with thick, unbroken rings and darkened centers.[16] Its fur is fairly soft with long and dense hair. The length of hair on the back is 20–25 mm (0.79–0.98 in) in summer and up to 70 mm (2.8 in) in winter. The winter coat varies from fairly light yellow to dense yellowish-red with a golden tinge or rusty-reddish-yellow. In summer, the fur is brighter with more vivid coloration pattern. It is rather small in body size, with males larger than females. Males measure from 107–136 cm (42–54 in) with an 82–90 cm (32–35 in) long tail, a shoulder height of 64–78 cm (25–31 in), and a weight of 32.2–48 kg (71–106 lb). Females weigh from 25–42.5 kg (55–94 lb).[19]
+The North Chinese leopard was first described on the basis of a single tanned skin which was fulvous above, and pale beneath, with large, roundish, oblong black spots on the back and limbs, and small black spots on the head. The spots on the back, shoulders and sides formed a ring around a central fulvous spot. The black spots on the nape were elongated, and large ones on the chest formed a necklace. The tail was spotted and had four black rings at the tip.[9]
+In the Russian Far East, the Amur leopard inhabits an area of about 7,000 km2 (2,700 sq mi) today.[20]
+It is well adapted to a cold climate and snow.[7]
+Leopards cross between Russia, China, and North Korea across the Tumen River despite a high and long wire fence marking the boundary.[21]
+The first camera trap image of an Amur leopard in northeastern China was taken in 2010 in Hunchun National Nature Reserve located in the Changbai Mountains in Jilin and Heilongjiang Provinces.[22]
+This habitat consists of broadleaved conifer and Korean pine forests at altitudes of 600–1,200 m (2,000–3,900 ft), where annual average temperature is about 1.5 °C (34.7 °F).[23]
+In this area, leopards were repeatedly photographed by camera traps set up between January 2013 and July 2014 covering up to 4,858 km2 (1,876 sq mi).[24][25][26]
+Elsewhere in China, leopard range is fragmented with small populations occurring foremost in isolated reserves. In Shanxi Province, leopards were recorded in 16 protected areas during camera trapping surveys between 2007 and 2014. In Shaanxi Province, leopards were recorded in six nature reserves, including Foping National Nature Reserve.[27]
+Leopard fossils from the Pleistocene have been excavated in Japan, but the species has not been identified with certainty.[28]
+Historic records from before 1930 indicate that the Amur leopard occurred in Eastern Siberia, in Northern China near Beijing and in the mountains to the northwest, and in Korea.[10][29]
+Its range once extended throughout Manchuria in northeastern China, including Jilin and Heilongjiang provinces, and throughout the Korean Peninsula. In Russia, its range was dramatically reduced during the 1970s to about 20% of its former range. The northern boundary of its occurrence commenced on the coast of the Sea of Japan at 44°N and ran south at a distance of 15–30 km (9.3–18.6 mi) from the coast to 43°10'N. There its range turned steeply westward, north of the Suchan River basin, then north to encompass the source of the Ussuri River and two right bank tributaries in the upper reaches of the Ussuri, and westward toward the bank of Khanka Lake. In the 1950s, leopards were observed 50 km (31 mi) north of Vladivostok and in Kedrovaya Pad Nature Reserve. The association of the leopard with mountains is fairly definite, and to snow-free south-facing rocky slopes in winter. The species is confined more to places where wild sika deer live or where deer husbandry is practiced.[19]
+Leopards were extirpated on the Korean Peninsula during the period of Korea under Japanese rule.[30] At least 624 leopards were killed during the Japanese occupation between 1910 and 1945. In South Korea, the last known leopard was captured in 1970.[17] The Amur leopard is considered locally extinct.[1]
+In China, Amur leopards occurred in the Lesser Khingan, Changbai Mountains and Wanda Mountains until the 1970s. In the following decades, the range decreased to a few areas in Jilin and Heilongjiang Provinces.[31]
+Today, only small and isolated populations remain in China.[18]
+Amur leopards are solitary, unless females have offspring.[19] Records from camera-traps indicate that they are more active during the day than at night and during  twilight, both in the summer and winter seasons. This activity pattern coincides with activity of prey species such as Siberian roe deer, sika deer and wild boar.[26]
+They are extremely conservative in their choice of territory. An individual's territory is usually located in a river basin which generally extends to the natural topographical borders of the area. The territory of two individuals overlaps sometimes, but only slightly. Depending on sex, age and family size, the size of an individual's territory varies from 5,000–30,000 ha (19–116 sq mi). They use the same hunting trails, migration routes and even rest places over the course of many years.[32]
+Leopards are resident at places where wild animals are abundant, and follow herds of ungulates. In the Ussuri region the main prey of leopards are Siberian roe deer (Capreolus pygargus) and Manchurian sika deer (Cervus nippon mantchuricus), Manchurian wapiti (Cervus canadensis xanthopygus), Siberian musk deer (Moschus moschiferus), Amur moose (Alces alces cameloides) and Ussuri wild boar (Sus scrofa ussuricus). They also catch hare (Lepus), Eurasian badger (Meles meles), fowl, and mice. In Kedrovaya Pad Nature Reserve roe deer is their main prey year-round, but they also prey on young Asian black bear if they are less than two years old.[19]
+When density of ungulates is low, leopards have large home ranges of up to 100 km2 (39 sq mi).[33]
+During a study of radio-collared Amur leopards in the early 1990s, a territorial dispute between two males at a deer farm was documented, suggesting that Amur leopards favour such farms for hunting.[34] Female leopards with cubs are often found in the proximity of deer farms. The large number of domesticated deer is a reliable food source in difficult times.[35]
+In 2011, an adult Amur leopard female was radio-collared in the vicinity of the Land of the Leopard National Park in the Khasansky region of Primorskii Krai. During three years of tracking, she used a home range of 161.7 km2 (62.4 sq mi) with a core area of 23.3 km2 (9.0 sq mi). During estrus, she moved in a core area of 52.9 km2 (20.4 sq mi). After giving birth in late June, she reduced her movements to an area of about 3 km2 (1.2 sq mi) for a month, in which she shifted her cubs three times. From autumn onwards, she gradually increased her home range. When the cubs were more than one year old, the family moved together in the initial home range of 161.7 km2 (62.4 sq mi).[36]
+Amur leopards share habitat with Siberian tigers, but in the Changbai Mountains have been recorded more often in higher altitudes and far from settlements and roads than tigers.[26]
+Amur leopards become sexually mature at the age of 2–3 years. They are able to reproduce up to 10–15 years of age. Estrus lasts 12–18 days, and in exceptional cases up to 25 days. Gestation lasts 90–105 days, and usually between 92 and 95 days. A newborn cub weighs 500–700 g (1.1–1.5 lb). The young open their eyes on the 7th–10th day and begin to crawl on the 12th–15th day. By the second month they emerge from their dens and also begin to eat meat. Cubs are weaned when three months old,  and then learn to hunt. Lactation continues for five or six months. Cubs reach independence at the approximate age of two to three years. They stay with their mother until they are around 18 months to two years old.[36] Juveniles sometimes stay with their mother until she comes into estrus again. Until the 1970s, cubs were seen in Kedrovaya Pad Nature Reserve and in northeastern China most often between the end of March and May. Litters comprised two to three cubs. In captivity some individuals have lived for 21 years.[19]
+During a population census in 1997, four females found with young had only one cub each. Results of radio telemetry studies confirmed that young stay with their mother for two years. In Kedrovaya Pad Nature Reserve, the young of two different litters were observed with their mothers at the same time.[32]
+The Amur leopard is threatened by poaching, poaching of prey species, habitat loss and deforestation or exploitation of forests. Its natural habitat is threatened by forest fires and construction of new roads.[1]
+Due to the small number of reproducing Amur leopards in the wild, the gene pool is so reduced that the population is at risk from inbreeding depression.[37]
+In 2015, a wild Amur leopard was found with canine distemper virus in Primorskii Krai. The small population is possibly exposed to domestic or wild disease carriers and transmitters.[38]
+Tigers can eliminate leopards if densities of large and medium-sized prey species are low. Competition between these predators supposedly decreases in summer, when small prey species are more available. In winter, conditions are less favorable for tigers and the extent of trophic niche overlap with that of leopards probably reaches its peak.[33]
+Poaching of leopards is a main threat to their survival. There are rumours but no evidence that Chinese traders buy leopard skins; no skins were confiscated at borders to China. In 14 months from February 2002 to April 2003, seven skins or part of skins were confiscated, six in Russia and one in China. Leopards are most often killed by local Russians from small villages in and around the leopard habitat. Most of these villagers hunt entirely illegally; they have no licenses for hunting nor for their guns, and they are not members of one of the local hunting leases.[35]
+In 1999, skins of poached leopards were offered for $500–1,000 near a protected area in Russia.[39]
+Human induced fires are a main threat to the survival of the leopard. Setting fire to fields is a habit of rural farmers who start them for a particular purpose such as improving fertility for livestock grazing, killing ticks and other insects, making scrap metals visible so that they can be easily collected, culling vegetation along train tracks, and stimulating fern growth. Young ferns are sold in shops, served in restaurants and also exported to China as a popular dish. Surveys using satellite images and GIS techniques revealed that on average 19% of south-west Primorye burns annually, and a total of 46% burned at least once in six years. Due to a long and frequent fire history, much of the land in south-west Primorye has been converted to permanent grasslands. These frequent fires cause degradation of suitable leopard habitat into unsuitable habitat. Repeated fires have created open "savannah" landscapes with grass, oak bushes and isolated trees that leopards seem to avoid, again probably because of low ungulate densities.[35]
+Large deer farms extended over thousands of hectares in leopard habitat. Deer farming used to be a large-scale business; the velvet of deer antlers was sold to Asian pharmacies.[34] The number of deer farms decreased considerably since the late 1990s.[35]
+A number of plans for economic activities in south-west Primorye were developed that posed a serious threat to the leopard's survival. A plan to build an oil pipeline from central Siberia through Primorye to the coast of the Sea of Japan has been shelved. A plan for an open pit coal mine in the heart of the leopard range was not carried out following pressure from environmentalists and the Ministry of Natural Resources. The strategic location of south-west Primorye, close to the main population centres of Primorsky Krai, the Japanese Sea and the borders of Korea and China, makes it more attractive for economic activities including transport, industries, tourism and development of infrastructure. Logging is not a major threat; the use of the road network established for the transport of logs from forests increases anthropogenic pressures in unprotected leopard habitat.[35]
+An acute problem is potential inbreeding. The remaining population could disappear as a result of genetic degeneration, even without direct human influence. The levels of diversity are remarkably low, indicative of a history of inbreeding in the population for several generations. Such levels of genetic reduction have been associated with severe reproductive and congenital abnormalities that impede the health, survival and reproduction of some but not all genetically diminished small populations. Cub survival has been declining from 1.9 cubs per one female in 1973 to 1.7 in 1984 and 1.0 in 1991. Besides a decline in natural replacement, there is a high probability of mortality for all age groups as a result of certain diseases or direct human impact.[40]
+Results of genetic analyses imply that the Amur leopard population lost genetic diversity over a short period of time.[7]
+The Amur leopard is listed in CITES Appendix I. It is stated to need better protection from illegal trade in skins and bones.[1]
+In 2001, a meeting was held in Vladivostok with the aim of devising and planning management recommendations and activities needed to ensure the recovery and continued survival of the wild Amur leopard population in range countries. Chinese participants announced the creation of a new protected area in Jilin Province, the Hunchun Nature Reserve.[41]
+Since 2014, Russian and Chinese biologists collaborate in transboundary monitoring of the Amur leopard population.[5]
+The Amur Leopard and Tiger Alliance (ALTA) is an initiative of Russian and western conservation organisations to conserve the Amur leopard and tiger, and secure a future for both species in the Russian Far East and Northeast China. ALTA operates across Northeast Asia under the guiding principle that only co-operative, co-ordinated conservation actions from all interested parties can save these endangered species from extinction. ALTA works in close co-operation with local, regional, and federal government and non-government organisations to protect the region's biological wealth through conservation, sustainable development, and local community involvement. The Phoenix Fund and the Wildlife Conservation Society provide a local framework for implementing ALTA projects, working closely with many Russian and Chinese agencies. With regard to conservation of leopards, ALTA aims at retaining a leopard population of 35 adult females (100 total) in south-west Primorye and the Jilin-Heilongjiang border region; and creating a second population of 20 adult females (60 total) in the former range of the leopard. Conservation projects for the leopard include:[35]
+An oil pipeline planned to be built through leopard habitat was rerouted, following a campaign by conservationists.[42]
+Since 1996, the idea of reintroducing leopards in the south of Sikhote-Alin has been discussed by ALTA members.[43] During a workshop in 2001, the outlines and principles of a plan for the development of a second population of the leopard in the Russian Far East was prepared. For reintroduction to be successful, one fundamental question needs to be answered: Why did leopards disappear from the southern Sikhote-Alin in the middle of the 20th century? It was recommended to assess reasons for localized extinctions, obtain support of local people, increase prey in areas proposed for reintroduction, ensure that conditions exist conducive for reintroduction in the selected area, and ensure survival of the existing population. There are two sources of leopards for reintroduction: leopards born and raised in zoos and leopards raised in a special reintroduction center passed through a rehabilitation program for life in the wild.[40]
+If this reintroduction is to succeed, it is clear that the design of the breeding and release centre, and the management of the leopards in it, must focus strongly on overcoming the difficulties imposed by the captive origin of the cats. Three necessary behaviours should be acquired prior to release: hunting and killing of live natural prey; avoidance of humans and avoidance of tigers.[44]
+In March 2009, the Minister of Natural Resources of Russia during his meeting with Vladimir Putin reassured that the ministry is planning to introduce new "imported" leopards into the area and creating suitable and safe habitat for them. The government already allocated all required funds for the project.[45]
+Contiguous patches of potential leopard habitat as potential reintroduction sites were identified in the southern Sikhote-Alin. Three coastal potential habitat patches could harbour approximately 72 adult leopards.[20]
+A captive breeding programme for the Amur leopard was established in 1961 from nine wild-born founders.[46] A molecular genetic survey revealed that at least two founders of the captive pedigree had genetic information that is not consistent with any wildborn Amur leopard.[37]
+Both the American and European zoo populations include contribution of genes from a male founder that was not an Amur leopard. It has been the strategy of the European Endangered Species Programme to minimize his contribution and maintain genetic diversity of the captive population.[44]
+As of December 2011[update], 173 captive Amur leopards were held in zoos worldwide. Within the EESP, 54 male, 40 female and 7 unsexed individuals are kept. In American and Canadian zoos, another 31 males and 41 females are kept within the Population Management Program.[47]
+The names 'Amurland leopard' and 'Amur leopard' were coined by Pocock in 1930, when he compared leopard specimens in the collection of the Natural History Museum, London. In particular, he referred to a leopard skin from the Amur Bay as 'Amur leopard'.[16] Since at least 1985, this name has been used for the leopard population in Eastern Siberia and for the captive population in zoos worldwide.[48][46]
+The Amur leopard is also known as the "Siberian leopard",[49] "Far Eastern leopard",[50][37][43] and "Korean leopard".[17]
+The Amur leopard were depicted in many Korean folk paintings in Joseon Dynasty.
+The Animal Planet documentary The Last Leopard (2008) is about the plight of Amur leopards in Russia. The television series "Wild Russia" showed a glimpse into the life of leopards. A female leopard and her cub were featured on Planet Earth episodes "Seasonal Forests".[3]
+Leopard subspecies: African leopard  · Arabian leopard  · Anatolian leopard  · Persian leopard  · Indian leopard  · Indochinese leopard  · Javan leopard  · Sri Lankan leopard  · Panthera pardus spelaea

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+A. f. fulgens F. Cuvier, 1825
+A. f. styani Thomas, 1902[2]
+The red panda (Ailurus fulgens) is a mammal species native to the eastern Himalayas and southwestern China. It is listed as Endangered on the IUCN Red List because the wild population is estimated at fewer than 10,000 mature individuals and continues to decline due to habitat loss and fragmentation, poaching, and inbreeding depression.[1] Despite its name, it is not closely related to the giant panda.[3]
+The red panda has reddish-brown fur, a long, shaggy tail, and a waddling gait due to its shorter front legs; it is roughly the size of a domestic cat, though with a longer body, and is somewhat heavier. It is arboreal and feeds mainly on bamboo, but also eats eggs, birds, and insects. It is a solitary animal, mainly active from dusk to dawn, and is largely sedentary during the day. It is also called the lesser panda, the red bear-cat, and the red cat-bear.[4]
+The red panda is the only living member of the genus Ailurus and the family Ailuridae. It has previously been placed in the raccoon and bear families, but the results of phylogenetic analysis provide strong support for its taxonomic classification in its own family, Ailuridae, which is part of the superfamily Musteloidea, along with the weasel, raccoon and skunk families.[5] Traditionally it was thought to consist of two subspecies.[6] However, results of genetic analysis indicate that there are probably two distinct red panda species, the Chinese red panda and the Himalayan red panda, which genetically diverged 0.22 million years ago.[7]
+The red panda has long, soft, reddish-brown fur on the upper parts, blackish fur on the lower parts, and a light face with tear markings and white badges similar to those of a raccoon, but each individual can have distinctive markings. Its skull is roundish with medium-sized upright ears, its nose is black, and its eyes are blackish. Its teeth are robust. Its long, bushy tail with six alternating transverse ochre rings provide balance and excellent camouflage in a habitat with moss- and lichen-covered trees. The legs are black and short with thick fur on the soles of the paws. This fur serves as thermal insulation on snow-covered or icy surfaces and conceals scent glands, which are also present on the anus.[8]
+The head and body length of a red panda measures 50 to 64 cm (20 to 25 in), and its tail is 28 to 59 cm (11 to 23 in) long. Males weigh 3.7 to 6.2 kg (8.2 to 13.7 lb) and females 3 to 6.0 kg (6.6 to 13.2 lb).[9]
+The red panda is specialized as a bamboo feeder with strong, curved and sharp semi-retractile claws[9] standing inward for grasping narrow tree branches, leaves, and fruit. Like the giant panda, it has a "false thumb", which is an extension of the wrist bone. When descending a tree head-first, the red panda rotates its ankle to control its descent, one of the few climbing species to do so.[10]
+The red panda is endemic to the temperate forests of the Himalayas, and ranges from the foothills of western Nepal to China in the east.[11] Its easternmost limit is the Qinling Mountains of the Shaanxi Province in China. Its range includes southern Tibet, Sikkim and Assam in India, Bhutan, the northern mountains of Burma, and in south-western China, in the Hengduan Mountains of Sichuan and the Gongshan Mountains in Yunnan. It may also live in south-west Tibet and northern Arunachal Pradesh, but this has not been documented. Locations with the highest density of red pandas include an area in the Himalayas that has been proposed as having been a refuge for a variety of endemic species in the Pleistocene. The distribution range of the red panda should be considered disjunct, rather than continuous.[9] A disjunct population inhabits the Meghalaya Plateau of north-eastern India.[12]
+The red panda lives between 2,200 and 4,800 m (7,200 and 15,700 ft) altitude, inhabiting areas of moderate temperature between 10 and 25 °C (50 and 77 °F) with little annual change. It prefers mountainous mixed deciduous and conifer forests, especially with old trees and dense understories of bamboo.[9][11]
+During a survey in the 1970s, signs of red pandas were found in Nepal's Dhorpatan Hunting Reserve.[13] Their presence was confirmed in spring 2007 when four red pandas were sighted at elevations ranging from 3,220 to 3,610 m (10,560 to 11,840 ft).[14] Its westernmost distribution is in Rara National Park.[15][16] In 2018, red pandas were sighted at elevations of 3,150–3,650 m (10,330–11,980 ft) in Nepal's Lamjung District.[17]
+The red panda population in Sichuan Province is larger and more stable than the Yunnan population, suggesting a southward expansion from Sichuan into Yunnan in the Holocene.[18]
+The red panda has become extirpated from the Chinese provinces of Guizhou, Gansu, Shaanxi, and Qinghai.[19]
+The red panda is territorial; it is solitary except during mating season. It is generally quiet except for some twittering, tweeting, and whistling communication sounds. It has been reported to be both nocturnal and crepuscular, sleeping on tree branches or in tree hollows during the day and increasing its activity in the late afternoon and early evening hours. It sleeps stretched out on a branch with legs dangling when it is hot, and curled up with its tail over the face when it is cold.[9] It is very heat-sensitive, with an optimal "well-being" temperature between 17 and 25 °C (63 and 77 °F).[20]
+Shortly after waking, red pandas clean their fur somewhat like a cat would, licking their front paws and then rubbing their backs, torsos, and sides. They also rub their backs and bellies along the sides of trees or rocks. Then they patrol their territories, marking with urine and a weak musk-smelling secretion from their anal glands. They search for food running along the ground or through the trees. Red pandas may use their forepaws alternately to bring food to their mouths or place food directly into their mouths.[9]
+Predators of the red panda include the snow leopard (Panthera uncia), mustelids, and humans. If they feel threatened or sense danger, they may try to escape by climbing a rock column or tree. If they can no longer flee, they stand on their hind legs to make themselves appear larger and use the sharp claws on their front paws to defend themselves. A red panda became a visitor attraction in Japan for his ability to stand upright for ten seconds at a time.[21]
+Red pandas are excellent climbers, and forage largely in trees. They eat mostly bamboo, and may eat small mammals, birds, eggs, flowers, and berries. In captivity, they were observed to eat birds, flowers, maple and mulberry leaves, and bark and fruits of maple, beech, and mulberry.[9]
+Like the giant panda, they cannot digest cellulose, so they must consume a large volume of bamboo to survive. Their diets consist of about two-thirds bamboo, but they also eat mushrooms, roots, acorns, lichens, and grasses. Occasionally, they supplement their diets with fish and insects. They do little more than eat and sleep due to their low-calorie diets.[22][23]
+Bamboo shoots are more easily digested than leaves, exhibiting the highest digestibility in summer and autumn, intermediate digestibility in the spring, and lowest digestibility in the winter. These variations correlate with the nutrient contents in the bamboo. Red pandas process bamboo poorly, especially the cellulose and cell wall components. This implies microbial digestion plays only a minor role in their digestive strategy. To survive on this poor-quality diet, they have to eat the high-quality sections of the bamboo plant, such as the tender leaves and shoots, in large quantities, over 1.5 kg (3.3 lb) of fresh leaves and 4 kg (8.8 lb) of fresh shoots daily. This food passes through the digestive tract fairly rapidly (about 2–4 hr) so as to maximize daily nutrient intake.[24] Red pandas can taste artificial sweeteners, such as aspartame, and are the only nonprimates known to be able to do so.[25]
+Red pandas are able to reproduce at around 18 months of age, and are fully mature at two to three years. Adults rarely interact in the wild except to mate. Both sexes may mate with more than one partner during the mating season from mid-January to early March.[26] A few days before birth, females begin to collect material, such as brushwood, grass, and leaves; to build a nest, which is normally located in a hollow tree or a rock crevice. After a gestation period of 112 to 158 days, the female gives birth in mid-June to late July to one to four (usually 1–2) blind and deaf cubs weighing 110 to 130 g (3.9 to 4.6 oz) each.[9]
+After birth, the mother cleans the cubs, and can then recognize each by their smell. At first, she spends 60% to 90% of her time with the cubs. After the first week, the mother starts spending more time outside the nest, returning every few hours to nurse and groom the cubs. She moves the young frequently among several nests, all of which she keeps clean. The cubs start to open their eyes at about 18 days of age. By about 90 days, they achieve full adult fur and coloring, and begin to venture out of the nest. They also start eating solid foods at this point, weaning at around six to eight months of age. The cubs stay with their mother until the next litter is born in the following summer. Males rarely help raise the young, and only if they live in pairs or in small groups.[9]
+A red panda's lifespan ranges between eight and 10 years, but individuals have been known to reach 15 years.[27]
+The primary threats to red pandas are direct harvest from the wild, live or dead, competition with domestic livestock resulting in habitat degradation, and deforestation resulting in habitat loss or fragmentation. The relative importance of these factors is different in each region, and is not well understood.[11] For instance, in India, the biggest threat seems to be habitat loss followed by poaching, while in China, the biggest threat seems to be hunting and poaching.[1] A 40% decrease in red panda populations has been reported in China over the last 50 years, and populations in western Himalayan areas are considered to be lower.[19]
+Deforestation can inhibit the spread of red pandas and exacerbate the natural population subdivision by topography and ecology, leading to severe fragmentation of the remaining wild population. Fewer than 40 animals in four separate groups share resources with humans in Nepal's Langtang National Park, where only 6% of 1,710 km2 (660 sq mi) is preferred red panda habitat. Although direct competition for food with domestic livestock is not significant, livestock can depress bamboo growth by trampling.[28]
+Small groups of animals with little opportunity for exchange between them face the risk of inbreeding, decreased genetic diversity, and even extinction. In addition, clearcutting for firewood or agriculture, including hillside terracing, removes old trees that provide maternal dens and decreases the ability of some species of bamboo to regenerate.[11]
+In south-west China, red pandas are hunted for their fur, especially for the highly valued bushy tails, from which hats are produced. In these areas, the fur is often used for local cultural ceremonies. In weddings, the bridegroom traditionally carries the hide. The "good-luck charm" red panda-tail hats are also used by local newly-weds.[19] This practice may be quite old, as the red panda seems to be depicted in a 13th-century Chinese pen-and-ink scroll showing a hunting scene. Little or no mention of the red panda is made in the culture and folklore of Nepal.[29]
+In the past, red pandas were captured and sold to zoos. In an article appearing in the International Zoo News in 1969, one reported he personally had handled 350 red pandas in 17 years.[30]
+Due to CITES, this zoo harvest has decreased substantially in recent years, but poaching continues, and red pandas are often sold to private collectors at exorbitant prices. In some parts of Nepal and India, red pandas are kept as pets.[31]
+The red panda has a naturally low birth rate (usually one single or twin birth per year), and a high death rate in the wild[citation needed].
+The red panda is listed as endangered on the IUCN Red List since 2008 because the global population is estimated at about 10,000 individuals, with a decreasing population trend; only about half of the total area of potential habitat of 142,000 km2 (55,000 sq mi) is actually being used by the species. Due to its shy and secretive nature, and its largely nocturnal habits, observation of red pandas is difficult. Therefore, population figures in the wild are determined by population density estimates and not direct counts. It is protected in all range countries, and hunting is illegal. It is listed in CITES Appendix I.[1]
+Worldwide population estimates range from fewer than 2,500[26] to between 16,000 and 20,000 individuals.[12] In 1999, the total population in China was estimated at between 3,000 and 7,000 individuals.[19] In 2001, the wild population in India was estimated at between 5,000 and 6,000 individuals.[12] Estimates for Nepal indicate only a few hundred individuals.[32]
+Reliable population numbers are hard to find, partly because other animals have been mistaken for the red panda. For instance, one report from Myanmar stated that red pandas were still fairly common in some areas; however, the accompanying photographic proof of the "red panda" was in fact a viverrid species.[33]
+Conservation efforts are highly variable between countries:
+A community-managed forest in Ilam District of eastern Nepal is home to 15 red pandas which generate household income through tourism activities, including homestays. Villagers in the high-altitude areas of Arunachal Pradesh have formed the Pangchen Red Panda Conservation Alliance comprising five villages with a community-conserved forest area of 200 km2 (77 sq mi) at an altitude of 2,500 m (8,200 ft) to over 4,000 m (13,000 ft).[35]
+The international red panda studbook is currently managed at Rotterdam Zoo in the Netherlands. In cooperation with the International Red Panda Management Group, they coordinate the Species Survival Plan in North America, the European Endangered Species Programme in Europe, and other captive-breeding programs in Australia, India, Japan, and China. As of 2006, more than 800 individuals were kept in zoos and parks around the world. Of these, 511 individuals of the Himalayan red panda were kept in 173 institutions[36] and 306 individuals of Styan's red panda were kept in 81 institutions.[37]
+Since 2009, the North American Red Panda Species Survival Plan is coordinated at the Knoxville Zoo, which by 2011 had 101 red panda births. Only the Rotterdam Zoo has had more captive births worldwide.[36][37]
+The Padmaja Naidu Himalayan Zoological Park in Darjeeling successfully released four captive-bred red pandas to the wild in August and November 2003.[38]
+The most often cited example of keeping red pandas as pets is the case of former Indian prime minister Indira Gandhi. Pandas were presented to her family as a gift, and they were then housed in "a special tree house".[39]
+Ailurus fulgens was the scientific name proposed by Frédéric Cuvier in 1825, who described a zoological specimen sent by Alfred Duvaucel "from the mountains north of India". He was the first to also use the vernacular name panda.[40][41] In the 19th and 20th centuries, the following specimens were described:
+Pocock distinguished A. f. styani from A. f. fulgens by its longer winter coat and greater blackness of the pelage, bigger skull, more strongly curved forehead, and more robust teeth. His description is based on skulls and skins collected in Sichuan, Myitkyina District, close to the border of Yunnan, and Upper Burma.[8]
+Two subspecies are usually recognised, although results from a recent genomic study has suggested that these should be considered separate species:[1][7]
+The Brahmaputra River is often considered the natural barrier between the two subspecies, where it makes a curve around the eastern end of the Himalayas, although some authors suggest A. f. fulgens extends farther eastward into China.[43]
+The name Ailurus fulgens refulgens is sometimes incorrectly used for A. f. styani. This stems from a lapsus made by Henri Milne-Edwards in 1874.[44] making A. f. refulgens a nomen nudum.[43] This has been corrected in later publications.[45][46]
+At various times, the red panda was placed in the Procyonidae, Ursidae, with Ailuropoda (giant panda) in the Ailuropodinae (until this family was moved into the Ursidae), and into its own family, the Ailuridae. This uncertainty comes from difficulty in determining whether certain characteristics of Ailurus are phylogenetically conservative or are derived and convergent with species of similar ecological habits.[9]
+Evidence based on the fossil record, serology, karyology, behavior, anatomy, and reproduction reflect closer affinities with Procyonidae than Ursidae. However, ecological and foraging specializations and distinct geographical distribution in relation to modern procyonids support classification in the separate family Ailuridae.[6][9][47]
+Recent molecular systematic DNA research also places the red panda into its own family, Ailuridae, a part of the broad superfamily Musteloidea that also includes the mephitids (skunks), procyonids (raccoons), and mustelids (weasels).[5][47][48] According to the most recent phylogenetic studies, the red panda's closest relatives within the Musteloidea superfamily are the procyonids and mustelids.[49]
+It is not a bear, nor closely related to the giant panda, nor a raccoon, nor a lineage of uncertain affinities. Rather it is a basal lineage of musteloid, with a long history of independence from its closest relatives (skunks, raccoons, and otters/weasels/badgers).
+The red panda is considered a living fossil and only distantly related to the giant panda (Ailuropoda melanoleuca), as it is naturally more closely related to the other members of the superfamily Musteloidea to which it belongs. The common ancestor of both pandas (which also was an ancestor for all living bears; pinnipeds like seals and walruses; and members of the family Musteloidea like weasels and otters) can be traced back to the Paleogene period tens of millions of years ago, with a wide distribution across Eurasia.
+Fossils of the extinct red panda Parailurus anglicus were excavated in sites from China in the east to Britain in the west.[50] In 1977, a single tooth of Parailurus was discovered in the Pliocene Ringold Formation of Washington. This first North American record is almost identical to European specimens and indicates the immigration of this species from Asia.[51] In 2004, a tooth from a red panda species never before recorded in North America was discovered at the Gray Fossil Site in Tennessee. The tooth dates from 4.5–7 million years ago. This species, described as Pristinailurus bristoli, indicates that a second, more primitive ailurine lineage inhabited North America during the Miocene. Cladistic analysis suggests that Parailurus and Ailurus are sister taxa.[50][52] Additional fossils of Pristinailurus bristoli were discovered at the Gray Fossil Site in 2010 and in 2012.[53][54]
+The discovery in Spain of the postcranial remains of Simocyon batalleri, a Miocene relative to the red panda, supports a sister-group relationship between red pandas and bears. The discovery suggests the red panda's "false thumb" was an adaptation to arboreal locomotion — independent of the giant panda's adaptation to manipulate bamboo — one of the most dramatic cases of convergent evolution among vertebrates.[55]
+In 2020, results of a phylogenetic analysis of red panda samples showed that red pandas in China and the Himalayas were separated by a river about 250,000 years ago. Therefore, the two subspecies should be treated as distinct species. The analysed samples showed high levels of population structure across the red panda's range.[7] However, the results of this research should be treated with caution because of the sampling gap of >500 km between the two proposed species, and the lack of isolation-by-distance and morphometric analyses. Additionally, the use of the phylogenetic species concept for species delimitation in mammals has been associated with the unnecessary splitting of subspecies into species.[56][57]
+Ailurus is adopted from the ancient Greek word αἴλουρος (ailouros), meaning "cat".[58] The specific epithet fulgens is Latin for "shining, bright".[59][60]
+Panda is a Roman goddess of peace and travellers, who was called upon before starting a difficult journey.[61]
+The Lepcha call it sak nam. In Nepal, it is called bhalu biralo (bear-cat) and habre. The Sherpa people of Nepal and Sikkim call it ye niglva ponva and wah donka.[62] The word wậː is Sunuwari meaning bear; in Tamang language, a small, red bear is called tāwām.[63] In the Kanchenjunga region of eastern Nepal, the Limbu people know red pandas as kaala (literally "dark") because of their underside pelage; villagers of Tibetan origin call them hoptongar.[64]
+Additionally, Pocock lists the vernacular names ye and nigálya ponya (Nepal); thokya and thongwa (Limbu); oakdonga or wakdonka and woker (Bhotia); saknam sunam (Lepcha).[8] Nigálya may originate from the Nepali word निङालो niṅālo or nĩgālo, a small bamboo, Arundinaria intermedia, but also refers to a kind of small leopard, or cat-bear.[65] The word pónya may originate from the Nepali पञ्जा pajā ("claw") or पौँजा paũjā ("paw").[66]
+'Poonya' also means "eater of bamboo".[67] The name panda could originate from panjā.[68]
+In modern Chinese, the red panda is called xiăoxióngmāo (小熊猫 and 小熊貓, lesser or small panda, or literally "little bear cat"),[69] or 红熊猫/紅熊貓 (hóngxióngmāo, red panda or literally "red bear cat").[70]
+In English, the red panda is also called the "lesser panda", "true panda" and "common panda".[71][72][73]
+The first known written record of the red panda occurs in a 13th-century Chinese scroll depicting a hunting scene between hunters and the red panda.[29]
+The red panda was recognized as the state animal of Sikkim in the early 1990s,[74] and was the mascot of the Darjeeling Tea Festival.[29]
+In 2005, Babu, a male red panda at Birmingham Nature Centre in Birmingham, England, escaped[75] and briefly became a media celebrity,[75][76] before being recaptured. He was subsequently voted "Brummie of the Year", the first animal to receive this honor.[75][76]
+Rusty, a male red panda at the National Zoo in Washington, DC, similarly attracted media attention when he briefly escaped in 2013.[77][78]
+The name of the open-source Firefox web browser is said to have been derived from a nickname of the red panda: "fire fox".[79][80]
+An anthropomorphic red panda was featured as Master Shifu, the kung fu teacher, in the 2008 film Kung Fu Panda, and its sequels Kung Fu Panda 2 in 2011 and Kung Fu Panda 3 in 2016.[81] The red panda Futa inspired the character of Pabu, the so-called "fire ferret" animal companion (primarily of Bolin), in the U.S. animated TV series The Legend of Korra.[82]
+Jetstar Japan uses a red panda mascot character named "Jetta" (ジェッ太).[83]
+An anthropomorphic red panda, Retsuko, is the main character of the TV anime and Netflix original series Aggretsuko.[84]

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+Aglossata
+Glossata
+Heterobathmiina
+Zeugloptera
+Lepidoptera (/ˌlɛpɪˈdɒptərə/ LEP-i-DOP-tər-ə, from Ancient Greek lepís “scale” + pterón “wing”) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families[1] and 46 superfamilies,[2] 10 per cent of the total described species of living organisms.[2][3] It is one of the most widespread and widely recognizable insect orders in the world.[4] The Lepidoptera show many variations of the basic body structure that have evolved to gain advantages in lifestyle and distribution. Recent estimates suggest the order may have more species than earlier thought,[5] and is among the four most speciose orders, along with the Hymenoptera, Diptera, and Coleoptera.[4]
+Lepidopteran species are characterized by more than three derived features. The most apparent is the presence of scales that cover the bodies, wings, and a proboscis. The scales are modified, flattened "hairs", and give butterflies and moths their wide variety of colors and patterns. Almost all species have some form of membranous wings, except for a few that have reduced wings or are wingless. Mating and the laying of eggs are carried out by adults, normally near or on host plants for the larvae.  Like most other insects, butterflies and moths are holometabolous, meaning they undergo complete metamorphosis.  The larvae are commonly called caterpillars, and are completely different from their adult moth or butterfly forms, having a cylindrical body with a well-developed head, mandible mouth parts, three pairs of thoracic legs and from none up to five pairs of prolegs. As they grow, these larvae change in appearance, going through a series of stages called instars. Once fully matured, the larva develops into a pupa. A few butterflies and many moth species spin a silk case or cocoon prior to pupating, while others do not, instead going underground.[4] A butterfly pupa, called a chrysalis, has a hard skin, usually with no cocoon. Once the pupa has completed its metamorphosis, a sexually mature adult emerges.
+The Lepidoptera have, over millions of years, evolved a wide range of wing patterns and coloration ranging from drab moths akin to the related order Trichoptera, to the brightly colored and complex-patterned butterflies.[1] Accordingly, this is the most recognized and popular of insect orders with many people involved in the observation, study, collection, rearing of, and commerce in these insects. A person who collects or studies this order is referred to as a lepidopterist.
+Butterflies and moths play an important role in the natural ecosystem as pollinators and as food in the food chain; conversely, their larvae are considered very problematic to vegetation in agriculture, as their main source of food is often live plant matter. In many species, the female may produce from 200 to 600 eggs, while in others, the number may approach 30,000 eggs in one day. The caterpillars hatching from these eggs can cause damage to large quantities of crops. Many moth and butterfly species are of economic interest by virtue of their role as pollinators, the silk they produce, or as pest species.
+The term was coined by Linnaeus in 1735 and is derived from Greek λεπίς, gen. λεπίδος ("scale") and πτερόν ("wing").[6] Sometimes, the term Rhopalocera is used for the clade of all butterfly species, derived from the Ancient Greek ῥόπαλον (rhopalon)[7]:4150 and κέρας (keras)[7]:3993 meaning "club" and "horn", respectively, coming from the shape of the antennae of butterflies.
+The origins of the common names "butterfly" and "moth" are varied and often obscure. The English word butterfly is from Old English buttorfleoge, with many variations in spelling. Other than that, the origin is unknown, although it could be derived from the pale yellow color of many species' wings suggesting the color of butter.[8][9] The species of Heterocera are commonly called moths. The origins of the English word moth are more clear, deriving from the Old English moððe" (cf. Northumbrian dialect mohðe) from Common Germanic (compare Old Norse motti, Dutch mot and German Motte all meaning "moth"). Perhaps its origins are related to Old English maða meaning "maggot" or from the root of "midge", which until the 16th century was used mostly to indicate the larva, usually in reference to devouring clothes.[10]
+The etymological origins of the word "caterpillar", the larval form of butterflies and moths, are from the early 16th century, from Middle English catirpel, catirpeller, probably an alteration of Old North French catepelose: cate, cat (from Latin cattus) + pelose, hairy (from Latin pilōsus).[11]
+The Lepidoptera are among the most successful groups of insects. They are found on all continents, except Antarctica, and inhabit all terrestrial habitats ranging from desert to rainforest, from lowland grasslands to mountain plateaus, but almost always associated with higher plants, especially angiosperms (flowering plants).[12] Among the most northern dwelling species of butterflies and moths is the Arctic Apollo (Parnassius arcticus), which is found in the Arctic Circle in northeastern Yakutia, at an altitude of 1500 m above sea level.[13] In the Himalayas, various Apollo species such as Parnassius epaphus  have been recorded to occur up to an altitude of 6,000 m above sea level.[14]:221
+Some lepidopteran species exhibit symbiotic, phoretic, or parasitic lifestyles, inhabiting the bodies of organisms rather than the environment. Coprophagous pyralid moth species, called sloth moths, such as Bradipodicola hahneli and Cryptoses choloepi, are unusual in that they are exclusively found inhabiting the fur of sloths, mammals found in Central and South America.[15][16]
+Two species of Tinea moths have been recorded as feeding on horny tissue and have been bred from the horns of cattle. The larva of Zenodochium coccivorella is an internal parasite of the coccid Kermes species. Many species have been recorded as breeding in natural materials or refuse such as owl pellets, bat caves, honeycombs or diseased fruit.[16]
+As of 2007, there was roughly 174,250 lepidopteran species described, with butterflies and skippers estimated to comprise around 17,950, and moths making up the rest.[2][17] The vast majority of Lepidoptera are to be found in the tropics, but substantial diversity exists on most continents. North America has over 700 species of butterflies and over 11,000 species of moths,[18][19] while about 400 species of butterflies and 14,000 species of moths are reported from Australia.[20] The diversity of Lepidoptera in each faunal region has been estimated by John Heppner in 1991 based partly on actual counts from the literature, partly on the card indices in the Natural History Museum (London) and the National Museum of Natural History (Washington), and partly on estimates:[5]
+Lepidoptera are morphologically distinguished from other orders principally by the presence of scales on the external parts of the body and appendages, especially the wings. Butterflies and moths vary in size from microlepidoptera only a few millimeters long, to conspicuous animals with a wingspan greater than 25 centimetres, such as the Queen Alexandra's birdwing and Atlas moth.[21]:246
+Lepidopterans undergo a four-stage life cycle: egg; larva or caterpillar; pupa or chrysalis; and imago (plural: imagines) / adult and show many variations of the basic body structure, which give these animals advantages for diverse lifestyles and environments.
+The head is where many sensing organs and the mouth parts are found. Like the adult, the larva also has a toughened, or sclerotized head capsule.[22] Here, two compound eyes, and chaetosema, raised spots or clusters of sensory bristles unique to Lepidoptera, occur, though many taxa have lost one or both of these spots. The antennae have a wide variation in form among species and even between different sexes. The antennae of butterflies are usually filiform and shaped like clubs, those of the skippers are hooked, while those of moths have flagellar segments variously enlarged or branched. Some moths have enlarged antennae or ones that are tapered and hooked at the ends.[23]:559–560
+The maxillary galeae are modified and form an elongated proboscis. The proboscis consists of one to five segments, usually kept coiled up under the head by small muscles when it is not being used to suck up nectar from flowers or other liquids. Some basal moths still have mandibles, or separate moving jaws, like their ancestors, and these form the family Micropterigidae.[22][23]:560[24]
+The larvae, called caterpillars, have a toughened head capsule. Caterpillars lack the proboscis and have separate chewing mouthparts.[22] These mouthparts, called mandibles, are used to chew up the plant matter that the larvae eat. The lower jaw, or labium, is weak, but may carry a spinneret, an organ used to create silk. The head is made of large lateral lobes, each having an ellipse of up to six simple eyes.[23]:562–563
+The thorax is made of three fused segments, the prothorax, mesothorax, and metathorax, each with a pair of legs. The first segment contains the first pair of legs. In some males of the butterfly family Nymphalidae, the forelegs are greatly reduced and are not used for walking or perching.[23]:586 The three pairs of legs are covered with scales. Lepidoptera also have olfactory organs on their feet, which aid the butterfly in "tasting" or "smelling" out its food.[25] In the larval form there are 3 pairs of true legs, with up to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[12]
+The two pairs of wings are found on the middle and third segments, or mesothorax and metathorax, respectively. In the more recent genera, the wings of the second segment are much more pronounced, although some more primitive forms have similarly sized wings of both segments. The wings are covered in scales arranged like shingles, which form an extraordinary variety of colors and patterns. The mesothorax has more powerful muscles to propel the moth or butterfly through the air, with the wing of this segment (forewing) having a stronger vein structure.[23]:560 The largest superfamily, the Noctuidae, has their wings modified to act as tympanal or hearing organs.[26]
+The caterpillar has an elongated, soft body that may have hair-like or other projections, three pairs of true legs, with none to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[12] The thorax usually has a pair of legs on each segment. The thorax is also lined with many spiracles on both the mesothorax and metathorax, except for a few aquatic species, which instead have a form of gills.[23]:563
+The abdomen, which is less sclerotized than the thorax, consists of 10 segments with membranes in between, allowing for articulated movement. The sternum, on the first segment, is small in some families and is completely absent in others. The last two or three segments form the external parts of the species' sex organs. The genitalia of Lepidoptera are highly varied and are often the only means of differentiating between species. Male genitals include a valva, which is usually large, as it is used to grasp the female during mating. Female genitalia include three distinct sections.
+The females of basal moths have only one sex organ, which is used for copulation and as an ovipositor, or egg-laying organ. About 98% of moth species have a separate organ for mating, and an external duct that carries the sperm from the male.[23]:561
+The abdomen of the caterpillar has four pairs of prolegs, normally located on the third to sixth segments of the abdomen, and a separate pair of prolegs by the anus, which have a pair of tiny hooks called crotchets. These aid in gripping and walking, especially in species that lack many prolegs (e. g. larvae of Geometridae). In some basal moths, these prolegs may be on every segment of the body, while prolegs may be completely absent in other groups, which are more adapted to boring and living in sand (e. g., Prodoxidae and Nepticulidae, respectively).[23]:563
+The wings, head, and parts of the thorax and abdomen of Lepidoptera are covered with minute scales, a feature from which the order derives its name. Most scales are lamellar, or blade-like, and attached with a pedicel, while other forms may be hair-like or specialized as secondary sexual characteristics.[27]
+The lumen or surface of the lamella has a complex structure. It gives color either by colored pigments it contains, or through structural coloration with mechanisms that include photonic crystals and diffraction gratings.[28]
+Scales function in insulation, thermoregulation, producing pheromones (in males only),[29] and aiding gliding flight, but the most important is the large diversity of vivid or indistinct patterns they provide, which help the organism protect itself by camouflage or mimicry, and which act as signals to other animals including rivals and potential mates.[27]
+In the reproductive system of butterflies and moths, the male genitalia are complex and unclear. In females the three types of genitalia are based on the relating taxa: 'monotrysian', 'exoporian', and 'ditrysian'. In the monotrysian type is an opening on the fused segments of the sterna 9 and 10, which act as insemination and oviposition. In the exoporian type (in Hepaloidae and Mnesarchaeoidea) are two separate places for insemination and oviposition, both occurring on the same sterna as the monotrysian type, i.e. 9 and 10.[21] The ditrysian groups have an internal duct that carries sperm, with separate openings for copulation and egg-laying.[4] In most species, the genitalia are flanked by two soft lobes, although they may be specialized and sclerotized in some species for ovipositing in area such as crevices and inside plant tissue.[21] Hormones and the glands that produce them run the development of butterflies and moths as they go through their life cycles, called the endocrine system. The first insect hormone prothoracicotropic hormone (PTTH) operates the species life cycle and diapause.[30] This hormone is produced by corpora allata and corpora cardiaca, where it is also stored. Some glands are specialized to perform certain task such as producing silk or producing saliva in the palpi.[31]:65, 75 While the corpora cardiaca produce PTTH, the corpora allata also produces juvenile hormones, and the prothorocic glands produce moulting hormones.
+In the digestive system, the anterior region of the foregut has been modified to form a pharyngeal  sucking pump as they need it for the food they eat, which are for the most part liquids. An esophagus follows and leads to the posterior of the pharynx and in some species forms a form of crop. The midgut is short and straight, with the hindgut being longer and coiled.[21] Ancestors of lepidopteran species, stemming from Hymenoptera, had midgut ceca, although this is lost in current butterflies and moths. Instead, all the digestive enzymes, other than initial digestion, are immobilized at the surface of the midgut cells. In larvae, long-necked and stalked goblet cells are found in the anterior and posterior midgut regions, respectively. In insects, the goblet cells excrete positive potassium ions, which are absorbed from leaves ingested by the larvae. Most butterflies and moths display the usual digestive cycle, but species with different diets require adaptations to meet these new demands.[23]:279
+In the circulatory system, hemolymph, or insect blood, is used to circulate heat in a form of thermoregulation, where muscles contraction produces heat, which is transferred to the rest of the body when conditions are unfavorable.[32]
+In lepidopteran species, hemolymph is circulated through the veins in the wings by some form of pulsating organ, either by the heart or by the intake of air into the trachea.[31]:69
+Air is taken in through spiracles along the sides of the abdomen and thorax supplying the trachea with oxygen as it goes through the lepidopteran's respiratory system. Three different tracheaes supply and diffuse oxygen throughout the species' bodies. The dorsal tracheae supply oxygen to the dorsal musculature and vessels, while the ventral tracheae supply the ventral musculature and nerve cord, and the visceral tracheae supply the guts, fat bodies, and gonads.[31]:71, 72
+Polymorphism is the appearance of forms or "morphs", which differ in color and number of attributes within a single species.[12]:163[33] In Lepidoptera, polymorphism can be seen not only between individuals in a population, but also between the sexes as sexual dimorphism, between geographically separated populations in geographical polymorphism, and between generations flying at different seasons of the year (seasonal polymorphism or polyphenism). In some species, the polymorphism is limited to one sex, typically the female. This often includes the phenomenon of mimicry when mimetic morphs fly alongside nonmimetic morphs in a population of a particular species. Polymorphism occurs both at specific level with heritable variation in the overall morphological adaptations of individuals, as well as in certain specific morphological or physiological traits within a species.[12]
+Environmental polymorphism, in which traits are not inherited, is often termed as polyphenism, which in Lepidoptera is commonly seen in the form of seasonal morphs, especially in the butterfly families of Nymphalidae and Pieridae. An Old World pierid butterfly, the common grass yellow (Eurema hecabe) has a darker summer adult morph, triggered by a long day exceeding 13 hours in duration, while the shorter diurnal period of 12 hours or less induces a paler morph in the postmonsoon period.[34] Polyphenism also occurs in caterpillars, an example being the peppered moth, Biston betularia.[35]
+Geographical isolation causes a divergence of a species into different morphs. A good example is the Indian white admiral Limenitis procris, which has five forms, each geographically separated from the other by large mountain ranges.[36]:26 An even more dramatic showcase of geographical polymorphism is the Apollo butterfly (Parnassius apollo). Because the Apollos live in small local populations, thus having no contact with each other, coupled with their strong stenotopic nature and weak migration ability, interbreeding between populations of one species practically does not occur; by this, they form over 600 different morphs, with the size of spots on the wings of which varies greatly.[37]
+Sexual dimorphism is the occurrence of differences between males and females in a species. In Lepidoptera, it is widespread and almost completely set by genetic determination.[34] Sexual dimorphism is present in all families of the Papilionoidea and more prominent in the Lycaenidae, Pieridae, and certain taxa of the Nymphalidae. Apart from color variation, which may differ from slight to completely different color-pattern combinations, secondary sexual characteristics may also be present.[36]:25 Different genotypes maintained by natural selection may also be expressed at the same time.[34] Polymorphic and/or mimetic females occur in the case of some taxa in the Papilionidae primarily to obtain a level of protection not available to the male of their species. The most distinct case of sexual dimorphism is that of adult females of many Psychidae species which have only vestigial wings, legs, and mouthparts as compared to the adult males that are strong fliers with well-developed wings and feathery antennae.[38]
+Species of Lepidoptera undergo holometabolism or "complete metamorphosis". Their life cycle normally consists of an egg, a larva, a pupa, and an imago or adult.[12] The larvae are commonly called caterpillars, and the pupae of moths encapsulated in silk are called cocoons, while the uncovered pupae of butterflies are called chrysalides.
+Unless the species reproduces year-round, a butterfly or moth may enter diapause, a state of dormancy that allows the insect to survive unfavorable environmental conditions.
+Males usually start eclosion (emergence) earlier than females and peak in numbers before females. Both of the sexes are sexually mature by the time of eclosion.[23]:564 Butterflies and moths normally do not associate with each other, except for migrating species, staying relatively asocial. Mating begins with an adult (female or male) attracting a mate, normally using visual stimuli, especially in diurnal species like most butterflies. However, the females of most nocturnal species, including almost all moth species, use pheromones to attract males, sometimes from long distances.[12] Some species engage in a form of acoustic courtship, or attract mates using sound or vibration such as the polka-dot wasp moth, Syntomeida epilais.[39]
+Adaptations include undergoing one seasonal generation, two or even more, called voltinism (Univoltism, bivoltism, and multivism, respectively). Most lepidopterans in temperate climates are univoltine, while in tropical climates most have two seasonal broods. Some others may take advantage of any opportunity they can get, and mate continuously throughout the year. These seasonal adaptations are controlled by hormones, and these delays in reproduction are called diapause.[23]:567 Many lepidopteran species, after mating and laying their eggs, die shortly afterwards, having only lived for a few days after eclosion. Others may still be active for several weeks and then overwinter and become sexually active again when the weather becomes more favorable, or diapause. The sperm of the male that mated most recently with the female is most likely to have fertilized the eggs, but the sperm from a prior mating may still prevail.[23]:564
+Lepidoptera usually reproduce sexually and are oviparous (egg-laying), though some species exhibit live birth in a process called ovoviviparity. A variety of differences in egg-laying and the number of eggs laid occur. Some species simply drop their eggs in flight (these species normally have polyphagous larvae, meaning they eat a variety of plants e. g., hepialids and some nymphalids)[40] while most lay their eggs near or on the host plant on which the larvae feed. The number of eggs laid may vary from only a few to several thousand.[12] The females of both butterflies and moths select the host plant instinctively, and primarily, by chemical cues.[23]:564
+The eggs are derived from materials ingested as a larva and in some species, from the spermatophores received from males during mating.[41] An egg can only be 1/1000 the mass of the female, yet she may lay up to her own mass in eggs. Females lay smaller eggs as they age. Larger females lay larger eggs.[42] The egg is covered by a hard-ridged protective outer layer of shell, called the chorion. It is lined with a thin coating of wax, which prevents the egg from drying out. Each egg contains a number of micropyles, or tiny funnel-shaped openings at one end, the purpose of which is to allow sperm to enter and fertilize the egg. Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate.
+The egg stage lasts a few weeks in most butterflies, but eggs laid prior to winter, especially in temperate regions, go through diapause, and hatching may be delayed until spring. Other butterflies may lay their eggs in the spring and have them hatch in the summer. These butterflies are usually temperate species (e. g. Nymphalis antiopa).
+The larvae or caterpillars are the first stage in the life cycle after hatching. Caterpillars, are "characteristic polypod larvae with cylindrical bodies, short thoracic legs, and abdominal prolegs (pseudopods)".[43] They have a toughened (sclerotised) head capsule with an adfrontal suture formed by medial fusion of the sclerites, mandibles (mouthparts) for chewing, and a soft tubular, segmented body, that may have hair-like or other projections, three pairs of true legs, and additional prolegs (up to five pairs).[44] The body consists of thirteen segments, of which three are thoracic and ten are abdominal.[45] Most larvae are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores.[44]
+Different herbivorous species have adapted to feed on every part of the plant and are normally considered pests to their host plants; some species have been found to lay their eggs on the fruit and other species lay their eggs on clothing or fur (e. g., Tineola bisselliella, the common clothes moth). Some species are carnivorous and others are even parasitic. Some lycaenid species such as Phengaris rebeli are social parasites of Myrmica ants nests.[46] A species of Geometridae from Hawaii has carnivorous larvae that catch and eat flies.[47] Some pyralid caterpillars are aquatic.[48]
+The larvae develop rapidly with several generations in a year; however, some species may take up to 3 years to develop, and exceptional examples like Gynaephora groenlandica take as long as seven years.[12] The larval stage is where the feeding and growing stages occur, and the larvae periodically undergo hormone-induced ecdysis, developing further with each instar, until they undergo the final larval-pupal molt.
+The larvae of both butterflies and moths exhibit mimicry to deter potential predators. Some caterpillars have the ability to inflate parts of their heads to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called osmeteria (family Papilionidae), which are exposed to produce smelly chemicals used in defense. Host plants often have toxic substances in them and caterpillars are able to sequester these substances and retain them into the adult stage. This helps make them unpalatable to birds and other predators. Such unpalatability is advertised using bright red, orange, black, or white warning colors. The toxic chemicals in plants are often evolved specifically to prevent them from being eaten by insects. Insects, in turn, develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants.[49]
+No form of wing is externally visible on the larva, but when larvae are dissected, developing wings can be seen as disks, which can be found on the second and third thoracic segments, in place of the spiracles that are apparent on abdominal segments. Wing disks develop in association with a trachea that runs along the base of the wing, and are surrounded by a thin peripodial membrane, which is linked to the outer epidermis of the larva by a tiny duct. Wing disks are very small until the last larval instar, when they increase dramatically in size, are invaded by branching tracheae from the wing base that precede the formation of the wing veins, and begin to develop patterns associated with several landmarks of the wing.[50]
+Near pupation, the wings are forced outside the epidermis under pressure from the hemolymph, and although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval cuticle, they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled without damage to the wings.[50]
+After about five to seven instars,[51]:26–28 or molts, certain hormones, like PTTH, stimulate the production of ecdysone, which initiates insect molting. Then, the larva puparium, a sclerotized or hardened cuticle of the last larval instar, develops into the pupa. Depending on the species, the pupa may be covered in a silk cocoon, attached to different types of substrates, buried in the ground, or may not be covered at all. Features of the imago are externally recognizable in the pupa. All the appendages on the adult head and thorax are found cased inside the cuticle (antennae, mouthparts, etc.), with the wings wrapped around, adjacent to the antennae.[23]:564 The pupae of some species have functional mandibles, while the pupal mandibles are not functional in others.[22]
+While encased, some of the lower segments are not fused, and are able to move using small muscles found in between the membrane. Moving may help the pupa, for example, escape the sun, which would otherwise kill it. The pupa of the Mexican jumping bean moth (Cydia deshaisiana) does this. The larvae cut a trapdoor in the bean (species of Sebastiania) and use the bean as a shelter. With a sudden rise in temperature, the pupa inside twitches and jerks, pulling on the threads inside. Wiggling may also help to deter parasitoid wasps from laying eggs on the pupa. Other species of moths are able to make clicks to deter predators.[23]:564, 566
+The length of time before the pupa ecloses (emerges) varies greatly. The monarch butterfly may stay in its chrysalis for two weeks, while other species may need to stay for more than 10 months in diapause. The adult emerges from the pupa either by using abdominal hooks or from projections located on the head. The mandibles found in the most primitive moth families are used to escape from their cocoon (e. g., Micropterigoidea).[12][23]:564
+Most lepidopteran species do not live long after eclosion, only needing a few days to find a mate and then lay their eggs. Others may remain active for a longer period (from one to several weeks), or go through diapause and overwintering as monarch butterflies do, or waiting out environmental stress. Some adult species of microlepidoptera go through a stage where no reproductive-related activity occurs, lasting through summer and winter, followed by mating and oviposition in the early spring.[23]:564
+While most butterflies and moths are terrestrial, many species of Pyralidae are truly aquatic with all stages except the adult occurring in water. Many species from other families such as Erebidae, Nepticulidae, Cosmopterygidae, Tortricidae, Olethreutidae, Noctuidae, Cossidae, and Sphingidae are aquatic or semiaquatic.[52]:22
+Flight is an important aspect of the lives of butterflies and moths, and is used for evading predators, searching for food, and finding mates in a timely manner, as most lepidopteran species do not live long after eclosion. It is the main form of locomotion in most species. In Lepidoptera, the forewings and hindwings are mechanically coupled and flap in synchrony. Flight is anteromotoric, or being driven primarily by action of the forewings. Although lepidopteran species reportedly can still fly when their hindwings are cut off, it reduces their linear flight and turning capabilities.[53]
+Lepidopteran species have to be warm, about 77 to 79 °F (25 to 26 °C), to fly. They depend on their body temperature being sufficiently high and since they cannot regulate it themselves, this is dependent on their environment. Butterflies living in cooler climates may use their wings to warm their bodies. They will bask in the sun, spreading out their wings so that they get maximum exposure to the sunlight. In hotter climates butterflies can easily overheat, so they are usually active only during the cooler parts of the day, early morning, late afternoon or early evening. During the heat of the day, they rest in the shade. Some larger thick-bodied moths (e.g. Sphingidae) can generate their own heat to a limited degree by vibrating their wings. The heat generated by the flight muscles warms the thorax while the temperature of the abdomen is unimportant for flight. To avoid overheating, some moths rely on hairy scales, internal air sacs, and other structures to separate the thorax and abdomen and keep the abdomen cooler.[54]
+Some species of butterflies can reach fast speeds, such as the southern dart, which can go as fast as 48.4 km/h. Sphingids are some of the fastest flying insects, some are capable of flying at over 50 km/h (30 mi/h), having a wingspan of 35–150 mm.[1][55] In some species, sometimes a gliding component to their flight exists. Flight occurs either as hovering, or as forward or backward motion.[56] In butterfly and moth species, such as hawk moths, hovering is important as they need to maintain a certain stability over flowers when feeding on the nectar.[1]
+Navigation is important to Lepidoptera species, especially for those that migrate. Butterflies, which have more species that migrate, have been shown to navigate using time-compensated sun compasses. They can see polarized light, so can orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is suggested to be particularly important.[57] Most migratory butterflies are those that live in semiarid areas where breeding seasons are short.[58] The life histories of their host plants also influence the strategies of the butterflies.[59] Other theories include the use of landscapes. Lepidoptera may use coastal lines, mountains, and even roads to orient themselves. Above sea, the flight direction is much more accurate if the coast is still visible.[60]
+Many studies have also shown that moths navigate. One study showed that many moths may use the Earth's magnetic field to navigate, as a study of the moth heart and dart suggests.[61] Another study, of the migratory behavior of the silver Y, showed, even at high altitudes, the species can correct its course with changing winds, and prefers flying with favourable winds, suggesting a great sense of direction.[62][63] Aphrissa statira in Panama loses its navigational capacity when exposed to a magnetic field, suggesting it uses the Earth's magnetic field.[64]
+Moths exhibit a tendency to circle artificial lights repeatedly. This suggests they use a technique of celestial navigation called transverse orientation. By maintaining a constant angular relationship to a bright celestial light, such as the Moon, they can fly in a straight line. Celestial objects are so far away, even after traveling great distances, the change in angle between the moth and the light source is negligible; further, the moon will always be in the upper part of the visual field or on the horizon. When a moth encounters a much closer artificial light and uses it for navigation, the angle changes noticeably after only a short distance, in addition to being often below the horizon. The moth instinctively attempts to correct by turning toward the light, causing airborne moths to come plummeting downwards, and at close range, which results in a spiral flight path that gets closer and closer to the light source.[65] Other explanations have been suggested, such as the idea that moths may be impaired with a visual distortion called a Mach band by Henry Hsiao in 1972. He stated that they fly towards the darkest part of the sky in pursuit of safety, thus are inclined to circle ambient objects in the Mach band region.[66]
+Lepidopteran migration is typically seasonal, as the insects moving to escape dry seasons or other disadvantageous conditions. Most lepidopterans that migrate are butterflies, and the distance travelled varies. Some butterflies that migrate include the mourning cloak, painted lady, American lady, red admiral, and the common buckeye.[51]:29–30 A notable species of moth that migrates long distances is the bogong moth.[67] The most well-known migrations are those of the eastern population of the monarch butterfly from Mexico to northern United States and southern Canada, a distance of about 4,000–4,800 km (2,500–3,000 mi). Other well-known migratory species include the painted lady and several of the danaine butterflies. Spectacular and large-scale migrations associated with the monsoons are seen in peninsular India.[68] Migrations have been studied in more recent times using wing tags and stable hydrogen isotopes.[69][70]
+Moths also undertake migrations, an example being the uraniids. Urania fulgens undergoes population explosions and massive migrations that may be not surpassed by any other insect in the Neotropics. In Costa Rica and Panama, the first population movements may begin in July and early August and depending on the year, may be very massive, continuing unabated for as long as five months.[71]
+Pheromones are commonly involved in mating rituals among species, especially moths, but they are also an important aspect of other forms of communication. Usually, the pheromones are produced by either the male or the female and detected by members of the opposite sex with their antennae.[72] In many species, a gland between the eighth and ninth segments under the abdomen in the female produces the pheromones.[12] Communication can also occur through stridulation, or producing sounds by rubbing various parts of the body together.[63]
+Moths are known to engage in acoustic forms of communication, most often as courtship, attracting mates using sound or vibration. Like most other insects, moths pick up these sounds using tympanic membranes in their abdomens.[73] An example is that of the polka-dot wasp moth (Syntomeida epilais), which produces sounds with a frequency above that normally detectable by humans (about 20 kHz). These sounds also function as tactile communication, or communication through touch, as they stridulate, or vibrate a substrate like leaves and stems.[39]
+Most moths lack bright colors, as many species use coloration as camouflage, but butterflies engage in visual communication. Female cabbage butterflies, for example, use ultraviolet light to communicate, with scales colored in this range on the dorsal wing surface. When they fly, each down stroke of the wing creates a brief flash of ultraviolet light which the males apparently recognize as the flight signature of a potential mate. These flashes from the wings may attract several males that engage in aerial courtship displays.[73]
+Moths and butterflies are important in the natural ecosystem. They are integral participants in the food chain; having co-evolved with flowering plants and predators, lepidopteran species have formed a network of trophic relationships between autotrophs and heterotrophs, which are included in the stages of Lepidoptera larvae, pupae, and adults. Larvae and pupae are links in the diets of birds and parasitic entomophagous insects. The adults are included in food webs in a much broader range of consumers (including birds, small mammals, reptiles, etc.).[23]:567
+Lepidopteran species are soft bodied, fragile, and almost defenseless, while the immature stages move slowly or are immobile, hence all stages are exposed to predation. Adult butterflies and moths are preyed upon by birds, bats, lizards, amphibians, dragonflies, and spiders. Caterpillars and pupae fall prey not only to birds, but also to invertebrate predators and small mammals, as well as fungi and bacteria. Parasitoid and parasitic wasps and flies may lay eggs in the caterpillar, which eventually kill it as they hatch inside its body and eat its tissues. Insect-eating birds are probably the largest predators. Lepidoptera, especially the immature stages, are an ecologically important food to many insectivorous birds, such as the great tit in Europe.
+An "evolutionary arms race" can be seen between predator and prey species. The Lepidoptera have developed a number of strategies for defense and protection, including evolution of morphological characters and changes in ecological lifestyles and behaviors. These include aposematism, mimicry, camouflage, and development of threat patterns and displays.[74] Only a few birds, such as the nightjars, hunt nocturnal lepidopterans. Their main predators are bats. Again, an "evolutionary race" exists, which has led to numerous evolutionary adaptations of moths to escape from their main predators, such as the ability to hear ultrasonic sounds, or even to emit sounds in some cases. Lepidopteran eggs are also preyed upon. Some caterpillars, such as the zebra swallowtail butterfly larvae, are cannibalistic.
+Some species of Lepidoptera are poisonous to predators, such as the monarch butterfly in the Americas, Atrophaneura species (roses, windmills, etc.) in Asia, as well as Papilio antimachus, and the birdwings, the largest butterflies in Africa and Asia, respectively. They obtain their toxicity by sequestering the chemicals from the plants they eat into their own tissues. Some Lepidoptera manufacture their own toxins. Predators that eat poisonous butterflies and moths may become sick and vomit violently, learning not to eat those species. A predator which has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future, thus saving many other species, as well.[74][75] Toxic butterflies and larvae tend to develop bright colors and striking patterns as an indicator to predators about their toxicity. This phenomenon is known as aposematism.[76] Some caterpillars, especially members of Papilionidae, contain an osmeterium, a Y-shaped protrusible gland found in the prothoracic segment of the larvae.[74] When threatened, the caterpillar emits unpleasant smells from the organ to ward off the predators.[77][78]
+Camouflage is also an important defense strategy, which involves the use of coloration or shape to blend into the surrounding environment. Some lepidopteran species blend with their surroundings, making them difficult to spot by predators. Caterpillars can exhibit shades of green that match its host plant. Others look like inedible objects, such as twigs or leaves. For instance, the mourning cloak fades into the backdrop of trees when it folds its wings back. The larvae of some species, such as the common Mormon (Papilio polytes) and the western tiger swallowtail look like bird droppings.[74][79] For example, adult Sesiidae species (also known as clearwing moths) have a general appearance sufficiently similar to a wasp or hornet to make it likely the moths gain a reduction in predation by Batesian mimicry.[80] Eyespots are a type of automimicry used by some butterflies and moths. In butterflies, the spots are composed of concentric rings of scales in different colors. The proposed role of the eyespots is to deflect attention of predators. Their resemblance to eyes provokes the predator's instinct to attack these wing patterns.[81]
+Batesian and Müllerian mimicry complexes are commonly found in Lepidoptera. Genetic polymorphism and natural selection give rise to otherwise edible species (the mimic) gaining a survival advantage by resembling inedible species (the model). Such a mimicry complex is referred to as Batesian and is most commonly known in the example between the limenitidine viceroy butterfly in relation to the inedible danaine monarch. The viceroy is, in fact, more toxic than the monarch and this resemblance should be considered as a case of Müllerian mimicry.[82] In Müllerian mimicry, inedible species, usually within a taxonomic order, find it advantageous to resemble each other so as to reduce the sampling rate by predators that need to learn about the insects' inedibility. Taxa from the toxic genus Heliconius form one of the most well-known Müllerian complexes.[83] The adults of the various species now resemble each other so well, the species cannot be distinguished without close morphological observation and, in some cases, dissection or genetic analysis.
+Moths evidently are able to hear the range emitted by bats, which in effect causes flying moths to make evasive maneuvers because bats are a main predator of moths. Ultrasonic frequencies trigger a reflex action in the noctuid moth that cause it to drop a few inches in its flight to evade attack.[84] Tiger moths in a defense emit clicks within the same range of the bats, which interfere with the bats and foil their attempts to echolocate it.[85]
+Most species of Lepidoptera engage in some form of entomophily (more specifically psychophily and phalaenophily for butterflies and moths, respectively), or the pollination of flowers.[86] Most adult butterflies and moths feed on the nectar inside flowers, using their probosces to reach the nectar hidden at the base of the petals. In the process, the adults brush against the flowers' stamens, on which the reproductive pollen is made and stored. The pollen is transferred on appendages on the adults, which fly to the next flower to feed and unwittingly deposit the pollen on the stigma of the next flower, where the pollen germinates and fertilizes the seeds.[23]:813–814
+Flowers pollinated by butterflies tend to be large and flamboyant, pink or lavender in color, frequently having a landing area, and usually scented, as butterflies are typically day-flying. Since butterflies do not digest pollen (except for heliconid species,[86]) more nectar is offered than pollen. The flowers have simple nectar guides, with the nectaries usually hidden in narrow tubes or spurs, reached by the long "tongue" of the butterflies. Butterflies such as Thymelicus flavus have been observed to engage in flower constancy, which means they are more likely to transfer pollen to other conspecific plants. This can be beneficial for the plants being pollinated, as flower constancy prevents the loss of pollen during different flights and the pollinators from clogging stigmas with pollen of other flower species.[87]
+Among the more important moth pollinator groups are the hawk moths of the family Sphingidae. Their behavior is similar to hummingbirds, i.e., using rapid wing beats to hover in front of flowers. Most hawk moths are nocturnal or crepuscular, so moth-pollinated flowers (e.g., Silene latifolia ) tend to be white, night-opening, large, and showy with tubular corollae and a strong, sweet scent produced in the evening, night, or early morning. A lot of nectar is produced to fuel the high metabolic rates needed to power their flight.[88] Other moths (e.g., noctuids, geometrids, pyralids) fly slowly and settle on the flower. They do not require as much nectar as the fast-flying hawk moths, and the flowers tend to be small (though they may be aggregated in heads).[89]
+Mutualism is a form of biological interaction wherein each individual involved benefits in some way. An example of a mutualistic relationship would be that shared by yucca moths (Tegeculidae) and their host, yucca flowers (Liliaceae). Female yucca moths enter the host flowers, collect the pollen into a ball using specialized maxillary palps, then move to the apex of the pistil, where pollen is deposited on the stigma, and lay eggs into the base of the pistil where seeds will develop. The larvae develop in the fruit pod and feed on a portion of the seeds. Thus, both insect and plant benefit, forming a highly mutualistic relationship.[23]:814 Another form of mutualism occurs between some larvae of butterflies and certain species of ants (e.g. Lycaenidae). The larvae communicate with the ants using vibrations transmitted through a substrate, such as the wood of a tree or stems, as well as using chemical signals.[90] The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions.[91]
+Only 42 species of parasitoid lepidopterans are known (1 Pyralidae; 40 Epipyropidae).[23]:748 The larvae of the greater and lesser wax moths feed on the honeycomb inside bee nests and may become pests; they are also found in bumblebee and wasp nests, albeit to a lesser extent. In northern Europe, the wax moth is regarded as the most serious parasitoid of the bumblebee, and is found only in bumblebee nests. In some areas in southern England, as much as 80% of nests can be destroyed.[92] Other parasitic larvae are known to prey upon cicadas and leaf hoppers.[93]
+In reverse, moths and butterflies may be subject to parasitic wasps and flies, which may lay eggs on the caterpillars, which hatch and feed inside its body, resulting in death. Although, in a form of parasitism called idiobiont, the adult paralyzes the host, so as not to kill it but for it to live as long as possible, for the parasitic larvae to benefit the most. In another form of parasitism, koinobiont, the species live off their hosts while inside (endoparasitic). These parasites live inside the host caterpillar throughout its life cycle, or may affect it later on as an adult. In other orders, koinobionts include flies, a majority of coleopteran, and many hymenopteran parasitoids.[23]:748–749 Some species may be subject to a variety of parasites, such as the gypsy moth (Lymantaria dispar), which is attacked by a series of 13 species, in six different taxa throughout its life cycle.[23]:750
+In response to a parasitoid egg or larva in the caterpillar's body, the plasmatocytes, or simply the host's cells can form a multilayered capsule that eventually causes the endoparasite to asphyxiate. The process, called encapsulation, is one of the caterpillar's only means of defense against parasitoids.[23]:748
+A few species of Lepidoptera are secondary consumers, or predators. These species typically prey upon the eggs of other insects, aphids, scale insects, or ant larvae.[23]:567 Some caterpillars are cannibals, and others prey on caterpillars of other species (e.g. Hawaiian Eupithecia ). Those of the 15 species in Eupithecia that mirror inchworms, are the only known species of butterflies and moths that are ambush predators.[94] Four species are known to eat snails. For example, the Hawaiian caterpillar (Hyposmocoma molluscivora) uses silk traps, in a manner similar to that of spiders, to capture certain species of snails (typically Tornatellides).[93]
+Larvae of some species of moths in the Tineidae, Gelechioidae, and Noctuidae (family/superfamily/families, respectively), besides others, feed on detritus, or dead organic material, such as fallen leaves and fruit, fungi, and animal products, and turn it into humus.[23]:567 Well-known species include the cloth moths (Tineola bisselliella, Tinea pellionella, and Trichophaga tapetzella), which feed on detritus containing keratin, including hair, feathers, cobwebs, bird nests (particularly of domestic pigeons, Columba livia domestica) and fruits or vegetables. These species are important to ecosystems as they remove substances that would otherwise take a long time to decompose.[95]
+In 2015 it was reported that wasp bracovirus DNA was present in Lepidoptera such as monarch butterflies, silkworms and moths.[96] These were described in some newspaper articles as examples of a naturally occurring genetically engineered insects.[97]
+Linnaeus in Systema Naturae (1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx and Phalaena, with seven subgroups in Phalaena.[98] These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Michael Denis & Ignaz Schiffermüller (1775), Johan Christian Fabricius (1775) and Pierre André Latreille (1796). Jacob Hübner described many genera, and the lepidopteran genera were catalogued by Ferdinand Ochsenheimer and Georg Friedrich Treitschke in a series of volumes on the lepidopteran fauna of Europe published between 1807 and 1835.[98] Gottlieb August Wilhelm Herrich-Schäffer (several volumes, 1843–1856), and Edward Meyrick (1895) based their classifications primarily on wing venation. Sir George Francis Hampson worked on the microlepidoptera during this period and Philipp Christoph Zeller published The Natural History of the Tineinae also on microlepidoptera (1855).
+Among the first entomologists to study fossil insects and their evolution was Samuel Hubbard Scudder (1837–1911), who worked on butterflies.[99] He published a study of the Florissant deposits of Colorado, including the exceptionally preserved Prodryas persephone. Andreas V. Martynov (1879–1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.[99]
+Major contributions in the 20th century included the creation of the monotrysia and ditrysia (based on female genital structure) by Borner in 1925 and 1939.[98] Willi Hennig (1913–1976) developed the cladistic methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among monotrysian families and Kristensen worked more generally on insect phylogeny and higher Lepidoptera too.[98][99] While it is often found that DNA-based phylogenies differ from those based on morphology, this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.[99]
+Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not monophyletic: Microlepidoptera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.[98]
+The fossil record for Lepidoptera is lacking in comparison to other winged species, and tends not to be as common as some other insects in habitats that are most conducive to fossilization, such as lakes and ponds; their juvenile stage has only the head capsule as a hard part that might be preserved. The location and abundance of the most common moth species are indicative that mass migrations of moths occurred over the Palaeogene North Sea, which is why there is a serious lack of moth fossils.[100] Yet there are fossils, some preserved in amber and some in very fine sediments. Leaf mines are also seen in fossil leaves, although the interpretation of them is tricky.[99]
+Putative fossil stem group representatives of Amphiesmenoptera (the clade comprising Trichoptera and Lepidoptera) are known from the Triassic.[23]:567 The earliest known lepidopteran fossils are fossilized scales from the Triassic-Jurassic boundary. They were found as rare palynological elements in the sediments of the Triassic-Jurassic boundary from the cored Schandelah-1 well, drilled near Braunschweig in northern Germany. This pushes back the fossil record and origin of glossatan lepidopterans by about 70 million years, supporting molecular estimates of a Norian (ca 212 million years) divergence of glossatan and non-glossatan lepidopterans. The findings were reported in 2018 in the journal Science Advances. The authors of the study proposed that lepidopterans evolved a proboscis as an adaptation to drink from droplets and thin films of water for maintaining their fluid balance in the hot and arid climate of the Triassic.[101]
+The earliest described lepidopteran taxon is Archaeolepis mane, a primitive moth-like species from the Jurassic, dated back to around 190 million years ago, and known only from three wings found in Dorset, UK. The wings show scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared with Trichoptera (caddisflies).[102][103] Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from the Cretaceous, which all belong to primitive moth-like families.[99]
+Many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber. The oldest genuine butterflies of the superfamily Papilionoidea have been found in the Paleocene MoClay or Fur Formation of Denmark. The best preserved fossil lepidopteran is the Eocene Prodryas persephone from the Florissant Fossil Beds.
+Lepidoptera and Trichoptera (caddisflies) are sister groups, sharing many similarities that are lacking in others; for example the females of both orders are heterogametic, meaning they have two different sex chromosomes, whereas in most species the males are heterogametic and the females have two identical sex chromosomes. The adults in both orders display a particular wing venation pattern on their forewings. The larvae in the two orders have mouth structures and glands with which they make and manipulate silk. Willi Hennig grouped the two orders into the superorder Amphiesmenoptera; together they are sister to the extinct order Tarachoptera.[104] Lepidoptera descend from a diurnal moth-like common ancestor that either fed on dead or living plants.[105]
+The cladogram, based on a 2008 DNA and protein analysis, shows the order as a clade, sister to the Trichoptera, and more distantly related to the Diptera (true flies) and Mecoptera (scorpionflies).[106][107][108][109]
+Diptera (true flies)
+Mecoptera (scorpionflies)
+Boreidae (snow scorpionflies)
+Siphonaptera (fleas)
+Trichoptera (caddisflies)
+Lepidoptera (butterflies and moths)
+Hymenoptera (sawflies, wasps, ants, bees)
+Micropterigidae, Agathiphagidae and Heterobathmiidae are the oldest and most basal lineages of Lepidoptera. The adults of these families do not have the curled tongue or proboscis, that are found in most members of the order, but instead have chewing mandibles adapted for a special diet. Micropterigidae larvae feed on leaves, fungi, or liverworts (much like the Trichoptera).[98] Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live inside kauri pines and feed on seeds. In Heterobathmiidae the larvae feed on the leaves of Nothofagus, the southern beech tree. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon after metamorphosis.[98]
+The Eriocraniidae have a short coiled proboscis in the adult stage, and though they retain their pupal mandibles with which they escaped the cocoon, their mandibles are non-functional thereafter.[98] Most of these non-ditrysian families, are primarily leaf miners in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.[99]
+With the evolution of the Ditrysia in the mid-Cretaceous, there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is one cloaca, or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, Gracillarioidea and Gelechioidea are mostly leaf miners, but more recent lineages feed externally. In the Tineoidea, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.[99]
+The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.[99] They evolved about the time that flowering plants underwent an expansive adaptive radiation in the mid-Cretaceous, and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved coevolution or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.
+In the so-called "macrolepidoptera", which constitutes about 60% of lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant.[99] Many also have tympanal organs, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.[99]
+The main lineages in the macrolepidoptera are the Noctuoidea, Bombycoidea, Lasiocampidae, Mimallonoidea, Geometroidea and Rhopalocera. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a monophyletic group.[99] The Rhopalocera, comprising the Papilionoidea (butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved.[98] There is quite a good fossil record for this group, with the oldest skipper dating from 56 million years ago.[99]
+Taxonomy is the classification of species in selected taxa, the process of naming being called nomenclature. There are over 120 families in Lepidoptera, in 45 to 48 superfamilies. Lepidoptera have always been, historically, classified in five suborders, one of which is of primitive moths that never lost the morphological features of their ancestors. The rest of the moths and butterflies make up ninety-eight percent of the other taxa, making Ditrysia. More recently, findings of new taxa, larvae and pupa have aided in detailing the relationships of primitive taxa, phylogenetic analysis showing the primitive lineages to be paraphyletic compared to the rest of Lepidoptera lineages. Recently lepidopterists have abandoned clades like suborders, and those between orders and superfamilies.[23]:569
+Artistic depictions of butterflies have been used in many cultures including as early as 3500 years ago, in Egyptian hieroglyphs.[114] Today, butterflies are widely used in various objects of art and jewelry: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and in some mixed media artworks and furnishings.[115] Butterflies have also inspired the "butterfly fairy" as an art and fictional character.
+In many cultures the soul of a dead person is associated with the butterfly, for example in Ancient Greece, where the word for butterfly ψυχή (psyche) also means soul and breath. In Latin, as in Ancient Greece, the word for "butterfly" papilio was associated with the soul of the dead.[116] The skull-like marking on the thorax of the death's-head hawkmoth has helped these moths, particularly A. atropos, earn a negative reputation, such as associations with the supernatural and evil. The moth has been prominently featured in art and movies such as Un Chien Andalou (by Buñuel and Dalí) and The Silence of the Lambs, and in the artwork of the Japanese metal band Sigh's album Hail Horror Hail. According to Kwaidan: Stories and Studies of Strange Things, by Lafcadio Hearn, a butterfly was seen in Japan as the personification of a person's soul; whether they be living, dying, or already dead. One Japanese superstition says that if a butterfly enters your guestroom and perches behind the bamboo screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad omens. When Taira no Masakado was secretly preparing for his famous revolt, there appeared in Kyoto so vast a swarm of butterflies that the people were frightened—thinking the apparition to be a portent of coming evil.[117]
+In the ancient Mesoamerican city of Teotihuacan, the brilliantly colored image of the butterfly was carved into many temples, buildings, jewelry, and emblazoned on incense burners in particular. The butterfly was sometimes depicted with the maw of a jaguar and some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies to fire and warfare persisted through to the Aztec civilization and evidence of similar jaguar-butterfly images has been found among the Zapotec, and Maya civilizations.[118]
+The larvae of many lepidopteran species are major pests in agriculture. Some of the major pests include Tortricidae, Noctuidae, and Pyralidae. The larvae of the Noctuidae genus Spodoptera (armyworms), Helicoverpa (corn earworm), or Pieris brassicae can cause extensive damage to certain crops.[98] Helicoverpa zea larvae (cotton bollworms or tomato fruitworms) are polyphagous, meaning they eat a variety of crops, including tomatoes and cotton.[119] Peridroma saucia (variegated cutworms) are described as one of the most damaging pests to gardens, with the ability to destroy entire gardens and fields in a matter of days.[120]
+Butterflies and moths are one of the largest taxa to solely feed and be dependent on living plants, in terms of the number of species, and they are in many ecosystems, making up the largest biomass to do so. In many species, the female may produce anywhere from 200 to 600 eggs, while in some others it may go as high as 30,000 eggs in one day. This can create many problems for agriculture, where many caterpillars can affect acres of vegetation. Some reports estimate that there have been over 80,000 caterpillars of several different taxa feeding on a single oak tree. In some cases, phytophagous larvae can lead to the destruction of entire trees in relatively short periods of time.[23]:567
+Ecological ways of removing pest Lepidoptera species are becoming more economically viable, as research has shown ways like introducing parasitic wasps and flies. For example, Sarcophaga aldrichi, a fly which deposited larvae feed upon the pupae of the forest tent caterpillar moth. Pesticides can affect other species other than the species they are targeted to eliminate, damaging the natural ecosystem.[121] Another good biological pest control method is the use of pheromone traps. A pheromone trap is a type of insect trap that uses pheromones to lure insects. Sex pheromones and aggregating pheromones are the most common types used. A pheromone-impregnated lure is encased in a conventional trap such as a Delta trap, water-pan trap, or funnel trap.[122]
+Species of moths that are detritivores would naturally eat detritus containing keratin, such as hairs or feathers. Well known species are cloth moths (T. bisselliella, T. pellionella, and T. tapetzella), feeding on foodstuffs that people find economically important, such as cotton, linen, silk and wool fabrics as well as furs; furthermore they have been found on shed feathers and hair, bran, semolina and flour (possibly preferring wheat flour), biscuits, casein, and insect specimens in museums.[95]
+Even though most butterflies and moths affect the economy negatively, some species are a valuable economic resource. The most prominent example is that of the domesticated silkworm moth (Bombyx mori), the larvae of which make their cocoons out of silk, which can be spun into cloth. Silk is and has been an important economic resource throughout history. The species Bombyx mori has been domesticated to the point where it is completely dependent on mankind for survival.[123] A number of wild moths such as Bombyx mandarina, and Antheraea species, besides others, provide commercially important silks.[124]
+The preference of the larvae of most lepidopteran species to feed on a single species or limited range of plants is used as a mechanism for biological control of weeds in place of herbicides. The pyralid cactus moth was introduced from Argentina to Australia, where it successfully suppressed millions of acres of prickly pear cactus.[23]:567 Another species of the Pyralidae, called the alligator weed stem borer (Arcola malloi), was used to control the aquatic plant known as alligator weed (Alternanthera philoxeroides) in conjunction with the alligator weed flea beetle; in this case, the two insects work in synergy and the weed rarely recovers.[125]
+Breeding butterflies and moths, or butterfly gardening/rearing, has become an ecologically viable process of introducing species into the ecosystem to benefit it. Butterfly ranching in Papua New Guinea permits nationals of that country to "farm" economically valuable insect species for the collectors market in an ecologically sustainable manner.[126]
+Lepidoptera feature prominently in entomophagy as food items on almost every continent. While in most cases, adults, larvae or pupae are eaten as staples by indigenous people, beondegi or silkworm pupae are eaten as a snack in Korean cuisine[127] while Maguey worm is considered a delicacy in Mexico.[128] In some parts of Huasteca, the silk nests of the Madrone butterfly are maintained on the edge of roof tops of houses for consumption.[129] In the Carnia region of Italy, children catch and eat ingluvies of the toxic Zygaena moths in early summer. The ingluvies, despite having a very low cyanogenic content, serve as a convenient, supplementary source of sugar to the children who can include this resource as a seasonal delicacy at minimum risk.[130]
+Some larvae of both moths and butterflies have a form of hair that has been known to be a cause of human health problems. Caterpillar hairs sometimes have toxins in them and species from approximately 12 families of moths or butterflies worldwide can inflict serious human injuries (urticarial dermatitis and atopic asthma to osteochondritis, consumption coagulopathy, renal failure, and intracerebral hemorrhage).[131] Skin rashes are the most common, but there have been fatalities.[132] Lonomia is a frequent cause of envenomation in humans in Brazil, with 354 cases reported between 1989 and 2005. Lethality ranging up to 20% with death caused most often by intracranial hemorrhage.[133]
+These hairs have also been known to cause keratoconjunctivitis. The sharp barbs on the end of caterpillar hairs can get lodged in soft tissues and mucous membranes such as the eyes. Once they enter such tissues, they can be difficult to extract, often exacerbating the problem as they migrate across the membrane.[134] This becomes a particular problem in an indoor setting. The hairs easily enter buildings through ventilation systems and accumulate in indoor environments because of their small size, which makes it difficult for them to be vented out. This accumulation increases the risk of human contact in indoor environments.[135]

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+Coordinates: 37°58′17″N 23°43′35″E / 37.9714°N 23.7265°E / 37.9714; 23.7265
+The Parthenon (/ˈpɑːrθəˌnɒn, -nən/; Ancient Greek: Παρθενών; Greek: Παρθενώνας, Parthenónas, [parθeˈnonas]) is a former temple[4][5] on the Athenian Acropolis, Greece, dedicated to the goddess Athena, whom the people of Athens considered their patron. Construction began in 447 BC when the Athenian Empire was at the peak of its power. It was completed in 438 BC, although decoration of the building continued until 432 BC. It is the most important surviving building of Classical Greece, generally considered the zenith of the Doric order[by whom?]. Its decorative sculptures are considered some of the high points of Greek art. The Parthenon is regarded as an enduring symbol of Ancient Greece, Athenian democracy and Western civilization,[6] and one of the world's greatest cultural monuments. To the Athenians who built it, the Parthenon and other Periclean monuments of the Acropolis were seen fundamentally as a celebration of Hellenic victory over the Persian invaders and as a thanksgiving to the gods for that victory.[7]
+The Parthenon itself replaced an older temple of Athena, which historians call the Pre-Parthenon or Older Parthenon, that was destroyed in the Persian invasion of 480 BC. Like most Greek temples, the Parthenon served a practical purpose as the city treasury.[8][9] For a time, it served as the treasury of the Delian League, which later became the Athenian Empire. In the final decade of the 6th century AD, the Parthenon was converted into a Christian church dedicated to the Virgin Mary.
+After the Ottoman conquest, it was turned into a mosque in the early 1460s. On 26 September 1687, an Ottoman ammunition dump inside the building was ignited by Venetian bombardment during a siege of the Acropolis. The resulting explosion severely damaged the Parthenon and its sculptures. From 1800 to 1803,[10] Thomas Bruce, 7th Earl of Elgin removed some of the surviving sculptures, now known as the Elgin Marbles, with the alleged permission of the Turks of the Ottoman Empire.[11]
+Since 1975 numerous large-scale restoration projects have been undertaken; the latest is expected to finish in 2020.[12]
+The origin of the Parthenon's name is from the Greek word παρθενών (parthenon), which referred to the "unmarried women's apartments" in a house and in the Parthenon's case seems to have been used at first only for a particular room of the temple;[13] it is debated which room this is and how the room acquired its name. The Liddell–Scott–Jones Greek–English Lexicon states that this room was the western cella of the Parthenon, as does J.B. Bury.[7] Jamauri D. Green holds that the parthenon was the room in which the peplos presented to Athena at the Panathenaic Festival was woven by the arrephoroi, a group of four young girls chosen to serve Athena each year.[14] Christopher Pelling asserts that Athena Parthenos may have constituted a discrete cult of Athena, intimately connected with, but not identical to, that of Athena Polias.[15] According to this theory, the name of the Parthenon means the "temple of the virgin goddess" and refers to the cult of Athena Parthenos that was associated with the temple.[16] The epithet parthénos (παρθένος) meant "maiden, girl" as well as "virgin, unmarried woman."[17] The term was especially used for Artemis, the goddess of wild animals, vegetation, and the hunt and for Athena, the goddess of strategy, tactics, handicraft, and practical reason.[18] It has also been suggested that the name of the temple alludes to the maidens (parthénoi), whose supreme sacrifice guaranteed the safety of the city.[19] Parthénos has also been applied to the Virgin Mary (Parthénos Maria) and the Parthenon was converted to a Christian church dedicated to the Virgin Mary in the final decade of the 6th century.[20]
+The first instance in which Parthenon definitely refers to the entire building is found in the writings of the 4th century BC orator Demosthenes.  In 5th-century building accounts, the structure is simply called ὁ νᾱός (ho naos; lit. "the temple"). The architects Iktinos and Callicrates are said to have called the building Ἑκατόμπεδος (Hekatómpedos; lit. "the hundred footer") in their lost treatise on Athenian architecture.[21] Harpocration writes that the Parthenon used to be called Hekatompedos by some, not due to its size but because of its beauty and fine proportions[21] and, in the 4th century and later, the building was referred to as the Hekatompedos or the Hekatompedon as well as the Parthenon; the 1st-century-AD writer Plutarch referred to the building as the Hekatompedos Parthenon.[22]
+Because the Parthenon was dedicated to the Greek goddess Athena it has sometimes been referred to as the Temple of Minerva, the Roman name for Athena, particularly during the 19th century.[23]
+Although the Parthenon is architecturally a temple and is usually called so, it is not really one in the conventional sense of the word.[24] A small shrine has been excavated within the building, on the site of an older sanctuary probably dedicated to Athena as a way to get closer to the goddess,[24] but the Parthenon never hosted the cult of Athena Polias, patron of Athens: the cult image, which was bathed in the sea and to which was presented the peplos, was an olive-wood xoanon, located at an older altar on the northern side of the Acropolis.[25]
+The colossal statue of Athena by Phidias was not related to any cult[26] and is not known to have inspired any religious fervor.[25] It did not seem to have any priestess, altar or cult name.[27]
+According to Thucydides, Pericles once referred to the statue as a gold reserve, stressing that it, "contained forty talents of pure gold and it was all removable".[28] The Athenian statesman thus implies that the metal, obtained from contemporary coinage,[29] could be used again without any impiety.[27] The Parthenon should then be viewed as a grand setting for Phidias' votive statue rather than a cult site.[30] It is said[by whom?] in many writings of the Greeks that there were many treasures stored inside the temple, such as Persian swords and small statue figures made of precious metals.
+Archaeologist Joan Breton Connelly has recently argued for the coherency of the Parthenon's sculptural program in presenting a succession of genealogical narratives that track Athenian identity back through the ages: from the birth of Athena, through cosmic and epic battles, to the final great event of the Athenian Bronze Age, the war of Erechtheus and Eumolpos.[31][32] She argues a pedagogical function for the Parthenon's sculptured decoration, one that establishes and perpetuates Athenian foundation myth, memory, values and identity.[33][34] While some classicists, including Mary Beard, Peter Green, and Garry Wills[35][36] have doubted or rejected Connelly's thesis, an increasing number of historians, archaeologists, and classical scholars support her work. They include: J.J. Pollitt,[37] Brunilde Ridgway,[38] Nigel Spivey,[39] Caroline Alexander,[40] and A. E. Stallings.[41]
+The first endeavor to build a sanctuary for Athena Parthenos on the site of the present Parthenon was begun shortly after the Battle of Marathon (c. 490–488 BC) upon a solid limestone foundation that extended and leveled the southern part of the Acropolis summit. This building replaced a hekatompedon ("hundred-footer") and would have stood beside the archaic temple dedicated to Athena Polias ("of the city"). The Older or Pre-Parthenon, as it is frequently referred to, was still under construction when the Persians sacked the city in 480 BC and razed the Acropolis.[42][43]
+The existence of both the proto-Parthenon and its destruction were known from Herodotus,[44] and the drums of its columns were plainly visible built into the curtain wall north of the Erechtheion. Further physical evidence of this structure was revealed with the excavations of Panagiotis Kavvadias of 1885–90. The findings of this dig allowed Wilhelm Dörpfeld, then director of the German Archaeological Institute, to assert that there existed a distinct substructure to the original Parthenon, called Parthenon I by Dörpfeld, not immediately below the present edifice as had been previously assumed.[45] Dörpfeld's observation was that the three steps of the first Parthenon consisted of two steps of Poros limestone, the same as the foundations, and a top step of Karrha limestone that was covered by the lowest step of the Periclean Parthenon. This platform was smaller and slightly to the north of the final Parthenon, indicating that it was built for a wholly different building, now completely covered over. This picture was somewhat complicated by the publication of the final report on the 1885–90 excavations, indicating that the substructure was contemporary with the Kimonian walls, and implying a later date for the first temple.[46]
+If the original Parthenon was indeed destroyed in 480, it invites the question of why the site was left a ruin for thirty-three years. One argument involves the oath sworn by the Greek allies before the Battle of Plataea in 479 BC[47] declaring that the sanctuaries destroyed by the Persians would not be rebuilt, an oath from which the Athenians were only absolved with the Peace of Callias in 450.[48] The mundane fact of the cost of reconstructing Athens after the Persian sack is at least as likely a cause. However, the excavations of Bert Hodge Hill led him to propose the existence of a second Parthenon, begun in the period of Kimon after 468 BC.[49] Hill claimed that the Karrha limestone step Dörpfeld thought was the highest of Parthenon I was in fact the lowest of the three steps of Parthenon II, whose stylobate dimensions Hill calculated at 23.51 by 66.888 metres (77.13 ft × 219.45 ft).
+One difficulty in dating the proto-Parthenon is that at the time of the 1885 excavation the archaeological method of seriation was not fully developed; the careless digging and refilling of the site led to a loss of much valuable information. An attempt to discuss and make sense of the potsherds found on the Acropolis came with the two-volume study by Graef and Langlotz published in 1925–33.[50] This inspired American archaeologist William Bell Dinsmoor to attempt to supply limiting dates for the temple platform and the five walls hidden under the re-terracing of the Acropolis. Dinsmoor concluded that the latest possible date for Parthenon I was no earlier than 495 BC, contradicting the early date given by Dörpfeld.[51] Further, Dinsmoor denied that there were two proto-Parthenons, and held that the only pre-Periclean temple was what Dörpfeld referred to as Parthenon II. Dinsmoor and Dörpfeld exchanged views in the American Journal of Archaeology in 1935.[52]
+In the mid-5th century BC, when the Athenian Acropolis became the seat of the Delian League and Athens was the greatest cultural center of its time, Pericles initiated an ambitious building project that lasted the entire second half of the century. The most important buildings visible on the Acropolis today – the Parthenon, the Propylaia, the Erechtheion and the temple of Athena Nike – were erected during this period. The Parthenon was built under the general supervision of the artist Phidias, who also had charge of the sculptural decoration. The architects Ictinos and Callicrates began their work in 447 BC, and the building was substantially completed by 432. However, work on the decorations continued until at least 431.
+The Parthenon was built primarily by men who knew how to work marble. These quarrymen had exceptional skills and were able to cut the blocks of marble to very specific measurements. The quarrymen also knew how to avoid the faults, which were numerous in the Pentelic marble. If the marble blocks were not up to standard, the architects would reject them. The marble was worked with iron tools -- picks, points, punches, chisels and drills. The quarrymen would hold their tools against the marble block and firmly tap the surface of the rock.[53]
+A big project like the Parthenon attracted stonemasons from far and wide who traveled to Athens to assist in the project. Slaves and foreigners worked together with the Athenian citizens in the building of the Parthenon, doing the same jobs for the same pay. Temple building was a very specialized craft, and there were not many men in Greece qualified to build temples like the Parthenon, so these men would travel around and work where they were needed.[54]
+Other craftsmen also were necessary in the building of the Parthenon, specifically carpenters and metalworkers. Unskilled laborers also had key roles in the building of the Parthenon. These laborers loaded and unloaded the marble blocks and moved the blocks from place to place. In order to complete a project like the Parthenon, a number of different laborers were needed, and each played a critical role in constructing the final building.[55]
+The Parthenon is a peripteral octastyle Doric temple with Ionic architectural features. It stands on a platform or stylobate of three steps. In common with other Greek temples, it is of post and lintel construction and is surrounded by columns ('peripteral') carrying an entablature. There are eight columns at either end ('octastyle') and 17 on the sides. There is a double row of columns at either end. The colonnade surrounds an inner masonry structure, the cella, which is divided into two compartments. At either end of the building the gable is finished with a triangular pediment originally occupied by sculpted figures. The columns are of the Doric order, with simple capitals, fluted shafts, and no bases. Above the architrave of the entablature is a frieze of carved pictorial panels (metopes), separated by formal architectural triglyphs, typical of the Doric order. Around the cella and across the lintels of the inner columns runs a continuous sculptured frieze in low relief. This element of the architecture is Ionic in style rather than Doric.[56]
+Measured at the stylobate, the dimensions of the base of the Parthenon are 69.5 by 30.9 metres (228 by 101 ft). The cella was 29.8 meters long by 19.2 meters wide (97.8 × 63.0 ft). On the exterior, the Doric columns measure 1.9 metres (6.2 ft) in diameter and are 10.4 metres (34 ft) high. The corner columns are slightly larger in diameter. The Parthenon had 46 outer columns and 23 inner columns in total, each column having 20 flutes. (A flute is the concave shaft carved into the column form.) The roof was covered with large overlapping marble tiles known as imbrices and tegulae.[57][58]
+The Parthenon is regarded as the finest example of Greek architecture. The temple, wrote John Julius Cooper, "Enjoys the reputation of being the most perfect Doric temple ever built. Even in antiquity, its architectural refinements were legendary, especially the subtle correspondence between the curvature of the stylobate, the taper of the naos walls and the entasis of the columns."[59] Entasis refers to the slight swelling, of 4 centimetres (1.6 in), in the center of the columns to counteract the appearance of columns having a waist, as the swelling makes them look straight from a distance. The stylobate is the platform on which the columns stand. As in many other classical Greek temples,[60] it has a slight parabolic upward curvature intended to shed rainwater and reinforce the building against earthquakes. The columns might therefore be supposed to lean outward, but they actually lean slightly inward so that if they carried on, they would meet almost exactly 2,400 metres (1.5 mi) above the center of the Parthenon.[61] Since they are all the same height, the curvature of the outer stylobate edge is transmitted to the architrave and roof above: "All follow the rule of being built to delicate curves", Gorham Stevens observed when pointing out that, in addition, the west front was built at a slightly higher level than that of the east front.[62]
+It is not universally agreed what the intended effect of these "optical refinements" was. They may serve as a sort of "reverse optical illusion."[63] As the Greeks may have been aware, two parallel lines appear to bow, or curve outward, when intersected by converging lines. In this case, the ceiling and floor of the temple may seem to bow in the presence of the surrounding angles of the building. Striving for perfection, the designers may have added these curves, compensating for the illusion by creating their own curves, thus negating this effect and allowing the temple to be seen as they intended. It is also suggested that it was to enliven what might have appeared an inert mass in the case of a building without curves. But the comparison ought to be, according to Smithsonian historian Evan Hadingham, with the Parthenon's more obviously curved predecessors than with a notional rectilinear temple.[64]
+Some studies of the Acropolis, including of the Parthenon and its façade, have conjectured that many of its proportions approximate the golden ratio.[65] However, such theories have been discredited by more recent studies, which have shown that the proportions of the Parthenon do not match the golden proportion.[66][67]
+The cella of the Parthenon housed the chryselephantine statue of Athena Parthenos sculpted by Phidias and dedicated in 439 or 438 BC. The appearance of this is known from other images. The decorative stonework was originally highly colored.[68] The temple was dedicated to Athena at that time, though construction continued until almost the beginning of the Peloponnesian War in 432. By the year 438, the sculptural decoration of the Doric metopes on the frieze above the exterior colonnade, and of the Ionic frieze around the upper portion of the walls of the cella, had been completed. In the opisthodomos (the back room of the cella) were stored the monetary contributions of the Delian League, of which Athens was the leading member.
+Only a very small number of the sculptures remain in situ; most of the surviving sculptures are today (controversially) in the British Museum in London (as with the Parthenon Marbles) and the Acropolis Museum in Athens, with a few pieces in the Louvre, National Museum of Denmark, and museums in Rome, Vienna, and Palermo.[69]
+The frieze of the Parthenon's entablature contained 92 metopes, 14 each on the east and west sides, 32 each on the north and south sides. They were carved in high relief, a practice employed until then only in treasuries (buildings used to keep votive gifts to the gods).[70]  According to the building records, the metope sculptures date to the years 446–440 BC. The metopes of the east side of the Parthenon, above the main entrance, depict the Gigantomachy (the mythical battle between the Olympian gods and the Giants). The metopes of the west end show the Amazonomachy (the mythical battle of the Athenians against the Amazons). The metopes of the south side show the Thessalian Centauromachy (battle of the Lapiths aided by Theseus against the half-man, half-horse Centaurs). Metopes 13–21 are missing, but drawings from 1674 attributed to Jaques Carrey indicate a series of humans; these have been variously interpreted as scenes from the Lapith wedding, scenes from the early history of Athens and various myths.[71] On the north side of the Parthenon, the metopes are poorly preserved, but the subject seems to be the sack of Troy.
+The mythological figures of the metopes of the East, North, and West sides of the Parthenon were deliberately mutilated by Christian iconoclasts in late antiquity.[72]
+The metopes present examples of the Severe Style in the anatomy of the figures' heads, in the limitation of the corporal movements to the contours and not to the muscles, and in the presence of pronounced veins in the figures of the Centauromachy. Several of the metopes still remain on the building, but, with the exception of those on the northern side, they are severely damaged. Some of them are located at the Acropolis Museum, others are in the British Museum, and one is at the Louvre museum.[73]
+In March 2011, archaeologists announced that they had discovered five metopes of the Parthenon in the south wall of the Acropolis, which had been extended when the Acropolis was used as a fortress. According to Eleftherotypia daily, the archaeologists claimed the metopes had been placed there in the 18th century when the Acropolis wall was being repaired. The experts discovered the metopes while processing 2,250 photos with modern photographic methods, as the white Pentelic marble they are made of differed from the other stone of the wall. It was previously presumed that the missing metopes were destroyed during the Morosini explosion of the Parthenon in 1687.[74]
+The most characteristic feature in the architecture and decoration of the temple is the Ionic frieze running around the exterior of the cella walls. The bas-relief frieze was carved in situ and is dated to 442 BC-438 BC.
+One interpretation is that it depicts an idealized version of the Panathenaic procession from the Dipylon Gate in the Kerameikos to the Acropolis. In this procession held every year, with a special procession taking place every four years, Athenians and foreigners participated in honoring the goddess Athena by offering her sacrifices and a new peplos (a dress woven by selected noble Athenian girls called ergastines). The procession is more crowded (appearing to slow in pace) as it nears the gods on the eastern side of the temple.[75]
+Joan Breton Connelly offers a mythological interpretation for the frieze, one that is in harmony with the rest of the temple's sculptural program which shows Athenian genealogy through a series of succession myths set in the remote past. She identifies the central panel above the door of the Parthenon as the pre-battle sacrifice of the daughter of king Erechtheus, a sacrifice that ensured Athenian victory over Eumolpos and his Thracian army. The great procession marching toward the east end of the Parthenon shows the post-battle thanksgiving sacrifice of cattle and sheep, honey and water, followed by the triumphant army of Erechtheus returning from their victory. This represents the very first Panathenaia set in mythical times, the model on which historic Panathenaic processions was based.[76][77]
+The traveller Pausanias, when he visited the Acropolis at the end of the 2nd century AD, only mentioned briefly the sculptures of the pediments (gable ends) of the temple, reserving the majority of his description for the gold and ivory statue of the goddess inside.[78]
+The figures on the corners of the pediment depict the passage of time over the course of a full day.  Tethrippa of Helios and Selene are located on the left and right corners of the pediment respectively. The horses of Helios's chariot are shown with livid expressions as they ascend into the sky at the start of the day; whereas the Selene's horses struggle to stay on the pediment scene as the day comes to an end.[79][80]
+The supporters of Athena are extensively illustrated at the back of the left chariot, while the defenders of Poseidon are shown trailing behind the right chariot. It is believed that the corners of the pediment are filled by Athenian water deities, such as the Kephisos river, the Ilissos river, and nymph Kallirhoe. This belief merges from the fluid character of the sculptures' body position which represents the effort of the artist to give the impression of a flowing river.[81][82] Next to the left river god, there are the sculptures of the mythical king of Athens (Cecrops or Kekrops) with his daughters ( Aglaurus, Pandrosos,  Herse). The statue of Poseidon was the largest sculpture in the pediment until it broke into pieces during Francesco Morosini's effort to remove it in 1688. The posterior piece of the torso was found by Lusieri in the groundwork of a Turkish house in 1801 and is currently held in British Museum. The anterior portion was revealed by Ross in 1835 and is now held in the Acropolis Museum of Athens.[83]
+Every statue on the west pediment has a fully completed back, which would have been impossible to see when the sculpture was on the temple; this indicates that the sculptors put great effort into accurately portraying the human body.[82]
+The only piece of sculpture from the Parthenon known to be from the hand of Phidias[84] was the statue of Athena housed in the naos. This massive chryselephantine sculpture is now lost and known only from copies, vase painting, gems, literary descriptions and coins.[85]
+A major fire broke out in the Parthenon shortly after the middle of the third century AD[86][87] which destroyed the Parthenon's roof and much of the sanctuary's interior.[88] Heruli pirates are also credited with sacking Athens in 276, and destroying most of the public buildings there, including the Parthenon.[89] Repairs were made in the fourth century AD, possibly during the reign of Julian the Apostate.[90] A new wooden roof overlaid with clay tiles was installed to cover the sanctuary. It sloped at a greater incline than the original roof and left the building's wings exposed.[88]
+The Parthenon survived as a temple dedicated to Athena for nearly 1,000 years until Theodosius II, during the Persecution of pagans in the late Roman Empire, decreed in 435 AD that all pagan temples in the Eastern Roman Empire be closed.[91] However, it is debated exactly when during the 5th-century that the closure of the Parthenon as a temple was actually put in practice. It is suggested to have occurred in c. 481–484, in the instructions against the remaining temples by order of Emperor Zeno, because the temple had been the focus of Pagan Hellenic opposition against Zeno in Athens in support of Illus, who had promised to restore Hellenic rites to the temples that were still standing.[92]
+At some point in the Fifth Century, Athena's great cult image was looted by one of the emperors and taken to Constantinople, where it was later destroyed, possibly during the siege and sack of Constantinople during the Fourth Crusade in 1204 AD.[93]
+The Parthenon was converted into a Christian church in the final decade of the sixth century AD[20] to become the Church of the Parthenos Maria (Virgin Mary), or the Church of the Theotokos (Mother of God). The orientation of the building was changed to face towards the east; the main entrance was placed at the building's western end, and the Christian altar and iconostasis were situated towards the building's eastern side adjacent to an apse built where the temple's pronaos was formerly located.[94][95][96] A large central portal with surrounding side-doors was made in the wall dividing the cella, which became the church's nave, from the rear chamber, the church's narthex.[94] The spaces between the columns of the opisthodomos and the peristyle were walled up, though a number of doorways still permitted access.[94] Icons were painted on the walls and many Christian inscriptions were carved into the Parthenon's columns.[90] These renovations inevitably led to the removal and dispersal of some of the sculptures.
+The Parthenon became the fourth most important Christian pilgrimage destination in the Eastern Roman Empire after Constantinople, Ephesos, and Thessaloniki.[97] In 1018, the emperor Basil II went on a pilgrimage to Athens directly after his final victory over the Bulgarians for the sole purpose of worshipping at the Parthenon.[97] In medieval Greek accounts it is called the Temple of Theotokos Atheniotissa and often indirectly referred to as famous without explaining exactly which temple they were referring to, thus establishing that it was indeed well known.[97]
+At the time of the Latin occupation, it became for about 250 years a Roman Catholic church of Our Lady. During this period a tower, used either as a watchtower or bell tower and containing a spiral staircase, was constructed at the southwest corner of the cella, and vaulted tombs were built beneath the Parthenon's floor.[98]
+In 1456, Ottoman Turkish forces invaded Athens and laid siege to a Florentine army defending the Acropolis until June 1458, when it surrendered to the Turks.[99] The Turks may have briefly restored the Parthenon to the Greek Orthodox Christians for continued use as a church.[100] Some time before the close of the fifteenth century, the Parthenon became a mosque.[101][102]
+The precise circumstances under which the Turks appropriated it for use as a mosque are unclear; one account states that Mehmed II ordered its conversion as punishment for an Athenian plot against Ottoman rule.[103] The apse became a mihrab,[104] the tower previously constructed during the Roman Catholic occupation of the Parthenon was extended upwards to become a minaret,[105] a minbar was installed,[94] the Christian altar and iconostasis were removed, and the walls were whitewashed to cover icons of Christian saints and other Christian imagery.[106]
+Despite the alterations accompanying the Parthenon's conversion into a church and subsequently a mosque, its structure had remained basically intact.[107] In 1667 the Turkish traveller Evliya Çelebi expressed marvel at the Parthenon's sculptures and figuratively described the building as "like some impregnable fortress not made by human agency".[108] He composed a poetic supplication stating that, as "a work less of human hands than of Heaven itself, should remain standing for all time".[109] The French artist Jacques Carrey in 1674 visited the Acropolis and sketched the Parthenon's sculptural decorations.[110] Early in 1687, an engineer named Plantier sketched the Parthenon for the Frenchman Graviers d’Ortières.[88] These depictions, particularly those made by Carrey, provide important, and sometimes the only, evidence of the condition of the Parthenon and its various sculptures prior to the devastation it suffered in late 1687 and the subsequent looting of its art objects.[110]
+In 1687, the Parthenon was extensively damaged in the greatest catastrophe to befall it in its long history.[90] As part of the Morean War (1684–1699), the Venetians sent an expedition led by Francesco Morosini to attack Athens and capture the Acropolis. The Ottoman Turks fortified the Acropolis and used the Parthenon as a gunpowder magazine – despite having been forewarned of the dangers of this use by the 1656 explosion that severely damaged the Propylaea – and as a shelter for members of the local Turkish community.[111]
+On 26 September a Venetian mortar round, fired from the Hill of Philopappos, blew up the magazine, and the building was partly destroyed.[112] The explosion blew out the building's central portion and caused the cella's walls to crumble into rubble.[107] Greek architect and archaeologist Kornilia Chatziaslani writes that "...three of the sanctuary’s four walls nearly collapsed and three-fifths of the sculptures from the frieze fell. Nothing of the roof apparently remained in place. Six columns from the south side fell, eight from the north, as well as whatever remained from eastern porch, except for one column. The columns brought down with them the enormous marble architraves, triglyphs and metopes."[88] About three hundred people were killed in the explosion, which showered marble fragments over nearby Turkish defenders[111] and caused large fires that burned until the following day and consumed many homes.[88]
+Accounts written at the time conflict over whether this destruction was deliberate or accidental; one such account, written by the German officer Sobievolski, states that a Turkish deserter revealed to Morosini the use to which the Turks had put the Parthenon; expecting that the Venetians would not target a building of such historic importance. Morosini was said to have responded by directing his artillery to aim at the Parthenon.[88][111] Subsequently, Morosini sought to loot sculptures from the ruin and caused further damage in the process. Sculptures of Poseidon and Athena's horses fell to the ground and smashed as his soldiers tried to detach them from the building's west pediment.[95][113]
+The following year, the Venetians abandoned Athens to avoid a confrontation with a large force the Turks had assembled at Chalcis; at that time, the Venetians had considered blowing up what remained of the Parthenon along with the rest of the Acropolis to deny its further use as a fortification to the Turks, but that idea was not pursued.[111]
+Once the Turks had recaptured the Acropolis, they used some of the rubble produced by this explosion to erect a smaller mosque within the shell of the ruined Parthenon.[114] For the next century and a half, parts of the remaining structure were looted for building material and especially valuable objects.[115]
+The 18th century was a period of Ottoman stagnation—so that many more Europeans found access to Athens, and the picturesque ruins of the Parthenon were much drawn and painted, spurring a rise in philhellenism and helping to arouse sympathy in Britain and France for Greek independence. Amongst those early travellers and archaeologists were James Stuart and Nicholas Revett, who were commissioned by the Society of Dilettanti to survey the ruins of classical Athens. What they produced was the first measured drawings of the Parthenon, published in 1787 in the second volume of Antiquities of Athens Measured and Delineated. In 1801, the British Ambassador at Constantinople, the Earl of Elgin, obtained a questionable firman (edict) from the Sultan, whose existence or legitimacy has not been proved to this day, to make casts and drawings of the antiquities on the Acropolis, to demolish recent buildings if this was necessary to view the antiquities, and to remove sculptures from them.[citation needed]
+When independent Greece gained control of Athens in 1832, the visible section of the minaret was demolished; only its base and spiral staircase up to the level of the architrave remain intact.[116] Soon all the medieval and Ottoman buildings on the Acropolis were destroyed. However, the image of the small mosque within the Parthenon's cella has been preserved in Joly de Lotbinière's photograph, published in Lerebours's Excursions Daguerriennes in 1842: the first photograph of the Acropolis.[117] The area became a historical precinct controlled by the Greek government. In the later 19th century, the Parthenon was widely considered by Americans and Europeans to be the pinnacle of human architectural achievement, and became a popular destination and subject of artists, including Frederic Edwin Church and Sanford Robinson Gifford.[118][119] Today it attracts millions of tourists every year, who travel up the path at the western end of the Acropolis, through the restored Propylaea, and up the Panathenaic Way to the Parthenon, which is surrounded by a low fence to prevent damage.[citation needed]
+The dispute centers around the Parthenon Marbles removed by Thomas Bruce, 7th Earl of Elgin, from 1801 to 1803, which are in the British Museum. A few sculptures from the Parthenon are also in the Louvre in Paris, in Copenhagen, and elsewhere, but more than half are in the Acropolis Museum in Athens.[16][120] A few can still be seen on the building itself. The Greek government has campaigned since 1983 for the British Museum to return the sculptures to Greece.[120] The British Museum has steadfastly refused to return the sculptures,[121] and successive British governments have been unwilling to force the Museum to do so (which would require legislation). Nevertheless, talks between senior representatives from Greek and British cultural ministries and their legal advisors took place in London on 4 May 2007. These were the first serious negotiations for several years, and there were hopes that the two sides might move a step closer to a resolution.[122]
+In 1975, the Greek government began a concerted effort to restore the Parthenon and other Acropolis structures. After some delay, a Committee for the Conservation of the Acropolis Monuments was established in 1983.[123] The project later attracted funding and technical assistance from the European Union. An archaeological committee thoroughly documented every artifact remaining on the site, and architects assisted with computer models to determine their original locations. Particularly important and fragile sculptures were transferred to the Acropolis Museum. A crane was installed for moving marble blocks; the crane was designed to fold away beneath the roofline when not in use.[citation needed] In some cases, prior re-constructions were found to be incorrect. These were dismantled, and a careful process of restoration began.[124] Originally, various blocks were held together by elongated iron H pins that were completely coated in lead, which protected the iron from corrosion. Stabilizing pins added in the 19th century were not so coated, and corroded. Since the corrosion product (rust) is expansive, the expansion caused further damage by cracking the marble.[125]

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+"Little Red Riding Hood" is a European fairy tale about a young girl and a Big Bad Wolf.[1] Its origins can be traced back to the 10th century to several European folk tales, including one from Italy called The False Grandmother. The best known version was written by Charles Perrault.[2]
+The story has been changed considerably in various retellings and subjected to numerous modern adaptations and readings. Other names for the story are: "Little Red Ridinghood", "Little Red Cap" or simply "Red Riding Hood". It is number 333 in the Aarne–Thompson classification system for folktales.[3]
+The story revolves around a girl called Little Red Riding Hood. In Perrault's versions of the tale, she is named after her red hooded cape/cloak that she wears. The girl walks through the woods to deliver food to her sickly grandmother (wine and cake depending on the translation). In Grimms' version, her mother had ordered her to stay strictly on the path.
+A Big Bad Wolf wants to eat the girl and the food in the basket. He secretly stalks her behind trees, bushes, shrubs, and patches of little and tall grass. He approaches Little Red Riding Hood, who naively tells him where she is going. He suggests that the girl pick some flowers as a present for her grandmother, which she does. In the meantime, he goes to the grandmother's house and gains entry by pretending to be her. He swallows the grandmother whole (in some stories, he locks her in the closet) and waits for the girl, disguised as the grandma.
+When the girl arrives, she notices that her grandmother looks very strange. Little Red then says, "What a deep voice you have!" ("The better to greet you with", responds the wolf), "Goodness, what big eyes you have!" ("The better to see you with", responds the wolf), "And what big hands you have!" ("The better to embrace you with", responds the wolf), and lastly, "What a big mouth you have" ("The better to eat you with!", responds the wolf), at which point the wolf jumps out of the bed and eats her, too. Then he falls asleep. In Charles Perrault's version of the story (the first version to be published), the tale ends here. However, in later versions, the story continues generally as follows:
+A woodcutter in the French version, but a hunter in the Brothers Grimm and traditional German versions, comes to the rescue with an axe, and cuts open the sleeping wolf. Little Red Riding Hood and her grandmother emerge shaken, but unharmed. Then they fill the wolf's body with heavy stones. The wolf awakens and attempts to flee, but the stones cause him to collapse and die. In Grimm's version, the wolf leaves the house and tries to drink out of a well, but the stones in his stomach cause him to fall in and drown. Sanitized versions of the story have the grandmother locked in the closet instead of being eaten and some have Little Red Riding Hood saved by the lumberjack as the wolf advances on her rather than after she gets eaten, where the woodcutter kills the wolf with his axe.[4]
+The tale makes the clearest contrast between the safe world of the village and the dangers of the forest, conventional antitheses that are essentially medieval, though no written versions are as old as that.[citation needed][6] It also warns about the dangers of not obeying one's mother (at least in Grimms' version).[citation needed]
+The story displays many similarities to stories from classical Greece and Rome. Scholar Graham Anderson has compared the story to a local legend recounted by Pausanias in which, each year, a virgin girl was offered to a malevolent spirit dressed in the skin of a wolf, who raped the girl. Then, one year, the boxer Euthymos came along, slew the spirit, and married the girl who had been offered as a sacrifice.[7] There are also a number of different stories recounted by Greek authors involving a woman named Pyrrha (literally "fire") and a man with some name meaning "wolf".[8] The Roman poet Horace alludes to a tale in which a male child is rescued alive from the belly of Lamia, an ogress in classical mythology.[9]
+The dialogue between the Big Bad Wolf and Little Red Riding Hood has its analogies to the Norse Þrymskviða from the Elder Edda; the giant Þrymr had stolen Mjölnir, Thor's hammer, and demanded Freyja as his bride for its return. Instead, the gods dressed Thor as a bride and sent him. When the giants note Thor's unladylike eyes, eating, and drinking, Loki explains them as Freyja's not having slept, eaten, or drunk, out of longing for the wedding.[10] A parallel to another Norse myth, the chase and eventual murder of the sun goddess by the wolf Sköll, has also been drawn.[11]
+A very similar story also belongs to the North African tradition, namely in Kabylia, where a number of versions are attested.[12] The theme of the little girl who visits her (grand)dad in his cabin and is recognized by the sound of her bracelets constitutes the refrain of a well-known song by the modern singer Idir, A Vava Inouva:
+The theme of the ravening wolf and of the creature released unharmed from its belly is also reflected in the Russian tale Peter and the Wolf and another Grimm tale The Wolf and the Seven Young Kids, but its general theme of restoration is at least as old as the biblical story, Jonah and the Whale. The theme also appears in the story of the life of Saint Margaret, wherein the saint emerges unharmed from the belly of a dragon, and in the epic "The Red Path" by Jim C. Hines.
+A Taiwanese story from the 16th century, known as Grandaunt Tiger bears several striking similarities. When the girl's mother goes out, the tigress comes to the girl's house and pretends to be their aunt, asking to come in. The girl says that her voice does not sound right, so the tigress attempts to disguise her voice. Then, the girl says that her hands feel too coarse, so the tigress attempts to make them smoother. When finally, the tigress gains entry, she eats the girl's sister's hand. The girl comes up with a ruse to go outside and fetch some food for her aunt. Grandaunt Tiger, suspicious of the girl, ties a rope to her leg. The girl ties a bucket to the rope to fool her, but Grandaunt Tiger realises this and chases after her, whereupon she climbs into a tree. The girl tells the tigress that she will let her eat her, but first she would like to feed her some fruit from the tree. The tigress comes closer to eat the food, whereupon, the girl pours boiling hot oil down her throat, killing her.[14]
+The origins of the Little Red Riding Hood story can be traced to several likely pre-17th century versions from various European countries. Some of these are significantly different from the currently known, Grimms-inspired version. It was told by French peasants in the 10th century[1] and recorded by the cathedral schoolmaster Egbert of Liège.[15] In Italy, Little Red Riding Hood was told by peasants in the fourteenth century, where a number of versions exist, including La finta nonna (The False Grandmother), written among others by Italo Calvino in the Italian Folktales collection.[16] It has also been called "The Story of Grandmother". It is also possible that this early tale has roots in very similar East Asian tales (e.g. "Grandaunt Tiger").[17]
+These early variations of the tale, do differ from the currently known version in several ways. The antagonist is not always a wolf, but sometimes a 'bzou' (werewolf), making these tales relevant to the werewolf trials (similar to witch trials) of the time (e.g. the trial of Peter Stumpp).[18] The wolf usually leaves the grandmother's blood and flesh for the girl to eat, who then unwittingly cannibalizes her own grandmother. Furthermore, the wolf was also known to ask her to remove her clothing and toss it into the fire.[19] In some versions, the wolf eats the girl after she gets into bed with him, and the story ends there.[20] In others, she sees through his disguise and tries to escape, complaining to her "grandmother" that she needs to defecate and would not wish to do so in the bed. The wolf reluctantly lets her go, tied to a piece of string so she does not get away. However, the girl slips the string over something else and runs off. In these stories she escapes with no help from any male or older female figure, instead using her own cunning, or in some versions the help of a younger boy who she happens to run into.[21] Sometimes, though more rarely, the red hood is even non-existent.[20]
+In other tellings of the story, the wolf chases after Little Red Riding Hood. She escapes with the help of some laundresses, who spread a sheet taut over a river so she may escape. When the wolf follows Red over the bridge of cloth, the sheet is released and the wolf drowns in the river.[22] And in another version the wolf is pushed into the fire, while he is preparing the flesh of the grandmother to be eaten by the girl.[20]
+The earliest known printed version[23] was known as Le Petit Chaperon Rouge and may have had its origins in 17th-century French folklore. It was included in the collection Tales and Stories of the Past with Morals. Tales of Mother Goose (Histoires et contes du temps passé, avec des moralités. Contes de ma mère l'Oye), in 1697, by Charles Perrault. As the title implies, this version[24] is both more sinister and more overtly moralized than the later ones. The redness of the hood, which has been given symbolic significance in many interpretations of the tale, was a detail introduced by Perrault.[25]
+The story had as its subject an "attractive, well-bred young lady", a village girl of the country being deceived into giving a wolf she encountered the information he needed to find her grandmother's house successfully and eat the old woman while at the same time avoiding being noticed by woodcutters working in the nearby forest. Then he proceeded to lay a trap for Red Riding Hood. Little Red Riding Hood ends up being asked to climb into the bed before being eaten by the wolf, where the story ends. The wolf emerges the victor of the encounter and there is no happy ending.
+Charles Perrault explained the 'moral' at the end of the tale:[26] so that no doubt is left to his intended meaning:
+From this story one learns that children, especially young lasses, pretty, courteous and well-bred, do very wrong to listen to strangers, And it is not an unheard thing if the Wolf is thereby provided with his dinner. I say Wolf, for all wolves are not of the same sort; there is one kind with an amenable disposition – neither noisy, nor hateful, nor angry, but tame, obliging and gentle, following the young maids in the streets, even into their homes. Alas! Who does not know that these gentle wolves are of all such creatures the most dangerous!
+This, the presumed original version of the tale was written for the late seventeenth-century French court of King Louis XIV. This audience, whom the King entertained with extravagant parties, presumably would take from the story the intended meaning.
+In the 19th century two separate German versions were retold to Jacob Grimm and his younger brother Wilhelm Grimm, known as the Brothers Grimm, the first by Jeanette Hassenpflug (1791–1860) and the second by Marie Hassenpflug (1788–1856). The brothers turned the first version to the main body of the story and the second into a sequel of it. The story as Rotkäppchen was included in the first edition of their collection Kinder- und Hausmärchen (Children's and Household Tales (1812)).[27]
+The earlier parts of the tale agree so closely with Perrault's variant that it is almost certainly the source of the tale.[28] However, they modified the ending; this version had the little girl and her grandmother saved by a huntsman who was after the wolf's skin; this ending is identical to that in the tale "The Wolf and the Seven Young Kids", which appears to be the source.[29] The second part featured the girl and her grandmother trapping and killing another wolf, this time anticipating his moves based on their experience with the previous one. The girl did not leave the path when the wolf spoke to her, her grandmother locked the door to keep it out, and when the wolf lurked, the grandmother had Little Red Riding Hood put a trough under the chimney and fill it with water that sausages had been cooked in; the smell lured the wolf down, and it drowned.[30]
+The Brothers further revised the story in later editions and it reached the above-mentioned final and better-known version in the 1857 edition of their work.[31] It is notably tamer than the older stories which contained darker themes.
+Numerous authors have rewritten or adapted this tale.
+Andrew Lang included a variant called "The True History of Little Goldenhood"[32] in The Red Fairy Book (1890). He derived it from the works of Charles Marelles,[33] in Contes of Charles Marelles. This version explicitly states that the story had been mistold earlier. The girl is saved, but not by the huntsman; when the wolf tries to eat her, its mouth is burned by the golden hood she wears, which is enchanted.
+James N. Barker wrote a variation of Little Red Riding Hood in 1827 as an approximately 1000-word story. It was later reprinted in 1858 in a book of collected stories edited by William E Burton, called the Cyclopedia of Wit and Humor. The reprint also features a wood engraving of a clothed wolf on a bended knee holding Little Red Riding Hood's hand.
+In the 20th century, the popularity of the tale appeared to snowball, with many new versions being written and produced, especially in the wake of Freudian analysis, deconstruction and feminist critical theory. (See "Modern uses and adaptations" below.) This trend has also led to a number of academic texts being written that focus on Little Red Riding Hood, including works by Alan Dundes and Jack Zipes.
+Apart from the overt warning about talking to strangers, there are many interpretations of the classic fairy tale, many of them sexual.[34] Some are listed below.
+Folklorists and cultural anthropologists, such as P. Saintyves and Edward Burnett Tylor, saw "Little Red Riding Hood" in terms of solar myths and other naturally occurring cycles. Her red hood could represent the bright sun which is ultimately swallowed by the terrible night (the wolf), and the variations in which she is cut out of the wolf's belly represent the dawn.[35] In this interpretation, there is a connection between the wolf of this tale and Sköll, the wolf in Norse mythology that will swallow the personified Sun at Ragnarök, or Fenrir.[36] Alternatively, the tale could be about the season of spring or the month of May, escaping the winter.[37]
+The tale has been interpreted as a puberty rite, stemming from a prehistoric origin (sometimes an origin stemming from a previous matriarchal era).[38] The girl, leaving home, enters a liminal state and by going through the acts of the tale, is transformed into an adult woman by the act of coming out of the wolf's stomach.[39]
+Bruno Bettelheim, in The Uses of Enchantment: The Meaning and Importance of Fairy Tales (1976), recast the Little Red Riding Hood motif in terms of classic Freudian analysis, that shows how fairy tales educate, support, and liberate children's emotions. The motif of the huntsman cutting open the wolf he interpreted as a "rebirth"; the girl who foolishly listened to the wolf has been reborn as a new person.[40]
+The poem "Þrymskviða" from the Poetic Edda mirrors some elements of Red Riding Hood. Loki's explanations for the strange behavior of "Freyja" (actually Thor disguised as Freyja) mirror the wolf's explanations for his strange appearance. The red hood has often been given great importance in many interpretations, with a significance from the dawn to blood.[41]
+A sexual analysis of the tale may also include negative connotations in terms of rape or abduction. In Against Our Will, Susan Brownmiller describes the fairy tale as a description of rape.[42] However, many revisionist retellings choose to focus on empowerment, and depict Little Red Riding Hood or the grandmother successfully defending herself against the wolf.[43]
+Such tellings bear some similarity to the "animal bridegroom" tales, such as Beauty and the Beast or The Frog Prince, but where the heroines of those tales revert the hero to a prince, these tellings of Little Red Riding Hood reveal to the heroine that she has a wild nature like the hero's.[44] These interpretations refuse to characterize Little Red Riding Hood as a victim; these are tales of female empowerment.

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+"Little Red Riding Hood" is a European fairy tale about a young girl and a Big Bad Wolf.[1] Its origins can be traced back to the 10th century to several European folk tales, including one from Italy called The False Grandmother. The best known version was written by Charles Perrault.[2]
+The story has been changed considerably in various retellings and subjected to numerous modern adaptations and readings. Other names for the story are: "Little Red Ridinghood", "Little Red Cap" or simply "Red Riding Hood". It is number 333 in the Aarne–Thompson classification system for folktales.[3]
+The story revolves around a girl called Little Red Riding Hood. In Perrault's versions of the tale, she is named after her red hooded cape/cloak that she wears. The girl walks through the woods to deliver food to her sickly grandmother (wine and cake depending on the translation). In Grimms' version, her mother had ordered her to stay strictly on the path.
+A Big Bad Wolf wants to eat the girl and the food in the basket. He secretly stalks her behind trees, bushes, shrubs, and patches of little and tall grass. He approaches Little Red Riding Hood, who naively tells him where she is going. He suggests that the girl pick some flowers as a present for her grandmother, which she does. In the meantime, he goes to the grandmother's house and gains entry by pretending to be her. He swallows the grandmother whole (in some stories, he locks her in the closet) and waits for the girl, disguised as the grandma.
+When the girl arrives, she notices that her grandmother looks very strange. Little Red then says, "What a deep voice you have!" ("The better to greet you with", responds the wolf), "Goodness, what big eyes you have!" ("The better to see you with", responds the wolf), "And what big hands you have!" ("The better to embrace you with", responds the wolf), and lastly, "What a big mouth you have" ("The better to eat you with!", responds the wolf), at which point the wolf jumps out of the bed and eats her, too. Then he falls asleep. In Charles Perrault's version of the story (the first version to be published), the tale ends here. However, in later versions, the story continues generally as follows:
+A woodcutter in the French version, but a hunter in the Brothers Grimm and traditional German versions, comes to the rescue with an axe, and cuts open the sleeping wolf. Little Red Riding Hood and her grandmother emerge shaken, but unharmed. Then they fill the wolf's body with heavy stones. The wolf awakens and attempts to flee, but the stones cause him to collapse and die. In Grimm's version, the wolf leaves the house and tries to drink out of a well, but the stones in his stomach cause him to fall in and drown. Sanitized versions of the story have the grandmother locked in the closet instead of being eaten and some have Little Red Riding Hood saved by the lumberjack as the wolf advances on her rather than after she gets eaten, where the woodcutter kills the wolf with his axe.[4]
+The tale makes the clearest contrast between the safe world of the village and the dangers of the forest, conventional antitheses that are essentially medieval, though no written versions are as old as that.[citation needed][6] It also warns about the dangers of not obeying one's mother (at least in Grimms' version).[citation needed]
+The story displays many similarities to stories from classical Greece and Rome. Scholar Graham Anderson has compared the story to a local legend recounted by Pausanias in which, each year, a virgin girl was offered to a malevolent spirit dressed in the skin of a wolf, who raped the girl. Then, one year, the boxer Euthymos came along, slew the spirit, and married the girl who had been offered as a sacrifice.[7] There are also a number of different stories recounted by Greek authors involving a woman named Pyrrha (literally "fire") and a man with some name meaning "wolf".[8] The Roman poet Horace alludes to a tale in which a male child is rescued alive from the belly of Lamia, an ogress in classical mythology.[9]
+The dialogue between the Big Bad Wolf and Little Red Riding Hood has its analogies to the Norse Þrymskviða from the Elder Edda; the giant Þrymr had stolen Mjölnir, Thor's hammer, and demanded Freyja as his bride for its return. Instead, the gods dressed Thor as a bride and sent him. When the giants note Thor's unladylike eyes, eating, and drinking, Loki explains them as Freyja's not having slept, eaten, or drunk, out of longing for the wedding.[10] A parallel to another Norse myth, the chase and eventual murder of the sun goddess by the wolf Sköll, has also been drawn.[11]
+A very similar story also belongs to the North African tradition, namely in Kabylia, where a number of versions are attested.[12] The theme of the little girl who visits her (grand)dad in his cabin and is recognized by the sound of her bracelets constitutes the refrain of a well-known song by the modern singer Idir, A Vava Inouva:
+The theme of the ravening wolf and of the creature released unharmed from its belly is also reflected in the Russian tale Peter and the Wolf and another Grimm tale The Wolf and the Seven Young Kids, but its general theme of restoration is at least as old as the biblical story, Jonah and the Whale. The theme also appears in the story of the life of Saint Margaret, wherein the saint emerges unharmed from the belly of a dragon, and in the epic "The Red Path" by Jim C. Hines.
+A Taiwanese story from the 16th century, known as Grandaunt Tiger bears several striking similarities. When the girl's mother goes out, the tigress comes to the girl's house and pretends to be their aunt, asking to come in. The girl says that her voice does not sound right, so the tigress attempts to disguise her voice. Then, the girl says that her hands feel too coarse, so the tigress attempts to make them smoother. When finally, the tigress gains entry, she eats the girl's sister's hand. The girl comes up with a ruse to go outside and fetch some food for her aunt. Grandaunt Tiger, suspicious of the girl, ties a rope to her leg. The girl ties a bucket to the rope to fool her, but Grandaunt Tiger realises this and chases after her, whereupon she climbs into a tree. The girl tells the tigress that she will let her eat her, but first she would like to feed her some fruit from the tree. The tigress comes closer to eat the food, whereupon, the girl pours boiling hot oil down her throat, killing her.[14]
+The origins of the Little Red Riding Hood story can be traced to several likely pre-17th century versions from various European countries. Some of these are significantly different from the currently known, Grimms-inspired version. It was told by French peasants in the 10th century[1] and recorded by the cathedral schoolmaster Egbert of Liège.[15] In Italy, Little Red Riding Hood was told by peasants in the fourteenth century, where a number of versions exist, including La finta nonna (The False Grandmother), written among others by Italo Calvino in the Italian Folktales collection.[16] It has also been called "The Story of Grandmother". It is also possible that this early tale has roots in very similar East Asian tales (e.g. "Grandaunt Tiger").[17]
+These early variations of the tale, do differ from the currently known version in several ways. The antagonist is not always a wolf, but sometimes a 'bzou' (werewolf), making these tales relevant to the werewolf trials (similar to witch trials) of the time (e.g. the trial of Peter Stumpp).[18] The wolf usually leaves the grandmother's blood and flesh for the girl to eat, who then unwittingly cannibalizes her own grandmother. Furthermore, the wolf was also known to ask her to remove her clothing and toss it into the fire.[19] In some versions, the wolf eats the girl after she gets into bed with him, and the story ends there.[20] In others, she sees through his disguise and tries to escape, complaining to her "grandmother" that she needs to defecate and would not wish to do so in the bed. The wolf reluctantly lets her go, tied to a piece of string so she does not get away. However, the girl slips the string over something else and runs off. In these stories she escapes with no help from any male or older female figure, instead using her own cunning, or in some versions the help of a younger boy who she happens to run into.[21] Sometimes, though more rarely, the red hood is even non-existent.[20]
+In other tellings of the story, the wolf chases after Little Red Riding Hood. She escapes with the help of some laundresses, who spread a sheet taut over a river so she may escape. When the wolf follows Red over the bridge of cloth, the sheet is released and the wolf drowns in the river.[22] And in another version the wolf is pushed into the fire, while he is preparing the flesh of the grandmother to be eaten by the girl.[20]
+The earliest known printed version[23] was known as Le Petit Chaperon Rouge and may have had its origins in 17th-century French folklore. It was included in the collection Tales and Stories of the Past with Morals. Tales of Mother Goose (Histoires et contes du temps passé, avec des moralités. Contes de ma mère l'Oye), in 1697, by Charles Perrault. As the title implies, this version[24] is both more sinister and more overtly moralized than the later ones. The redness of the hood, which has been given symbolic significance in many interpretations of the tale, was a detail introduced by Perrault.[25]
+The story had as its subject an "attractive, well-bred young lady", a village girl of the country being deceived into giving a wolf she encountered the information he needed to find her grandmother's house successfully and eat the old woman while at the same time avoiding being noticed by woodcutters working in the nearby forest. Then he proceeded to lay a trap for Red Riding Hood. Little Red Riding Hood ends up being asked to climb into the bed before being eaten by the wolf, where the story ends. The wolf emerges the victor of the encounter and there is no happy ending.
+Charles Perrault explained the 'moral' at the end of the tale:[26] so that no doubt is left to his intended meaning:
+From this story one learns that children, especially young lasses, pretty, courteous and well-bred, do very wrong to listen to strangers, And it is not an unheard thing if the Wolf is thereby provided with his dinner. I say Wolf, for all wolves are not of the same sort; there is one kind with an amenable disposition – neither noisy, nor hateful, nor angry, but tame, obliging and gentle, following the young maids in the streets, even into their homes. Alas! Who does not know that these gentle wolves are of all such creatures the most dangerous!
+This, the presumed original version of the tale was written for the late seventeenth-century French court of King Louis XIV. This audience, whom the King entertained with extravagant parties, presumably would take from the story the intended meaning.
+In the 19th century two separate German versions were retold to Jacob Grimm and his younger brother Wilhelm Grimm, known as the Brothers Grimm, the first by Jeanette Hassenpflug (1791–1860) and the second by Marie Hassenpflug (1788–1856). The brothers turned the first version to the main body of the story and the second into a sequel of it. The story as Rotkäppchen was included in the first edition of their collection Kinder- und Hausmärchen (Children's and Household Tales (1812)).[27]
+The earlier parts of the tale agree so closely with Perrault's variant that it is almost certainly the source of the tale.[28] However, they modified the ending; this version had the little girl and her grandmother saved by a huntsman who was after the wolf's skin; this ending is identical to that in the tale "The Wolf and the Seven Young Kids", which appears to be the source.[29] The second part featured the girl and her grandmother trapping and killing another wolf, this time anticipating his moves based on their experience with the previous one. The girl did not leave the path when the wolf spoke to her, her grandmother locked the door to keep it out, and when the wolf lurked, the grandmother had Little Red Riding Hood put a trough under the chimney and fill it with water that sausages had been cooked in; the smell lured the wolf down, and it drowned.[30]
+The Brothers further revised the story in later editions and it reached the above-mentioned final and better-known version in the 1857 edition of their work.[31] It is notably tamer than the older stories which contained darker themes.
+Numerous authors have rewritten or adapted this tale.
+Andrew Lang included a variant called "The True History of Little Goldenhood"[32] in The Red Fairy Book (1890). He derived it from the works of Charles Marelles,[33] in Contes of Charles Marelles. This version explicitly states that the story had been mistold earlier. The girl is saved, but not by the huntsman; when the wolf tries to eat her, its mouth is burned by the golden hood she wears, which is enchanted.
+James N. Barker wrote a variation of Little Red Riding Hood in 1827 as an approximately 1000-word story. It was later reprinted in 1858 in a book of collected stories edited by William E Burton, called the Cyclopedia of Wit and Humor. The reprint also features a wood engraving of a clothed wolf on a bended knee holding Little Red Riding Hood's hand.
+In the 20th century, the popularity of the tale appeared to snowball, with many new versions being written and produced, especially in the wake of Freudian analysis, deconstruction and feminist critical theory. (See "Modern uses and adaptations" below.) This trend has also led to a number of academic texts being written that focus on Little Red Riding Hood, including works by Alan Dundes and Jack Zipes.
+Apart from the overt warning about talking to strangers, there are many interpretations of the classic fairy tale, many of them sexual.[34] Some are listed below.
+Folklorists and cultural anthropologists, such as P. Saintyves and Edward Burnett Tylor, saw "Little Red Riding Hood" in terms of solar myths and other naturally occurring cycles. Her red hood could represent the bright sun which is ultimately swallowed by the terrible night (the wolf), and the variations in which she is cut out of the wolf's belly represent the dawn.[35] In this interpretation, there is a connection between the wolf of this tale and Sköll, the wolf in Norse mythology that will swallow the personified Sun at Ragnarök, or Fenrir.[36] Alternatively, the tale could be about the season of spring or the month of May, escaping the winter.[37]
+The tale has been interpreted as a puberty rite, stemming from a prehistoric origin (sometimes an origin stemming from a previous matriarchal era).[38] The girl, leaving home, enters a liminal state and by going through the acts of the tale, is transformed into an adult woman by the act of coming out of the wolf's stomach.[39]
+Bruno Bettelheim, in The Uses of Enchantment: The Meaning and Importance of Fairy Tales (1976), recast the Little Red Riding Hood motif in terms of classic Freudian analysis, that shows how fairy tales educate, support, and liberate children's emotions. The motif of the huntsman cutting open the wolf he interpreted as a "rebirth"; the girl who foolishly listened to the wolf has been reborn as a new person.[40]
+The poem "Þrymskviða" from the Poetic Edda mirrors some elements of Red Riding Hood. Loki's explanations for the strange behavior of "Freyja" (actually Thor disguised as Freyja) mirror the wolf's explanations for his strange appearance. The red hood has often been given great importance in many interpretations, with a significance from the dawn to blood.[41]
+A sexual analysis of the tale may also include negative connotations in terms of rape or abduction. In Against Our Will, Susan Brownmiller describes the fairy tale as a description of rape.[42] However, many revisionist retellings choose to focus on empowerment, and depict Little Red Riding Hood or the grandmother successfully defending herself against the wolf.[43]
+Such tellings bear some similarity to the "animal bridegroom" tales, such as Beauty and the Beast or The Frog Prince, but where the heroines of those tales revert the hero to a prince, these tellings of Little Red Riding Hood reveal to the heroine that she has a wild nature like the hero's.[44] These interpretations refuse to characterize Little Red Riding Hood as a victim; these are tales of female empowerment.

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+Mesothelae
+Opisthothelae
+ See Spider taxonomy.
+Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom,[2] and spinnerets that extrude silk.[3] They are the largest order of arachnids and rank seventh in total species diversity among all orders of organisms.[4] Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of July 2019[update], at least 48,200 spider species, and 120 families have been recorded by taxonomists.[1] However, there has been dissension within the scientific community as to how all these families should be classified, as evidenced by the over 20 different classifications that have been proposed since 1900.[5]
+Anatomically, spiders (as with all arachnids) differ from other arthropods in that the usual body segments are fused into two tagmata, the prosoma, or cephalothorax, and opisthosoma, or abdomen, and joined by a small, cylindrical pedicel (however, as there is currently neither paleontological nor embryological evidence that spiders ever had a separate thorax-like division, there exists an argument against the validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly, arguments can be formed against use of the term abdomen, as the opisthosoma of all spiders contains a heart and respiratory organs, organs atypical of an abdomen[6]). Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.
+Their abdomens bear appendages that have been modified into spinnerets that extrude silk from up to six types of glands. Spider webs vary widely in size, shape and the amount of sticky thread used. It now appears that the spiral orb web may be one of the earliest forms, and spiders that produce tangled cobwebs are more abundant and diverse than orb-web spiders. Spider-like arachnids with silk-producing spigots appeared in the Devonian period about 386 million years ago, but these animals apparently lacked spinnerets. True spiders have been found in Carboniferous rocks from 318 to 299 million years ago, and are very similar to the most primitive surviving suborder, the Mesothelae. The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, before 200 million years ago.
+The species Bagheera kiplingi was described as herbivorous in 2008,[7] but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. It is estimated that the world's 25 million tons of spiders kill 400–800 million tons of prey per year.[8] Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, so they liquefy their food by flooding it with digestive enzymes. They also grind food with the bases of their pedipalps, as arachnids do not have the mandibles that crustaceans and insects have.
+To avoid being eaten by the females, which are typically much larger, male spiders identify themselves to potential mates by a variety of complex courtship rituals. Males of most species survive a few matings, limited mainly by their short life spans. Females weave silk egg-cases, each of which may contain hundreds of eggs. Females of many species care for their young, for example by carrying them around or by sharing food with them. A minority of species are social, building communal webs that may house anywhere from a few to 50,000 individuals. Social behavior ranges from precarious toleration, as in the widow spiders, to co-operative hunting and food-sharing. Although most spiders live for at most two years, tarantulas and other mygalomorph spiders can live up to 25 years in captivity.
+While the venom of a few species is dangerous to humans, scientists are now researching the use of spider venom in medicine and as non-polluting pesticides. Spider silk provides a combination of lightness, strength and elasticity that is superior to that of synthetic materials, and spider silk genes have been inserted into mammals and plants to see if these can be used as silk factories. As a result of their wide range of behaviors, spiders have become common symbols in art and mythology symbolizing various combinations of patience, cruelty and creative powers. An abnormal fear of spiders is called arachnophobia.
+1: pedipalp
+ 2: trichobothria
+ 3: carapace of prosoma (cephalothorax)
+ 4: opisthosoma (abdomen)
+ 5: eyes – AL (anterior lateral)
+    AM (anterior median)
+    PL (posterior lateral)
+    PM (posterior median)
+Leg segments:
+ 6: costa
+ 7: trochanter
+ 8: patella
+ 9: tibia
+10: metatarsus
+11: tarsus
+13: claw
+14: chelicera
+15: sternum of prosoma
+16: pedicel (also called pedicle)
+17: book lung sac
+18: book lung stigma
+19: epigastric fold
+20: epigyne
+21: anterior spinneret
+22: posterior spinneret
+Spiders are chelicerates and therefore arthropods.[9] As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo.[10] Being chelicerates, their bodies consist of two tagmata, sets of segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete fusion of the segments that in an insect would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma.[9] In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel.[11] The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws".[10][12] The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.[9]
+Spiders and scorpions are members of one chelicerate group, the arachnids.[12] Scorpions' chelicerae have three sections and are used in feeding.[13] Spiders' chelicerae have two sections and terminate in fangs that are generally venomous, and fold away behind the upper sections while not in use. The upper sections generally have thick "beards" that filter solid lumps out of their food, as spiders can take only liquid food.[11] Scorpions' pedipalps generally form large claws for capturing prey,[13] while those of spiders are fairly small appendages whose bases also act as an extension of the mouth; in addition, those of male spiders have enlarged last sections used for sperm transfer.[11]
+In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex carapace, while the underside is covered by two rather flat plates. The abdomen is soft and egg-shaped. It shows no sign of segmentation, except that the primitive Mesothelae, whose living members are the Liphistiidae, have segmented plates on the upper surface.[11]
+Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end.[14] However, in spiders, it occupies only the upper part of the abdomen, and blood is discharged into the hemocoel by one artery that opens at the rear end of the abdomen and by branching arteries that pass through the pedicle and open into several parts of the cephalothorax. Hence spiders have open circulatory systems.[11] The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.[12]
+Spiders have developed several different respiratory anatomies, based on book lungs, a tracheal system, or both. Mygalomorph and Mesothelae spiders have two pairs of book lungs filled with haemolymph, where openings on the ventral surface of the abdomen allow air to enter and diffuse oxygen. This is also the case for some basal araneomorph spiders, like the family Hypochilidae, but the remaining members of this group have just the anterior pair of book lungs intact while the posterior pair of breathing organs are partly or fully modified into tracheae, through which oxygen is diffused into the haemolymph or directly to the tissue and organs.[11] The tracheal system has most likely evolved in small ancestors to help resist desiccation.[12] The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets.[11] Spiders that have tracheae generally have higher metabolic rates and better water conservation.[15] Spiders are ectotherms, so environmental temperatures affect their activity.[16]
+Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae.[11] The families Uloboridae and Holarchaeidae, and some Liphistiidae spiders, have lost their venom glands, and kill their prey with silk instead.[17] Like most arachnids, including scorpions,[12] spiders have a narrow gut that can only cope with liquid food and two sets of filters to keep solids out.[11] They use one of two different systems of external digestion. Some pump digestive enzymes from the midgut into the prey and then suck the liquified tissues of the prey into the gut, eventually leaving behind the empty husk of the prey. Others grind the prey to pulp using the chelicerae and the bases of the pedipalps, while flooding it with enzymes; in these species, the chelicerae and the bases of the pedipalps form a preoral cavity that holds the food they are processing.[11]
+The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The midgut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.[11]
+Most spiders convert nitrogenous waste products into uric acid, which can be excreted as a dry material. Malphigian tubules ("little tubes") extract these wastes from the blood in the hemocoel and dump them into the cloacal chamber, from which they are expelled through the anus.[11] Production of uric acid and its removal via Malphigian tubules are a water-conserving feature that has evolved independently in several arthropod lineages that can live far away from water,[18] for example the tubules of insects and arachnids develop from completely different parts of the embryo.[12] However, a few primitive spiders, the suborder Mesothelae and infraorder Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"),[11] which use large amounts of water to excrete nitrogenous waste products as ammonia.[18]
+The basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia.[19] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen;[11][12][19] in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.[15]
+Despite the relatively small central nervous system, some spiders (like Portia) exhibit complex behaviour, including the ability to use a trial-and-error approach.[20][21]
+Spiders have primarily four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another.[11] The principal pair at the front are of the type called pigment-cup ocelli ("little eyes"), which in most arthropods are only capable of detecting the direction from which light is coming, using the shadow cast by the walls of the cup. However, in spiders these eyes are capable of forming images.[22][23] The other pairs, called secondary eyes, are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the principal eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torchlight reflected from the tapeta. On the other hand, jumping spiders' secondary eyes have no tapeta.[11]
+Other differences between the principal and secondary eyes are that the latter have rhabdomeres that point away from incoming light, just like in vertebrates, while the arrangement is the opposite in the former. The principal eyes are also the only ones with eye muscles, allowing them to move the retina. Having no muscles, the secondary eyes are immobile.[24]
+Some jumping spiders' visual acuity exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects; in fact the human eye is only about five times sharper than a jumping spider's. They achieve this by a telephotographic series of lenses, a four-layer retina and the ability to swivel their eyes and integrate images from different stages in the scan. The downside is that the scanning and integrating processes are relatively slow.[20]
+There are spiders with a reduced number of eyes. Of these, those with six eyes (such as Periegops suterii) are the most numerous and are missing a pair of eyes on the anterior median line;[25] other species have four eyes and some just two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.
+As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae.[22] An adult Araneus may have up to 1,000 such chemosensitive setae, most on the tarsi of the first pair of legs. Males have more chemosensitive bristles on their pedipalps than females. They have been shown to be responsive to sex pheromones produced by females, both contact and air-borne.[26] The jumping spider Evarcha culicivora uses the scent of blood from mammals and other vertebrates, which is obtained by capturing blood-filled mosquitoes, to attract the opposite sex. Because they are able to tell the sexes apart, it is assumed the blood scent is mixed with pheromones.[27] Spiders also have in the joints of their limbs slit sensillae that detect force and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.[11]
+Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' proprioceptors, sensors that report the force exerted by muscles and the degree of bending in the body and joints, are well-understood. On the other hand, little is known about what other internal sensors spiders or other arthropods may have.[22]
+Each of the eight legs of a spider consists of seven distinct parts. The part closest to and attaching the leg to the cephalothorax is the coxa; the next segment is the short trochanter that works as a hinge for the following long segment, the femur; next is the spider's knee, the patella, which acts as the hinge for the tibia; the metatarsus is next, and it connects the tibia to the tarsus (which may be thought of as a foot of sorts); the tarsus ends in a claw made up of either two or three points, depending on the family to which the spider belongs. Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors.[28] The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter).[29] As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up.[11] Spiders can generate pressures up to eight times their resting level to extend their legs,[30] and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs.[11] Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.[29]
+Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine bristles between the paired claws at the tips of their legs. These tufts, known as scopulae, consist of bristles whose ends are split into as many as 1,000 branches, and enable spiders with scopulae to walk up vertical glass and upside down on ceilings. It appears that scopulae get their grip from contact with extremely thin layers of water on surfaces.[11] Spiders, like most other arachnids, keep at least four legs on the surface while walking or running.[31]
+The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk.[11]
+Silk is mainly composed of a protein very similar to that used in insect silk. It is initially a liquid, and hardens not by exposure to air but as a result of being drawn out, which changes the internal structure of the protein.[32] It is similar in tensile strength to nylon and biological materials such as chitin, collagen and cellulose, but is much more elastic. In other words, it can stretch much further before breaking or losing shape.[11]
+Some spiders have a cribellum, a modified spinneret with up to 40,000 spigots, each of which produces a single very fine fiber. The fibers are pulled out by the calamistrum, a comblike set of bristles on the jointed tip of the cribellum, and combined into a composite woolly thread that is very effective in snagging the bristles of insects. The earliest spiders had cribella, which produced the first silk capable of capturing insects, before spiders developed silk coated with sticky droplets. However, most modern groups of spiders have lost the cribellum.[11]
+Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a "safety rope"; for nest-building; and as "parachutes" by the young of some species.[11]
+Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods,[33] male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs onto which they ejaculate and then transfer the sperm to special syringe-styled structures, palpal bulbs or palpal organs, borne on the tips of the pedipalps of mature males. When a male detects signs of a female nearby he checks whether she is of the same species and whether she is ready to mate; for example in species that produce webs or "safety ropes", the male can identify the species and sex of these objects by "smell".[11]
+Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female via one or two openings on the underside of her abdomen.[11]
+Female spiders' reproductive tracts are arranged in one of two ways. The ancestral arrangement ("haplogyne" or "non-entelegyne") consists of a single genital opening, leading to two seminal receptacles (spermathecae) in which females store sperm. In the more advanced arrangement ("entelegyne" ), there are two further openings leading directly to the spermathecae, creating a "flow through" system rather than a "first-in first-out" one. Eggs are as a general rule only fertilized during oviposition when the stored sperm is released from its chamber, rather than in the ovarian cavity.[34] A few exceptions exist, such as Parasteatoda tepidariorum. In these species the female appears to be able to activate the dormant sperm before oviposition, allowing them to migrate to the ovarian cavity where fertilization occurs.[35][36][37] The only known example of direct fertilization between male and female is an Israeli spider, Harpactea sadistica, which has evolved traumatic insemination. In this species the male will penetrate its pedipalps through the female's body wall and inject his sperm directly into her ovaries, where the embryos inside the fertilized eggs will start to develop before being laid.[38]
+Males of the genus Tidarren amputate one of their palps before maturation and enter adult life with one palp only. The palps are 20% of the male's body mass in this species, and detaching one of the two improves mobility. In the Yemeni species Tidarren argo, the remaining palp is then torn off by the female. The separated palp remains attached to the female's epigynum for about four hours and apparently continues to function independently. In the meantime, the female feeds on the palpless male.[39] In over 60% of cases, the female of the Australian redback spider kills and eats the male after it inserts its second palp into the female's genital opening; in fact, the males co-operate by trying to impale themselves on the females' fangs. Observation shows that most male redbacks never get an opportunity to mate, and the "lucky" ones increase the likely number of offspring by ensuring that the females are well-fed.[40] However, males of most species survive a few matings, limited mainly by their short life spans. Some even live for a while in their mates' webs.[41]
+The tiny male of the Golden orb weaver (Trichonephila clavipes) (near the top of the leaf) is protected from the female by producing the right vibrations in the web, and may be too small to be worth eating.
+Orange spider egg sac hanging from ceiling
+Gasteracantha mammosa spiderlings next to their eggs capsule
+Wolf spider carrying its young on its abdomen
+Females lay up to 3,000 eggs in one or more silk egg sacs,[11] which maintain a fairly constant humidity level.[41] In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.[11]
+Baby spiders pass all their larval stages inside the egg and hatch as spiderlings, very small and sexually immature but similar in shape to adults. Some spiders care for their young, for example a wolf spider's brood clings to rough bristles on the mother's back,[11] and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.[41]
+Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch.[42] In some species males mate with newly-molted females, which are too weak to be dangerous to the males.[41] Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years,[11][43] and an Australian female trapdoor spider was documented to have lived in the wild for 43 years, dying of a parasitic wasp attack.[44]
+Spiders occur in a large range of sizes. The smallest, Patu digua from Colombia, are less than 0.37 mm (0.015 in) in body length. The largest and heaviest spiders occur among tarantulas, which can have body lengths up to 90 mm (3.5 in) and leg spans up to 250 mm (9.8 in).[45]
+Only three classes of pigment (ommochromes, bilins and guanine) have been identified in spiders, although other pigments have been detected but not yet characterized. Melanins, carotenoids and pterins, very common in other animals, are apparently absent. In some species, the exocuticle of the legs and prosoma is modified by a tanning process, resulting in a brown coloration.[46]
+Bilins are found, for example, in Micrommata virescens, resulting in its green color. Guanine is responsible for the white markings of the European garden spider Araneus diadematus. It is in many species accumulated in specialized cells called guanocytes. In genera such as Tetragnatha, Leucauge, Argyrodes or Theridiosoma, guanine creates their silvery appearance. While guanine is originally an end-product of protein metabolism, its excretion can be blocked in spiders, leading to an increase in its storage.[46] Structural colors occur in some species, which are the result of the diffraction, scattering or interference of light, for example by modified setae or scales. The white prosoma of Argiope results from bristles reflecting the light, Lycosa and Josa both have areas of modified cuticle that act as light reflectors.[46]
+While in many spiders color is fixed throughout their lifespan, in some groups, color may be variable in response to environmental and internal conditions.[46] Choice of prey may be able to alter the color of spiders. For example, the abdomen of Theridion grallator will become orange if the spider ingests certain species of Diptera and adult Lepidoptera, but if it consumes Homoptera or larval Lepidoptera, then the abdomen becomes green.[47] Environmentally induced color changes may be morphological (occurring over several days) or physiological (occurring near instantly). Morphological changes require pigment synthesis and degradation. In contrast to this, physiological changes occur by changing the position of pigment-containing cells.[46] An example of morphological color changes is background matching. Misumena vatia for instance can change its body color to match the substrate it lives on which makes it more difficult to be detected by prey.[48] An example of physiological color change is observed in Cyrtophora cicatrosa, which can change its body color from white to brown near instantly.[46]
+Although spiders are generally regarded as predatory, the jumping spider Bagheera kiplingi gets over 90% of its food from fairly solid plant material produced by acacias as part of a mutually beneficial relationship with a species of ant.[49]
+Juveniles of some spiders in the families Anyphaenidae, Corinnidae, Clubionidae, Thomisidae and Salticidae feed on plant nectar. Laboratory studies show that they do so deliberately and over extended periods, and periodically clean themselves while feeding. These spiders also prefer sugar solutions to plain water, which indicates that they are seeking nutrients. Since many spiders are nocturnal, the extent of nectar consumption by spiders may have been underestimated. Nectar contains amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.[50]
+Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.[50]
+The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.[11]
+Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it.[51] A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.[11]
+Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus Hyptiotes and the family Theridiosomatidae stretch their webs and then release them when prey strike them, but do not actively move their webs. Those of the family Deinopidae weave even smaller webs, hold them outstretched between their first two pairs of legs, and lunge and push the webs as much as twice their own body length to trap prey, and this move may increase the webs' area by a factor of up to ten. Experiments have shown that Deinopis spinosus has two different techniques for trapping prey: backwards strikes to catch flying insects, whose vibrations it detects; and forward strikes to catch ground-walking prey that it sees. These two techniques have also been observed in other deinopids. Walking insects form most of the prey of most deinopids, but one population of Deinopis subrufa appears to live mainly on tipulid flies that they catch with the backwards strike.[52]
+Mature female bolas spiders of the genus Mastophora build "webs" that consist of only a single "trapeze line", which they patrol. They also construct a bolas made of a single thread, tipped with a large ball of very wet sticky silk. They emit chemicals that resemble the pheromones of moths, and then swing the bolas at the moths. Although they miss on about 50% of strikes, they catch about the same weight of insects per night as web-weaving spiders of similar size. The spiders eat the bolas if they have not made a kill in about 30 minutes, rest for a while, and then make new bolas.[53][54] Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.[55]
+The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush predators that lurk in burrows, often closed by trapdoors and often surrounded by networks of silk threads that alert these spiders to the presence of prey.[15] Other ambush predators do without such aids, including many crab spiders,[11] and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking.[46] Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.[11]
+Some jumping spiders of the genus Portia hunt other spiders in ways that seem intelligent,[20] outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species.[56] However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.[20]
+Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective bristles to resemble the shiny bodies of ants. In some spider species, males and females mimic different ant species, as female spiders are usually much larger than males. Ant-mimicking spiders also modify their behavior to resemble that of the target species of ant; for example, many adopt a zig-zag pattern of movement, ant-mimicking jumping spiders avoid jumping, and spiders of the genus Synemosyna walk on the outer edges of leaves in the same way as Pseudomyrmex. Ant mimicry in many spiders and other arthropods may be for protection from predators that hunt by sight, including birds, lizards and spiders. However, several ant-mimicking spiders prey either on ants or on the ants' "livestock", such as aphids. When at rest, the ant-mimicking crab spider Amyciaea does not closely resemble Oecophylla, but while hunting it imitates the behavior of a dying ant to attract worker ants. After a kill, some ant-mimicking spiders hold their victims between themselves and large groups of ants to avoid being attacked.[57]
+There is strong evidence that spiders' coloration is camouflage that helps them to evade their major predators, birds and parasitic wasps, both of which have good color vision. Many spider species are colored so as to merge with their most common backgrounds, and some have disruptive coloration, stripes and blotches that break up their outlines. In a few species, such as the Hawaiian happy-face spider, Theridion grallator, several coloration schemes are present in a ratio that appears to remain constant, and this may make it more difficult for predators to recognize the species. Most spiders are insufficiently dangerous or unpleasant-tasting for warning coloration to offer much benefit. However, a few species with powerful venom, large jaws or irritant bristles have patches of warning colors, and some actively display these colors when threatened.[46][58]
+Many of the family Theraphosidae, which includes tarantulas and baboon spiders, have urticating hairs on their abdomens and use their legs to flick them at attackers. These bristles are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom.[59] A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles.[60] The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.[61]
+A few spider species that build webs live together in large colonies and show social behavior, although not as complex as in social insects. Anelosimus eximius (in the family Theridiidae) can form colonies of up to 50,000 individuals.[62] The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social.[63] Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food.[64] Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae).[65] Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this.[66] The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings.[49] Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.[67]
+There is no consistent relationship between the classification of spiders and the types of web they build: species in the same genus may build very similar or significantly different webs. Nor is there much correspondence between spiders' classification and the chemical composition of their silks. Convergent evolution in web construction, in other words use of similar techniques by remotely related species, is rampant. Orb web designs and the spinning behaviors that produce them are the best understood. The basic radial-then-spiral sequence visible in orb webs and the sense of direction required to build them may have been inherited from the common ancestors of most spider groups.[68] However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a subgroup that evolved from orb-web spiders, and non-orb spiders have over 40% more species and are four times as abundant as orb-web spiders. Their greater success may be because sphecid wasps, which are often the dominant predators of spiders, much prefer to attack spiders that have flat webs.[69]
+About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey.[68]
+The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.[68]
+Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the backlighting from the sky; enabling oscillations to catch insects in slow horizontal flight. However, there is no single explanation for the common use of horizontal orb webs.[68]
+Spiders often attach highly visible silk bands, called decorations or stabilimenta, to their webs. Field research suggests that webs with more decorative bands captured more prey per hour.[70] However, a laboratory study showed that spiders reduce the building of these decorations if they sense the presence of predators.[71]
+There are several unusual variants of orb web, many of them convergently evolved, including: attachment of lines to the surface of water, possibly to trap insects in or on the surface; webs with twigs through their centers, possibly to hide the spiders from predators; "ladderlike" webs that appear most effective in catching moths. However, the significance of many variations is unclear.[68]
+In 1973, Skylab 3 took two orb-web spiders into space to test their web-spinning capabilities in zero gravity. At first, both produced rather sloppy webs, but they adapted quickly.[72]
+Members of the family Theridiidae weave irregular, tangled, three-dimensional webs, popularly known as cobwebs. There seems to be an evolutionary trend towards a reduction in the amount of sticky silk used, leading to its total absence in some species. The construction of cobwebs is less stereotyped than that of orb-webs, and may take several days.[69]
+The Linyphiidae generally make horizontal but uneven sheets, with tangles of stopping threads above. Insects that hit the stopping threads fall onto the sheet or are shaken onto it by the spider, and are held by sticky threads on the sheet until the spider can attack from below.[73]
+Although the fossil record of spiders is considered poor,[74] almost 1000 species have been described from fossils.[75] Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber.[75] The oldest known amber that contains fossil arthropods dates from 130 million years ago in the Early Cretaceous period. In addition to preserving spiders' anatomy in very fine detail, pieces of amber show spiders mating, killing prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached;[76] the oldest fossil web found so far is 100 million years old.[77] Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.[76]
+The oldest known exclusively terrestrial arachnid is the trigonotarbid Palaeotarbus jerami, from about 420 million years ago in the Silurian period, and had a triangular cephalothorax and segmented abdomen, as well as eight legs and a pair of pedipalps.[78] Attercopus fimbriunguis, from 386 million years ago in the Devonian period, bears the earliest known silk-producing spigots, and was therefore hailed as a spider at the time of its discovery.[79] However, these spigots may have been mounted on the underside of the abdomen rather than on spinnerets, which are modified appendages and whose mobility is important in the building of webs. Hence Attercopus and the similar Permian arachnid Permarachne may not have been true spiders, and probably used silk for lining nests or producing egg cases rather than for building webs.[3] The largest known fossil spider as of 2011 is the araneid Nephila jurassica, from about 165 million years ago, recorded from Daohuogo, Inner Mongolia in China.[80] Its body length is almost 25 mm, (i.e., almost one inch).
+Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae.[79]
+The mesothelid Paleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets.[81] Although the Permian period 299 to 251 million years ago saw rapid diversification of flying insects, there are very few fossil spiders from this period.[79]
+The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appear in the Triassic well before 200 million years ago. Some Triassic mygalomorphs appear to be members of the family Hexathelidae, whose modern members include the notorious Sydney funnel-web spider, and their spinnerets appear adapted for building funnel-shaped webs to catch jumping insects. Araneomorphae account for the great majority of modern spiders, including those that weave the familiar orb-shaped webs. The Jurassic and Cretaceous periods provide a large number of fossil spiders, including representatives of many modern families.[79]
+Xiphosura (horseshoe crabs)
+Eurypterida†
+Chasmataspidida†
+Scorpiones
+Opiliones (harvestmen)
+Pseudoscorpiones
+Solifugae (sun spiders)
+Palpigradi (microwhip scorpions)
+Trigonotarbida†
+Araneae (spiders)
+Haptopoda†
+Amblypygi (whip spiders)
+Thelyphonida (whip scorpions)
+Schizomida
+Ricinulei (hooded tickspiders)
+Anactinotrichida
+Acariformes (mites)
+It is now agreed that spiders (Araneae) are monophyletic (i.e., members of a group of organisms that form a clade, consisting of a last common ancestor and all of its descendants).[83] There has been debate about what their closest evolutionary relatives are, and how all of these evolved from the ancestral chelicerates, which were marine animals. The cladogram on the right is based on J. W. Shultz' analysis (2007). Other views include proposals that: scorpions are more closely related to the extinct marine scorpioid eurypterids than to spiders; spiders and Amblypygi are a monophyletic group. The appearance of several multi-way branchings in the tree on the right shows that there are still uncertainties about relationships between the groups involved.[83]
+Arachnids lack some features of other chelicerates, including backward-pointing mouths and gnathobases ("jaw bases") at the bases of their legs;[83] both of these features are part of the ancestral arthropod feeding system.[84] Instead, they have mouths that point forwards and downwards, and all have some means of breathing air.[83] Spiders (Araneae) are distinguished from other arachnid groups by several characteristics, including spinnerets and, in males, pedipalps that are specially adapted for sperm transfer.[85]
+Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two infraorders, Mygalomorphae and Araneomorphae. Over 48,000 living species of spiders (order Araneae) have been identified and as of 2019 grouped into 120 families and about 4,100 genera by arachnologists.[1]
+The only living members of the primitive Mesothelae are the family Liphistiidae, found only in Southeast Asia, China, and Japan.[85] Most of the Liphistiidae construct silk-lined burrows with thin trapdoors, although some species of the genus Liphistius build camouflaged silk tubes with a second trapdoor as an emergency exit. Members of the genus Liphistius run silk "tripwires" outwards from their tunnels to help them detect approaching prey, while those of the genus Heptathela do not and instead rely on their built-in vibration sensors.[88] Spiders of the genus Heptathela have no venom glands, although they do have venom gland outlets on the fang tip.[89]
+The extinct families Arthrolycosidae, found in Carboniferous and Permian rocks, and Arthromygalidae, so far found only in Carboniferous rocks, have been classified as members of the Mesothelae.[90]
+The Mygalomorphae, which first appeared in the Triassic period,[79] are generally heavily built and ″hairy″, with large, robust chelicerae and fangs (technically, spiders do not have true hairs, but rather setae).[91][85] Well-known examples include tarantulas, ctenizid trapdoor spiders and the Australasian funnel-web spiders.[11] Most spend the majority of their time in burrows, and some run silk tripwires out from these, but a few build webs to capture prey. However, mygalomorphs cannot produce the pirifom silk that the Araneomorphae use as an instant adhesive to glue silk to surfaces or to other strands of silk, and this makes web construction more difficult for mygalomorphs. Since mygalomorphs rarely "balloon" by using air currents for transport, their populations often form clumps.[85] In addition to arthropods, some mygalomorphs are known to prey on frogs, small mammals, lizards, snakes, snails, and small birds.[92][93]
+In addition to accounting for over 90% of spider species, the Araneomorphae, also known as the "true spiders", include orb-web spiders, the cursorial wolf spiders, and jumping spiders,[85] as well as the only known herbivorous spider, Bagheera kiplingi.[49] They are distinguished by having fangs that oppose each other and cross in a pinching action, in contrast to the Mygalomorphae, which have fangs that are nearly parallel in alignment.[94]
+Although spiders are widely feared, only a few species are dangerous to people.[96] Spiders will only bite humans in self-defense, and few produce worse effects than a mosquito bite or bee sting.[97] Most of those with medically serious bites, such as recluse spiders (genus Loxosceles) and widow spiders (genus Latrodectus), would rather flee and bite only when trapped, although this can easily arise by accident.[98][99] The defensive tactics of Australian funnel-web spiders (family Atracidae) include fang display. Their venom, although they rarely inject much, has resulted in 13 attributed human deaths over 50 years.[100] They have been deemed to be the world's most dangerous spiders on clinical and venom toxicity grounds,[96] though this claim has also been attributed to the Brazilian wandering spider (genus Phoneutria).[101]
+There were about 100 reliably reported deaths from spider bites in the 20th century,[102] compared to about 1,500 from jellyfish stings.[103] Many alleged cases of spider bites may represent incorrect diagnoses,[104] which would make it more difficult to check the effectiveness of treatments for genuine bites.[105] A review published in 2016 agreed with this conclusion, showing that 78% of 134 published medical case studies of supposed spider bites did not meet the necessary criteria for a spider bite to be verified. In the case of the two genera with the highest reported number of bites, Loxosceles and Latrodectus, spider bites were not verified in over 90% of the reports. Even when verification had occurred, details of the treatment and its effects were often lacking.[106]
+Spider venoms may be a less polluting alternative to conventional pesticides, as they are deadly to insects but the great majority are harmless to vertebrates. Australian funnel web spiders are a promising source, as most of the world's insect pests have had no opportunity to develop any immunity to their venom, and funnel web spiders thrive in captivity and are easy to "milk". It may be possible to target specific pests by engineering genes for the production of spider toxins into viruses that infect species such as cotton bollworms.[107]
+The Ch'ol Maya use a beverage created from the tarantula species Brachypelma vagans for the treatment of a condition they term 'tarantula wind', the symptoms of which include chest pain, asthma and coughing.[108]
+Possible medical uses for spider venoms are being investigated, for the treatment of cardiac arrhythmia,[109] Alzheimer's disease,[110] strokes,[111] and erectile dysfunction.[112]
+The peptide GsMtx-4, found in the venom of Brachypelma vagans, is being researched to determine whether or not it could effectively be used for the treatment of cardiac arrhythmia, muscular dystrophy or glioma.[108]
+Because spider silk is both light and very strong, attempts are being made to produce it in goats' milk and in the leaves of plants, by means of genetic engineering.[113][114]
+Spiders can also be used as food. Cooked tarantulas are considered a delicacy in Cambodia,[115] and by the Piaroa Indians of southern Venezuela – provided the highly irritant bristles, the spiders' main defense system, are removed first.[116]
+Arachnophobia is a specific phobia—it is the abnormal fear of spiders or anything reminiscent of spiders, such as webs or spiderlike shapes. It is one of the most common specific phobias,[117][118] and some statistics show that 50% of women and 10% of men show symptoms.[119] It may be an exaggerated form of an instinctive response that helped early humans to survive,[120] or a cultural phenomenon that is most common in predominantly European societies.[121]
+Spiders have been the focus of stories and mythologies of various cultures for centuries.[122] Uttu, the ancient Sumerian goddess of weaving, was envisioned as a spider spinning her web.[123][124] According to her main myth, she resisted her father Enki's sexual advances by ensconcing herself in her web,[124] but let him in after he promised her fresh produce as a marriage gift,[124] thereby allowing him to intoxicate her with beer and rape her.[124] Enki's wife Ninhursag heard Uttu's screams and rescued her,[124] removing Enki's semen from her vagina and planting it in the ground to produce eight previously-nonexistent plants.[124] In a story told by the Roman poet Ovid in his Metamorphoses, Arachne was a Lydian girl who challenged the goddess Athena to a weaving contest.[125][126] Arachne won, but Athena destroyed her tapestry out of jealousy,[126][127] causing Arachne to hang herself.[126][127] In an act of mercy, Athena brought Arachne back to life as the first spider.[126][127] Stories about the trickster-spider Anansi are prominent in the folktales of West Africa and the Caribbean.[128]
+In some cultures, spiders have symbolized patience due to their hunting technique of setting webs and waiting for prey, as well as mischief and malice due to their venomous bites.[129] The Italian tarantella is a dance to rid the young woman of the lustful effects of a spider bite. Web-spinning also caused the association of the spider with creation myths, as they seem to have the ability to produce their own worlds.[130] Dreamcatchers are depictions of spiderwebs. The Moche people of ancient Peru worshipped nature.[131] They placed emphasis on animals and often depicted spiders in their art.[132]

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+Petroleum (pronounced /pəˈtroʊliəm/) is a naturally occurring, yellowish-black liquid found in geological formations beneath the Earth's surface. It is commonly refined into various types of fuels. Components of petroleum are separated using a technique called fractional distillation, i.e. separation of a liquid mixture into fractions differing in boiling point by means of distillation, typically using a fractionating column.
+It consists of naturally occurring  hydrocarbons of various molecular weights and may contain miscellaneous organic compounds.[1] The name petroleum covers both naturally occurring unprocessed crude oil and petroleum products that are made up of refined crude oil. A fossil fuel, petroleum is formed when large quantities of dead organisms, mostly zooplankton and algae, are buried underneath sedimentary rock and subjected to both intense heat and pressure.
+Petroleum has mostly been recovered by oil drilling (natural petroleum springs are rare). Drilling is carried out after studies of structural geology (at the reservoir scale), sedimentary basin analysis, and reservoir characterisation (mainly in terms of the porosity and permeability of geologic reservoir structures) have been completed.[2][3] It is refined and separated, most easily by distillation, into numerous consumer products, from gasoline (petrol) and kerosene to asphalt and chemical reagents (ethylene, propylene, butene[4], acrylic acid[5][6][7], para-xylene[8]) used to make plastics, pesticides and pharmaceuticals.[9] Petroleum is used in manufacturing a wide variety of materials,[10] and it is estimated that the world consumes about 95 million barrels each day.
+The use of petroleum as fuel contributes to global warming and ocean acidification. According to the UN's Intergovernmental Panel on Climate Change, without fossil fuel phase-out, including petroleum, there will be "severe, pervasive, and irreversible impacts for people and ecosystems".[11]
+The word petroleum comes from Medieval Latin petroleum (literally "rock oil"), which comes from Latin petra, "rock", (from Ancient Greek: πέτρα, romanized: petra, "rock") and Latin oleum, "oil", (from Ancient Greek: ἔλαιον, romanized: élaion, "oil").[12][13]
+The term was used in the treatise De Natura Fossilium, published in 1546 by the German mineralogist Georg Bauer, also known as Georgius Agricola.[14] In the 19th century, the term petroleum was often used to refer to mineral oils produced by distillation from mined organic solids such as cannel coal (and later oil shale) and refined oils produced from them; in the United Kingdom, storage (and later transport) of these oils were regulated by a series of Petroleum Acts, from the Petroleum Act 1863 onwards.
+Petroleum, in one form or another, has been used since ancient times, and is now important across society, including in economy, politics and technology. The rise in importance was due to the invention of the internal combustion engine, the rise in commercial aviation, and the importance of petroleum to industrial organic chemistry, particularly the synthesis of plastics, fertilisers, solvents, adhesives and pesticides.
+More than 4000 years ago, according to Herodotus and Diodorus Siculus, asphalt was used in the construction of the walls and towers of Babylon; there were oil pits near Ardericca (near Babylon), and a pitch spring on Zacynthus.[15] Great quantities of it were found on the banks of the river Issus, one of the tributaries of the Euphrates. Ancient Persian tablets indicate the medicinal and lighting uses of petroleum in the upper levels of their society.
+The use of petroleum in ancient China dates back to more than 2000 years ago. In I Ching, one of the earliest Chinese writings cites that oil in its raw state, without refining, was first discovered, extracted, and used in China in the first century BCE. In addition, the Chinese were the first to record the use of petroleum as fuel as early as the fourth century BCE.[16][17][18] By 347 CE, oil was produced from bamboo-drilled wells in China.[19][20]
+Crude oil was often distilled by Persian chemists, with clear descriptions given in Arabic handbooks such as those of Muhammad ibn Zakarīya Rāzi (Rhazes).[21] The streets of Baghdad were paved with tar, derived from petroleum that became accessible from natural fields in the region. In the 9th century, oil fields were exploited in the area around modern Baku, Azerbaijan. These fields were described by the Arab geographer Abu al-Hasan 'Alī al-Mas'ūdī in the 10th century, and by Marco Polo in the 13th century, who described the output of those wells as hundreds of shiploads.[22] Arab and Persian chemists also distilled crude oil in order to produce flammable products for military purposes. Through Islamic Spain, distillation became available in Western Europe by the 12th century.[23] It has also been present in Romania since the 13th century, being recorded as păcură.[24]
+Early British explorers to Myanmar documented a flourishing oil extraction industry based in Yenangyaung that, in 1795, had hundreds of hand-dug wells under production.[25]
+Pechelbronn (Pitch fountain) is said to be the first European site where petroleum has been explored and used. The still active Erdpechquelle, a spring where petroleum appears mixed with water has been used since 1498, notably for medical purposes. Oil sands have been mined since the 18th century.[26]
+In Wietze in lower Saxony, natural asphalt/bitumen has been explored since the 18th century.[27] Both in Pechelbronn as in Wietze, the coal industry dominated the petroleum technologies.[28]
+Chemist James Young noticed a natural petroleum seepage in the Riddings colliery at Alfreton, Derbyshire from which he distilled a light thin oil suitable for use as lamp oil, at the same time obtaining a more viscous oil suitable for lubricating machinery. In 1848, Young set up a small business refining the crude oil.[29]
+Young eventually succeeded, by distilling cannel coal at a low heat, in creating a fluid resembling petroleum, which when treated in the same way as the seep oil gave similar products. Young found that by slow distillation he could obtain a number of useful liquids from it, one of which he named "paraffine oil" because at low temperatures it congealed into a substance resembling paraffin wax.[29]
+The production of these oils and solid paraffin wax from coal formed the subject of his patent dated 17 October 1850. In 1850 Young & Meldrum and Edward William Binney entered into partnership under the title of E.W. Binney & Co. at Bathgate in West Lothian and E. Meldrum & Co. at Glasgow; their works at Bathgate were completed in 1851 and became the first truly commercial oil-works in the world with the first modern oil refinery.[30]
+The world's first oil refinery was built in 1856 by Ignacy Łukasiewicz.[31] His achievements also included the discovery of how to distill kerosene from seep oil, the invention of the modern kerosene lamp (1853), the introduction of the first modern street lamp in Europe (1853), and the construction of the world's first modern oil well (1854).[32]
+The demand for petroleum as a fuel for lighting in North America and around the world quickly grew.[33] Edwin Drake's 1859 well near Titusville, Pennsylvania, is popularly considered the first modern well. Already 1858 Georg Christian Konrad Hunäus had found a significant amount of petroleum while drilling for lignite 1858 in Wietze, Germany. Wietze later provided about 80% of the German consumption in the Wilhelminian Era.[34] The production stopped in 1963, but Wietze has hosted a Petroleum Museum since 1970.[35]
+Drake's well is probably singled out because it was drilled, not dug; because it used a steam engine; because there was a company associated with it; and because it touched off a major boom.[36] However, there was considerable activity before Drake in various parts of the world in the mid-19th century. A group directed by Major Alexeyev of the Bakinskii Corps of Mining Engineers hand-drilled a well in the Baku region in 1848.[37] There were engine-drilled wells in West Virginia in the same year as Drake's well.[38] An early commercial well was hand dug in Poland in 1853, and another in nearby Romania in 1857. At around the same time the world's first, small, oil refinery was opened at Jasło in Poland, with a larger one opened at Ploiești in Romania shortly after. Romania is the first country in the world to have had its annual crude oil output officially recorded in international statistics: 275 tonnes for 1857.[39][40]
+The first commercial oil well in Canada became operational in 1858 at Oil Springs, Ontario (then Canada West).[41] Businessman James Miller Williams dug several wells between 1855 and 1858 before discovering a rich reserve of oil four metres below ground.[42][specify] Williams extracted 1.5 million litres of crude oil by 1860, refining much of it into kerosene lamp oil. Williams's well became commercially viable a year before Drake's Pennsylvania operation and could be argued to be the first commercial oil well in North America.[43] The discovery at Oil Springs touched off an oil boom which brought hundreds of speculators and workers to the area. Advances in drilling continued into 1862 when local driller Shaw reached a depth of 62 metres using the spring-pole drilling method.[44] On January 16, 1862, after an explosion of natural gas Canada's first oil gusher came into production, shooting into the air at a recorded rate of 3,000 barrels per day.[45] By the end of the 19th century the Russian Empire, particularly the Branobel company in Azerbaijan, had taken the lead in production.[46]
+Access to oil was and still is a major factor in several military conflicts of the twentieth century, including World War II, during which oil facilities were a major strategic asset and were extensively bombed.[47] The German invasion of the Soviet Union included the goal to capture the Baku oilfields, as it would provide much needed oil-supplies for the German military which was suffering from blockades.[48] Oil exploration in North America during the early 20th century later led to the US becoming the leading producer by mid-century. As petroleum production in the US peaked during the 1960s, however, the United States was surpassed by Saudi Arabia and the Soviet Union.[49][50][51]
+In 1973, Saudi Arabia and other Arab nations imposed an oil embargo against the United States, United Kingdom, Japan and other Western nations which supported Israel in the Yom Kippur War of October 1973.[52] The embargo caused an oil crisis with many short- and long-term effects on global politics and the global economy.[53]
+Today, about 90 percent of vehicular fuel needs are met by oil. Petroleum also makes up 40 percent of total energy consumption in the United States, but is responsible for only 1 percent of electricity generation.[54] Petroleum's worth as a portable, dense energy source powering the vast majority of vehicles and as the base of many industrial chemicals makes it one of the world's most important commodities. Viability of the oil commodity is controlled by several key parameters: number of vehicles in the world competing for fuel; quantity of oil exported to the world market (Export Land Model); net energy gain (economically useful energy provided minus energy consumed); political stability of oil exporting nations; and ability to defend oil supply lines.[citation needed]
+The top three oil producing countries are Russia, Saudi Arabia and the United States.[55] In 2018, due in part to developments in hydraulic fracturing and horizontal drilling, the United States became the world's largest producer.[56][57]
+About 80 percent of the world's readily accessible reserves are located in the Middle East, with 62.5 percent coming from the Arab 5: Saudi Arabia, United Arab Emirates, Iraq, Qatar and Kuwait. A large portion of the world's total oil exists as unconventional sources, such as bitumen in Athabasca oil sands and extra heavy oil in the Orinoco Belt. While significant volumes of oil are extracted from oil sands, particularly in Canada, logistical and technical hurdles remain, as oil extraction requires large amounts of heat and water, making its net energy content quite low relative to conventional crude oil. Thus, Canada's oil sands are not expected to provide more than a few million barrels per day in the foreseeable future.[58][59][60]
+Petroleum includes not only crude oil, but all liquid, gaseous and solid hydrocarbons. Under surface pressure and temperature conditions, lighter hydrocarbons methane, ethane, propane and butane exist as gases, while pentane and heavier hydrocarbons are in the form of liquids or solids. However, in an underground oil reservoir the proportions of gas, liquid, and solid depend on subsurface conditions and on the phase diagram of the petroleum mixture.[61]
+An oil well produces predominantly crude oil, with some natural gas dissolved in it. Because the pressure is lower at the surface than underground, some of the gas will come out of solution and be recovered (or burned) as associated gas or solution gas. A gas well produces predominantly natural gas. However, because the underground temperature and pressure are higher than at the surface, the gas may contain heavier hydrocarbons such as pentane, hexane, and heptane in the gaseous state. At surface conditions these will condense out of the gas to form "natural gas condensate", often shortened to condensate. Condensate resembles gasoline in appearance and is similar in composition to some volatile light crude oils.[62][63][64]
+The proportion of light hydrocarbons in the petroleum mixture varies greatly among different oil fields, ranging from as much as 97 percent by weight in the lighter oils to as little as 50 percent in the heavier oils and bitumens.[citation needed]
+The hydrocarbons in crude oil are mostly alkanes, cycloalkanes and various aromatic hydrocarbons, while the other organic compounds contain nitrogen, oxygen and sulfur, and trace amounts of metals such as iron, nickel, copper and vanadium. Many oil reservoirs contain live bacteria.[65] The exact molecular composition of crude oil varies widely from formation to formation but the proportion of chemical elements varies over fairly narrow limits as follows:[66]
+Four different types of hydrocarbon molecules appear in crude oil. The relative percentage of each varies from oil to oil, determining the properties of each oil.[61]
+Crude oil varies greatly in appearance depending on its composition. It is usually black or dark brown (although it may be yellowish, reddish, or even greenish). In the reservoir it is usually found in association with natural gas, which being lighter forms a "gas cap" over the petroleum, and saline water which, being heavier than most forms of crude oil, generally sinks beneath it. Crude oil may also be found in a semi-solid form mixed with sand and water, as in the Athabasca oil sands in Canada, where it is usually referred to as crude bitumen. In Canada, bitumen is considered a sticky, black, tar-like form of crude oil which is so thick and heavy that it must be heated or diluted before it will flow.[68] Venezuela also has large amounts of oil in the Orinoco oil sands, although the hydrocarbons trapped in them are more fluid than in Canada and are usually called extra heavy oil. These oil sands resources are called unconventional oil to distinguish them from oil which can be extracted using traditional oil well methods. Between them, Canada and Venezuela contain an estimated 3.6 trillion barrels (570×10^9 m3) of bitumen and extra-heavy oil, about twice the volume of the world's reserves of conventional oil.[69]
+Petroleum is used mostly, by volume, for refining into fuel oil and gasoline, both important "primary energy" sources. 84 percent by volume of the hydrocarbons present in petroleum is converted into energy-rich fuels (petroleum-based fuels), including gasoline, diesel, jet, heating, and other fuel oils, and liquefied petroleum gas.[70] The lighter grades of crude oil produce the best yields of these products, but as the world's reserves of light and medium oil are depleted, oil refineries are increasingly having to process heavy oil and bitumen, and use more complex and expensive methods to produce the products required. Because heavier crude oils have too much carbon and not enough hydrogen, these processes generally involve removing carbon from or adding hydrogen to the molecules, and using fluid catalytic cracking to convert the longer, more complex molecules in the oil to the shorter, simpler ones in the fuels.[citation needed]
+Due to its high energy density, easy transportability and relative abundance, oil has become the world's most important source of energy since the mid-1950s. Petroleum is also the raw material for many chemical products, including pharmaceuticals, solvents, fertilizers, pesticides, and plastics; the 16 percent not used for energy production is converted into these other materials. Petroleum is found in porous rock formations in the upper strata of some areas of the Earth's crust. There is also petroleum in oil sands (tar sands). Known oil reserves are typically estimated at around 190 km3 (1.2 trillion (short scale) barrels) without oil sands,[71] or 595 km3 (3.74 trillion barrels) with oil sands.[72] Consumption is currently around 84 million barrels (13.4×10^6 m3) per day, or 4.9 km3 per year, yielding a remaining oil supply of only about 120 years, if current demand remains static.[73]More recent studies, however, put the number at around 50 years[74][75].
+Petroleum is a mixture of a very large number of different hydrocarbons; the most commonly found molecules are alkanes (paraffins), cycloalkanes (naphthenes), aromatic hydrocarbons, or more complicated chemicals like asphaltenes. Each petroleum variety has a unique mix of molecules, which define its physical and chemical properties, like color and viscosity.
+The alkanes, also known as paraffins, are saturated hydrocarbons with straight or branched chains which contain only carbon and hydrogen and have the general formula CnH2n+2. They generally have from 5 to 40 carbon atoms per molecule, although trace amounts of shorter or longer molecules may be present in the mixture.
+The alkanes from pentane (C5H12) to octane (C8H18) are refined into gasoline, the ones from nonane (C9H20) to hexadecane (C16H34) into diesel fuel, kerosene and jet fuel. Alkanes with more than 16 carbon atoms can be refined into fuel oil and lubricating oil. At the heavier end of the range, paraffin wax is an alkane with approximately 25 carbon atoms, while asphalt has 35 and up, although these are usually cracked by modern refineries into more valuable products. The shortest molecules, those with four or fewer carbon atoms, are in a gaseous state at room temperature. They are the petroleum gases. Depending on demand and the cost of recovery, these gases are either flared off, sold as liquefied petroleum gas under pressure, or used to power the refinery's own burners. During the winter, butane (C4H10), is blended into the gasoline pool at high rates, because its high vapour pressure assists with cold starts. Liquified under pressure slightly above atmospheric, it is best known for powering cigarette lighters, but it is also a main fuel source for many developing countries. Propane can be liquified under modest pressure, and is consumed for just about every application relying on petroleum for energy, from cooking to heating to transportation.
+The cycloalkanes, also known as naphthenes, are saturated hydrocarbons which have one or more carbon rings to which hydrogen atoms are attached according to the formula CnH2n. Cycloalkanes have similar properties to alkanes but have higher boiling points.
+The aromatic hydrocarbons are unsaturated hydrocarbons which have one or more planar six-carbon rings called benzene rings, to which hydrogen atoms are attached with the formula CnH2n-6. They tend to burn with a sooty flame, and many have a sweet aroma. Some are carcinogenic.
+These different molecules are separated by fractional distillation at an oil refinery to produce gasoline, jet fuel, kerosene, and other hydrocarbons. For example, 2,2,4-trimethylpentane (isooctane), widely used in gasoline, has a chemical formula of C8H18 and it reacts with oxygen exothermically:[76]
+The number of various molecules in an oil sample can be determined by laboratory analysis. The molecules are typically extracted in a solvent, then separated in a gas chromatograph, and finally determined with a suitable detector, such as a flame ionization detector or a mass spectrometer.[77] Due to the large number of co-eluted hydrocarbons within oil, many cannot be resolved by traditional gas chromatography and typically appear as a hump in the chromatogram. This Unresolved Complex Mixture (UCM) of hydrocarbons is particularly apparent when analysing weathered oils and extracts from tissues of organisms exposed to oil. Some of the component of oil will mix with water: the water associated fraction of the oil.
+Incomplete combustion of petroleum or gasoline results in production of toxic byproducts. Too little oxygen during combustion results in the formation of carbon monoxide. Due to the high temperatures and high pressures involved, exhaust gases from gasoline combustion in car engines usually include nitrogen oxides which are responsible for creation of photochemical smog.
+At a constant volume, the heat of combustion of a petroleum product can be approximated as follows:
+where
+Q
+v
+{\displaystyle Q_{v}}
+ is measured in calories per gram and
+d
+{\displaystyle d}
+ is the specific gravity at 60 °F (16 °C).
+The thermal conductivity of petroleum based liquids can be modeled as follows:[78]
+where
+K
+{\displaystyle K}
+ is measured in BTU · °F−1hr−1ft−1 ,
+t
+{\displaystyle t}
+ is measured in °F and
+A
+P
+I
+{\displaystyle API}
+ is degrees API gravity.
+The specific heat of petroleum oils can be modeled as follows:[79]
+where
+c
+{\displaystyle c}
+ is measured in BTU/(lb °F),
+t
+{\displaystyle t}
+ is the temperature in Fahrenheit and
+d
+{\displaystyle d}
+ is the specific gravity at 60 °F (16 °C).
+In units of kcal/(kg·°C), the formula is:
+where the temperature
+t
+{\displaystyle t}
+ is in Celsius and
+d
+{\displaystyle d}
+ is the specific gravity at 15 °C.
+The latent heat of vaporization can be modeled under atmospheric conditions as follows:
+where
+L
+{\displaystyle L}
+ is measured in BTU/lb,
+t
+{\displaystyle t}
+ is measured in °F and
+d
+{\displaystyle d}
+ is the specific gravity at 60 °F (16 °C).
+In units of kcal/kg, the formula is:
+where the temperature
+t
+{\displaystyle t}
+ is in Celsius and
+d
+{\displaystyle d}
+ is the specific gravity at 15 °C.[80]
+Petroleum is a fossil fuel derived from ancient fossilized organic materials, such as zooplankton and algae.[83][84] Vast amounts of these remains settled to sea or lake bottoms where they were covered in stagnant water (water with no dissolved oxygen) or sediments such as mud and silt faster than they could decompose aerobically. Approximately 1 m below this sediment or[clarification needed] water oxygen concentration was low, below 0.1 mg/l, and anoxic conditions existed. Temperatures also remained constant.[84]
+As further layers settled to the sea or lake bed, intense heat and pressure built up in the lower regions. This process caused the organic matter to change, first into a waxy material known as kerogen, found in various oil shales around the world, and then with more heat into liquid and gaseous hydrocarbons via a process known as catagenesis. Formation of petroleum occurs from hydrocarbon pyrolysis in a variety of mainly endothermic reactions at high temperature or pressure, or both.[84][85] These phases are described in detail below.
+In the absence of plentiful oxygen, aerobic bacteria were prevented from decaying the organic matter after it was buried under a layer of sediment or water. However, anaerobic bacteria were able to reduce sulfates and nitrates among the matter to H2S and N2 respectively by using the matter as a source for other reactants. Due to such anaerobic bacteria, at first this matter began to break apart mostly via hydrolysis: polysaccharides and proteins were hydrolyzed to simple sugars and amino acids respectively. These were further anaerobically oxidized at an accelerated rate by the enzymes of the bacteria: e.g., amino acids went through oxidative deamination to imino acids, which in turn reacted further to ammonia and α-keto acids. Monosaccharides in turn ultimately decayed to CO2 and methane. The anaerobic decay products of amino acids, monosaccharides, phenols and aldehydes combined to fulvic acids. Fats and waxes were not extensively hydrolyzed under these mild conditions.[84]
+Some phenolic compounds produced from previous reactions worked as bactericides and the actinomycetales order of bacteria  also produced antibiotic compounds (e.g., streptomycin). Thus the action of anaerobic bacteria ceased at about 10 m below the water or sediment. The mixture at this depth contained fulvic acids, unreacted and partially reacted fats and waxes, slightly modified lignin, resins and other hydrocarbons.[84] As more layers of organic matter settled to the sea or lake bed, intense heat and pressure built up in the lower regions.[85] As a consequence, compounds of this mixture began to combine in poorly understood ways to kerogen. Combination happened in a similar fashion as phenol and formaldehyde molecules react to urea-formaldehyde resins, but kerogen formation occurred in a more complex manner due to a bigger variety of reactants. The total process of kerogen formation from the beginning of anaerobic decay is called diagenesis, a word that means a transformation of materials by dissolution and recombination of their constituents.[84]
+Kerogen formation continued to the depth of about 1 km from the Earth's surface where temperatures may reach around 50 °C. Kerogen formation represents a halfway point between organic matter and fossil fuels: kerogen can be exposed to oxygen, oxidize and thus be lost or it could be buried deeper inside the Earth's crust and be subjected to conditions which allow it to slowly transform into fossil fuels like petroleum. The latter happened through catagenesis in which the reactions were mostly radical rearrangements of kerogen. These reactions took thousands to millions of years and no external reactants were involved. Due to radical nature of these reactions, kerogen reacted towards two classes of products: those with low H/C ratio (anthracene or products similar to it) and those with high H/C ratio (methane or products similar to it); i.e., carbon-rich or hydrogen-rich products. Because catagenesis was closed off from external reactants, the resulting composition of the fuel mixture was dependent on the composition of the kerogen via reaction stoichiometry. 3 main types of kerogen exist: type I (algal), II (liptinic) and III (humic), which were formed mainly from algae, plankton and woody plants (this term includes trees, shrubs and lianas) respectively.[84]
+Catagenesis was pyrolytic despite of the fact that it happened at relatively low temperatures (when compared to commercial pyrolysis plants) of 60 to several hundred °C. Pyrolysis was possible because of the long reaction times involved. Heat for catagenesis came from the decomposition of radioactive materials of the crust, especially 40K, 232Th, 235U and 238U. The heat varied with geothermal gradient and was typically 10-30 °C per km of depth from the Earth's surface. Unusual magma intrusions, however, could have created greater localized heating.[84]
+Geologists often refer to the temperature range in which oil forms as an "oil window".[86][84] Below the minimum temperature oil remains trapped in the form of kerogen. Above the maximum temperature the oil is converted to natural gas through the process of thermal cracking. Sometimes, oil formed at extreme depths may migrate and become trapped at a much shallower level. The Athabasca Oil Sands are one example of this.[84]
+An alternative mechanism to the one described above was proposed by Russian scientists in the mid-1850s, the hypothesis of abiogenic petroleum origin (petroleum formed by inorganic means), but this is contradicted by geological and geochemical evidence.[87] Abiogenic sources of oil have been found, but never in commercially profitable amounts. "The controversy isn't over whether abiogenic oil reserves exist," said Larry Nation of the American Association of Petroleum Geologists. "The controversy is over how much they contribute to Earth's overall reserves and how much time and effort geologists should devote to seeking them out."[88]
+Three conditions must be present for oil reservoirs to form:
+The reactions that produce oil and natural gas are often modeled as first order breakdown reactions, where hydrocarbons are broken down to oil and natural gas by a set of parallel reactions, and oil eventually breaks down to natural gas by another set of reactions. The latter set is regularly used in petrochemical plants and oil refineries.
+Wells are drilled into oil reservoirs to extract the crude oil. "Natural lift" production methods that rely on the natural reservoir pressure to force the oil to the surface are usually sufficient for a while after reservoirs are first tapped. In some reservoirs, such as in the Middle East, the natural pressure is sufficient over a long time. The natural pressure in most reservoirs, however, eventually dissipates. Then the oil must be extracted using "artificial lift" means. Over time, these "primary" methods become less effective and "secondary" production methods may be used. A common secondary method is "waterflood" or injection of water into the reservoir to increase pressure and force the oil to the drilled shaft or "wellbore." Eventually "tertiary" or "enhanced" oil recovery methods may be used to increase the oil's flow characteristics by injecting steam, carbon dioxide and other gases or chemicals into the reservoir. In the United States, primary production methods account for less than 40 percent of the oil produced on a daily basis, secondary methods account for about half, and tertiary recovery the remaining 10 percent. Extracting oil (or "bitumen") from oil/tar sand and oil shale deposits requires mining the sand or shale and heating it in a vessel or retort, or using "in-situ" methods of injecting heated liquids into the deposit and then pumping the liquid back out saturated with oil.
+Oil-eating bacteria biodegrade oil that has escaped to the surface. Oil sands are reservoirs of partially biodegraded oil still in the process of escaping and being biodegraded, but they contain so much migrating oil that, although most of it has escaped, vast amounts are still present—more than can be found in conventional oil reservoirs. The lighter fractions of the crude oil are destroyed first, resulting in reservoirs containing an extremely heavy form of crude oil, called crude bitumen in Canada, or extra-heavy crude oil in Venezuela. These two countries have the world's largest deposits of oil sands.[citation needed]
+On the other hand, oil shales are source rocks that have not been exposed to heat or pressure long enough to convert their trapped hydrocarbons into crude oil. Technically speaking, oil shales are not always shales and do not contain oil, but are fined-grain sedimentary rocks containing an insoluble organic solid called kerogen. The kerogen in the rock can be converted into crude oil using heat and pressure to simulate natural processes. The method has been known for centuries and was patented in 1694 under British Crown Patent No. 330 covering, "A way to extract and make great quantities of pitch, tar, and oil out of a sort of stone." Although oil shales are found in many countries, the United States has the world's largest deposits.[89]
+The petroleum industry generally classifies crude oil by the geographic location it is produced in (e.g., West Texas Intermediate, Brent, or Oman), its API gravity (an oil industry measure of density), and its sulfur content. Crude oil may be considered light if it has low density or heavy if it has high density; and it may be referred to as sweet if it contains relatively little sulfur or sour if it contains substantial amounts of sulfur.[citation needed]
+The geographic location is important because it affects transportation costs to the refinery. Light crude oil is more desirable than heavy oil since it produces a higher yield of gasoline, while sweet oil commands a higher price than sour oil because it has fewer environmental problems and requires less refining to meet sulfur standards imposed on fuels in consuming countries. Each crude oil has unique molecular characteristics which are revealed by the use of Crude oil assay analysis in petroleum laboratories.[90]
+Barrels from an area in which the crude oil's molecular characteristics have been determined and the oil has been classified are used as pricing references throughout the world. Some of the common reference crudes are:[citation needed]
+There are declining amounts of these benchmark oils being produced each year, so other oils are more commonly what is actually delivered. While the reference price may be for West Texas Intermediate delivered at Cushing, the actual oil being traded may be a discounted Canadian heavy oil—Western Canadian Select—delivered at Hardisty, Alberta, and for a Brent Blend delivered at Shetland, it may be a discounted Russian Export Blend delivered at the port of Primorsk.[93]
+The petroleum industry is involved in the global processes of exploration, extraction, refining, transporting (often with oil tankers and pipelines), and marketing petroleum products. The largest volume products of the industry are fuel oil and gasoline. Petroleum is also the raw material for many chemical products, including pharmaceuticals, solvents, fertilizers, pesticides, and plastics. The industry is usually divided into three major components: upstream, midstream and downstream. Midstream operations are usually included in the downstream category.[citation needed]
+Petroleum is vital to many industries, and is of importance to the maintenance of industrialized civilization itself, and thus is a critical concern to many nations. Oil accounts for a large percentage of the world's energy consumption, ranging from a low of 32 percent for Europe and Asia, up to a high of 53 percent for the Middle East, South and Central America (44%), Africa (41%), and North America (40%). The world at large consumes 30 billion barrels (4.8 km3) of oil per year, and the top oil consumers largely consist of developed nations. In fact, 24 percent of the oil consumed in 2004 went to the United States alone,[95] though by 2007 this had dropped to 21 percent of world oil consumed.[96]
+In the US, in the states of Arizona, California, Hawaii, Nevada, Oregon and Washington, the Western States Petroleum Association (WSPA) represents companies responsible for producing, distributing, refining, transporting and marketing petroleum. This non-profit trade association was founded in 1907, and is the oldest petroleum trade association in the United States.[97]
+In the 1950s, shipping costs made up 33 percent of the price of oil transported from the Persian Gulf to the United States,[98] but due to the development of supertankers in the 1970s, the cost of shipping dropped to only 5 percent of the price of Persian oil in the US.[98] Due to the increase of the value of the crude oil during the last 30 years, the share of the shipping cost on the final cost of the delivered commodity was less than 3% in 2010. For example, in 2010 the shipping cost from the Persian Gulf to the US was in the range of 20 $/t and the cost of the delivered crude oil around 800 $/t.[citation needed]
+After the collapse of the OPEC-administered pricing system in 1985, and a short-lived experiment with netback pricing, oil-exporting countries adopted a market-linked pricing mechanism. First adopted by PEMEX in 1986, market-linked pricing was widely accepted, and by 1988 became and still is the main method for pricing crude oil in international trade. The current reference, or pricing markers, are Brent, WTI, and Dubai/Oman.[99] Other important market is in Shanghai, because of high consumption in China(rising in XXI century). It is also uncommon, because it shifted trading from USD to RMB.[100]
+The chemical structure of petroleum is heterogeneous, composed of hydrocarbon chains of different lengths. Because of this, petroleum may be taken to oil refineries and the hydrocarbon chemicals separated by distillation and treated by other chemical processes, to be used for a variety of purposes. The total cost per plant is about 9 billion dollars.
+The most common distillation fractions of petroleum are fuels. Fuels include (by increasing boiling temperature range):[66]
+Petroleum classification according to chemical composition.[101]
+Certain types of resultant hydrocarbons may be mixed with other non-hydrocarbons, to create other end products:
+Since the 1940s, agricultural productivity has increased dramatically, due largely to the increased use of energy-intensive mechanization, fertilizers and pesticides.
+Rate of world energy usage per year from 1970.[102]
+Daily oil consumption from 1980 to 2006.
+Oil consumption 1980 to 2007 by region.
+According to the US Energy Information Administration (EIA) estimate for 2011, the world consumes 87.421 million barrels of oil each day.
+This table orders the amount of petroleum consumed in 2011 in thousand barrels (1000 bbl) per day and in thousand cubic metres (1000 m3) per day:[103][104]
+Source: US Energy Information Administration
+Population Data:[105]
+1 peak production of oil already passed in this state
+2 This country is not a major oil producer
+In petroleum industry parlance, production refers to the quantity of crude extracted from reserves, not the literal creation of the product.
+In order of net exports in 2011, 2009 and 2006 in thousand bbl/d and thousand m3/d:
+Source: US Energy Information Administration
+1 peak production already passed in this state
+2 Canadian statistics are complicated by the fact it is both an importer and exporter of crude oil, and refines large amounts of oil for the U.S. market. It is the leading source of U.S. imports of oil and products, averaging 2,500,000 bbl/d (400,000 m3/d) in August 2007.[108]
+Total world production/consumption (as of 2005) is approximately 84 million barrels per day (13,400,000 m3/d).
+In order of net imports in 2011, 2009 and 2006 in thousand bbl/d and thousand m3/d:
+Source: US Energy Information Administration
+1 peak production of oil expected in 2020[109]
+Countries whose oil production is 10% or less of their consumption.
+Source: CIA World Factbook[failed verification]
+Because petroleum is a naturally occurring substance, its presence in the environment need not be the result of human causes such as accidents and routine activities (seismic exploration, drilling, extraction, refining and combustion). Phenomena such as seeps[110] and tar pits are examples of areas that petroleum affects without man's involvement. Regardless of source, petroleum's effects when released into the environment are similar.
+When burned, petroleum releases carbon dioxide, a greenhouse gas. Along with the burning of coal, petroleum combustion is the largest contributor to the increase in atmospheric CO2.[111][112]  Atmospheric CO2 has risen over the last 150 years to current levels of over 415 ppmv,[113] from the 180–300 ppmv of the prior 800 thousand years.[114][115][116] This rise in temperature has reduced the minimum Arctic ice pack to 4,320,000 km2 (1,670,000 sq mi), a loss of almost half since satellite measurements started in 1979.[117][118] Because of this melt, more oil reserves have been revealed. About 13 percent of the world's undiscovered oil resides in the Arctic.[119]
+Ocean acidification is the increase in the acidity of the Earth's oceans caused by the uptake of carbon dioxide (CO2) from the atmosphere. This increase in acidity inhibits all marine life—having a greater impact on smaller organisms as well as shelled organisms (see scallops).[120]
+Oil extraction is simply the removal of oil from the reservoir (oil pool). Oil is often recovered as a water-in-oil emulsion, and specialty chemicals called demulsifiers are used to separate the oil from water. Oil extraction is costly and often environmentally damaging. Offshore exploration and extraction of oil disturb the surrounding marine environment.[121]
+Crude oil and refined fuel spills from tanker ship accidents have damaged natural ecosystems and human livelihoods in Alaska, the Gulf of Mexico, the Galápagos Islands, France and many other places.
+The quantity of oil spilled during accidents has ranged from a few hundred tons to several hundred thousand tons (e.g., Deepwater Horizon oil spill, SS Atlantic Empress, Amoco Cadiz). Smaller spills have already proven to have a great impact on ecosystems, such as the Exxon Valdez oil spill.
+Oil spills at sea are generally much more damaging than those on land, since they can spread for hundreds of nautical miles in a thin oil slick which can cover beaches with a thin coating of oil. This can kill sea birds, mammals, shellfish and other organisms it coats. Oil spills on land are more readily containable if a makeshift earth dam can be rapidly bulldozed around the spill site before most of the oil escapes, and land animals can avoid the oil more easily.
+Control of oil spills is difficult, requires ad hoc methods, and often a large amount of manpower. The dropping of bombs and incendiary devices from aircraft on the SS Torrey Canyon wreck produced poor results;[122] modern techniques would include pumping the oil from the wreck, like in the Prestige oil spill or the Erika oil spill.[123]
+Though crude oil is predominantly composed of various hydrocarbons, certain nitrogen heterocyclic compounds, such as pyridine, picoline, and quinoline are reported as contaminants associated with crude oil, as well as facilities processing oil shale or coal, and have also been found at legacy wood treatment sites. These compounds have a very high water solubility, and thus tend to dissolve and move with water. Certain naturally occurring bacteria, such as Micrococcus, Arthrobacter, and Rhodococcus have been shown to degrade these contaminants.[124]
+A tarball is a blob of crude oil (not to be confused with tar, which is a man-made product derived from pine trees or refined from petroleum) which has been weathered after floating in the ocean. Tarballs are an aquatic pollutant in most environments, although they can occur naturally, for example in the Santa Barbara Channel of California[125][126] or in the Gulf of Mexico off Texas.[127] Their concentration and features have been used to assess the extent of oil spills. Their composition can be used to identify their sources of origin,[128][129] and tarballs themselves may be dispersed over long distances by deep sea currents.[126] They are slowly decomposed by bacteria, including Chromobacterium violaceum, Cladosporium resinae, Bacillus submarinus, Micrococcus varians, Pseudomonas aeruginosa, Candida marina and Saccharomyces estuari.[125]
+James S. Robbins has argued that the advent of petroleum-refined kerosene saved some species of great whales from extinction by providing an inexpensive substitute for whale oil, thus eliminating the economic imperative for open-boat whaling.[130]
+In the United States in 2007 about 70 percent of petroleum was used for transportation (e.g. gasoline, diesel, jet fuel), 24 percent by industry (e.g. production of plastics), 5 percent for residential and commercial uses, and 2 percent for electricity production.[131] Outside of the US, a higher proportion of petroleum tends to be used for electricity.[132]
+Petroleum-based vehicle fuels can be replaced by either alternative fuels, or other methods of propulsion such as electric or nuclear.
+Alternative fuel vehicles refers to both:
+Biological feedstocks do exist for industrial uses such as Bioplastic production.[133]
+In oil producing countries with little refinery capacity, oil is sometimes burned to produce electricity. Renewable energy technologies such as solar power, wind power, micro hydro, biomass and biofuels are used, but the primary alternatives remain large-scale hydroelectricity, nuclear and coal-fired generation.
+Consumption in the twentieth and twenty-first centuries has been abundantly pushed by automobile sector growth. The 1985–2003 oil glut even fueled the sales of low fuel economy vehicles in OECD countries. The 2008 economic crisis seems to have had some impact on the sales of such vehicles; still, in 2008 oil consumption showed a small increase.
+In 2016 Goldman Sachs predicted lower demand for oil due to emerging economies concerns, especially China.[134] The BRICS (Brasil, Russia, India, China, South Africa) countries might also kick in, as China briefly was the first automobile market in December 2009.[135] The immediate outlook still hints upwards. In the long term, uncertainties linger; the OPEC believes that the OECD countries will push low consumption policies at some point in the future; when that happens, it will definitely curb oil sales, and both OPEC and the Energy Information Administration (EIA) kept lowering their 2020 consumption estimates during the past five years.[136] A detailed review of International Energy Agency oil projections have revealed that revisions of world oil production, price and investments have been motivated by a combination of demand and supply factors.[137] All together, Non-OPEC conventional projections have been fairly stable the last 15 years, while downward revisions were mainly allocated to OPEC. Recent upward revisions are primarily a result of US tight oil.
+Production will also face an increasingly complex situation; while OPEC countries still have large reserves at low production prices, newly found reservoirs often lead to higher prices; offshore giants such as Tupi, Guara and Tiber demand high investments and ever-increasing technological abilities. Subsalt reservoirs such as Tupi were unknown in the twentieth century, mainly because the industry was unable to probe them. Enhanced Oil Recovery (EOR) techniques (example: DaQing, China[138]) will continue to play a major role in increasing the world's recoverable oil.
+The expected availability of petroleum resources has always been around 35 years or even less since the start of the modern exploration. The oil constant, an insider pun in the German industry, refers to that effect.[139]
+A growing number of divestment campaigns from major funds pushed by newer generations who question the sustainability of petroleum may hinder the financing of future oil prospection and production.
+Peak oil is a term applied to the projection that future petroleum production (whether for individual oil wells, entire oil fields, whole countries, or worldwide production) will eventually peak and then decline at a similar rate to the rate of increase before the peak as these reserves are exhausted. The peak of oil discoveries was in 1965, and oil production per year has surpassed oil discoveries every year since 1980.[140] However, this does not mean that potential oil production has surpassed oil demand.
+Hubbert applied his theory to accurately predict the peak of U.S. conventional oil production at a date between 1966 and 1970. This prediction was based on data available at the time of his publication in 1956. In the same paper, Hubbert predicts world peak oil in "half a century" after his publication, which would be 2006.[141]
+It is difficult to predict the oil peak in any given region, due to the lack of knowledge and/or transparency in accounting of global oil reserves.[142] Based on available production data, proponents have previously predicted the peak for the world to be in years 1989, 1995, or 1995–2000. Some of these predictions date from before the recession of the early 1980s, and the consequent reduction in global consumption, the effect of which was to delay the date of any peak by several years. Just as the 1971 U.S. peak in oil production was only clearly recognized after the fact, a peak in world production will be difficult to discern until production clearly drops off.[143] The peak is also a moving target as it is now measured as "liquids", which includes synthetic fuels, instead of just conventional oil.[144]
+The International Energy Agency (IEA) said in 2010 that production of conventional crude oil had peaked in 2006 at 70 MBBL/d, then flattened at 68 or 69 thereafter.[145][146] Since virtually all economic sectors rely heavily on petroleum, peak oil, if it were to occur, could lead to a "partial or complete failure of markets".[147] In the mid-2000s, widespread fears of an imminent peak led to the "peak oil movement," in which over one hundred thousand Americans prepared, individually and collectively, for the "post-carbon" future.[148]
+While there has been much focus historically on peak oil supply, focus is increasingly shifting to peak demand as more countries seek to transition to renewable energy. The GeGaLo index of geopolitical gains and losses assesses how the geopolitical position of 156 countries may change if the world fully transitions to renewable energy resources. Former oil exporters are expected to lose power, while the positions of former oil importers and countries rich in renewable energy resources is expected to strengthen.[149]
+Unconventional oil is petroleum produced or extracted using techniques other than the conventional methods.[150][151] The calculus for peak oil has changed with the introduction of unconventional production methods. In particular, the combination of horizontal drilling and hydraulic fracturing has resulted in a significant increase in production from previously uneconomic plays.[152] Analysts expected that $150 billion would be spent on further developing North American tight oil fields in 2015. The large increase in tight oil production is one of the reasons behind the price drop in late 2014.[153] Certain rock strata contain hydrocarbons but have low permeability and are not thick from a vertical perspective. Conventional vertical wells would be unable to economically retrieve these hydrocarbons. Horizontal drilling, extending horizontally through the strata, permits the well to access a much greater volume of the strata. Hydraulic fracturing creates greater permeability and increases hydrocarbon flow to the wellbore.

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+Aglossata
+Glossata
+Heterobathmiina
+Zeugloptera
+Lepidoptera (/ˌlɛpɪˈdɒptərə/ LEP-i-DOP-tər-ə, from Ancient Greek lepís “scale” + pterón “wing”) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families[1] and 46 superfamilies,[2] 10 per cent of the total described species of living organisms.[2][3] It is one of the most widespread and widely recognizable insect orders in the world.[4] The Lepidoptera show many variations of the basic body structure that have evolved to gain advantages in lifestyle and distribution. Recent estimates suggest the order may have more species than earlier thought,[5] and is among the four most speciose orders, along with the Hymenoptera, Diptera, and Coleoptera.[4]
+Lepidopteran species are characterized by more than three derived features. The most apparent is the presence of scales that cover the bodies, wings, and a proboscis. The scales are modified, flattened "hairs", and give butterflies and moths their wide variety of colors and patterns. Almost all species have some form of membranous wings, except for a few that have reduced wings or are wingless. Mating and the laying of eggs are carried out by adults, normally near or on host plants for the larvae.  Like most other insects, butterflies and moths are holometabolous, meaning they undergo complete metamorphosis.  The larvae are commonly called caterpillars, and are completely different from their adult moth or butterfly forms, having a cylindrical body with a well-developed head, mandible mouth parts, three pairs of thoracic legs and from none up to five pairs of prolegs. As they grow, these larvae change in appearance, going through a series of stages called instars. Once fully matured, the larva develops into a pupa. A few butterflies and many moth species spin a silk case or cocoon prior to pupating, while others do not, instead going underground.[4] A butterfly pupa, called a chrysalis, has a hard skin, usually with no cocoon. Once the pupa has completed its metamorphosis, a sexually mature adult emerges.
+The Lepidoptera have, over millions of years, evolved a wide range of wing patterns and coloration ranging from drab moths akin to the related order Trichoptera, to the brightly colored and complex-patterned butterflies.[1] Accordingly, this is the most recognized and popular of insect orders with many people involved in the observation, study, collection, rearing of, and commerce in these insects. A person who collects or studies this order is referred to as a lepidopterist.
+Butterflies and moths play an important role in the natural ecosystem as pollinators and as food in the food chain; conversely, their larvae are considered very problematic to vegetation in agriculture, as their main source of food is often live plant matter. In many species, the female may produce from 200 to 600 eggs, while in others, the number may approach 30,000 eggs in one day. The caterpillars hatching from these eggs can cause damage to large quantities of crops. Many moth and butterfly species are of economic interest by virtue of their role as pollinators, the silk they produce, or as pest species.
+The term was coined by Linnaeus in 1735 and is derived from Greek λεπίς, gen. λεπίδος ("scale") and πτερόν ("wing").[6] Sometimes, the term Rhopalocera is used for the clade of all butterfly species, derived from the Ancient Greek ῥόπαλον (rhopalon)[7]:4150 and κέρας (keras)[7]:3993 meaning "club" and "horn", respectively, coming from the shape of the antennae of butterflies.
+The origins of the common names "butterfly" and "moth" are varied and often obscure. The English word butterfly is from Old English buttorfleoge, with many variations in spelling. Other than that, the origin is unknown, although it could be derived from the pale yellow color of many species' wings suggesting the color of butter.[8][9] The species of Heterocera are commonly called moths. The origins of the English word moth are more clear, deriving from the Old English moððe" (cf. Northumbrian dialect mohðe) from Common Germanic (compare Old Norse motti, Dutch mot and German Motte all meaning "moth"). Perhaps its origins are related to Old English maða meaning "maggot" or from the root of "midge", which until the 16th century was used mostly to indicate the larva, usually in reference to devouring clothes.[10]
+The etymological origins of the word "caterpillar", the larval form of butterflies and moths, are from the early 16th century, from Middle English catirpel, catirpeller, probably an alteration of Old North French catepelose: cate, cat (from Latin cattus) + pelose, hairy (from Latin pilōsus).[11]
+The Lepidoptera are among the most successful groups of insects. They are found on all continents, except Antarctica, and inhabit all terrestrial habitats ranging from desert to rainforest, from lowland grasslands to mountain plateaus, but almost always associated with higher plants, especially angiosperms (flowering plants).[12] Among the most northern dwelling species of butterflies and moths is the Arctic Apollo (Parnassius arcticus), which is found in the Arctic Circle in northeastern Yakutia, at an altitude of 1500 m above sea level.[13] In the Himalayas, various Apollo species such as Parnassius epaphus  have been recorded to occur up to an altitude of 6,000 m above sea level.[14]:221
+Some lepidopteran species exhibit symbiotic, phoretic, or parasitic lifestyles, inhabiting the bodies of organisms rather than the environment. Coprophagous pyralid moth species, called sloth moths, such as Bradipodicola hahneli and Cryptoses choloepi, are unusual in that they are exclusively found inhabiting the fur of sloths, mammals found in Central and South America.[15][16]
+Two species of Tinea moths have been recorded as feeding on horny tissue and have been bred from the horns of cattle. The larva of Zenodochium coccivorella is an internal parasite of the coccid Kermes species. Many species have been recorded as breeding in natural materials or refuse such as owl pellets, bat caves, honeycombs or diseased fruit.[16]
+As of 2007, there was roughly 174,250 lepidopteran species described, with butterflies and skippers estimated to comprise around 17,950, and moths making up the rest.[2][17] The vast majority of Lepidoptera are to be found in the tropics, but substantial diversity exists on most continents. North America has over 700 species of butterflies and over 11,000 species of moths,[18][19] while about 400 species of butterflies and 14,000 species of moths are reported from Australia.[20] The diversity of Lepidoptera in each faunal region has been estimated by John Heppner in 1991 based partly on actual counts from the literature, partly on the card indices in the Natural History Museum (London) and the National Museum of Natural History (Washington), and partly on estimates:[5]
+Lepidoptera are morphologically distinguished from other orders principally by the presence of scales on the external parts of the body and appendages, especially the wings. Butterflies and moths vary in size from microlepidoptera only a few millimeters long, to conspicuous animals with a wingspan greater than 25 centimetres, such as the Queen Alexandra's birdwing and Atlas moth.[21]:246
+Lepidopterans undergo a four-stage life cycle: egg; larva or caterpillar; pupa or chrysalis; and imago (plural: imagines) / adult and show many variations of the basic body structure, which give these animals advantages for diverse lifestyles and environments.
+The head is where many sensing organs and the mouth parts are found. Like the adult, the larva also has a toughened, or sclerotized head capsule.[22] Here, two compound eyes, and chaetosema, raised spots or clusters of sensory bristles unique to Lepidoptera, occur, though many taxa have lost one or both of these spots. The antennae have a wide variation in form among species and even between different sexes. The antennae of butterflies are usually filiform and shaped like clubs, those of the skippers are hooked, while those of moths have flagellar segments variously enlarged or branched. Some moths have enlarged antennae or ones that are tapered and hooked at the ends.[23]:559–560
+The maxillary galeae are modified and form an elongated proboscis. The proboscis consists of one to five segments, usually kept coiled up under the head by small muscles when it is not being used to suck up nectar from flowers or other liquids. Some basal moths still have mandibles, or separate moving jaws, like their ancestors, and these form the family Micropterigidae.[22][23]:560[24]
+The larvae, called caterpillars, have a toughened head capsule. Caterpillars lack the proboscis and have separate chewing mouthparts.[22] These mouthparts, called mandibles, are used to chew up the plant matter that the larvae eat. The lower jaw, or labium, is weak, but may carry a spinneret, an organ used to create silk. The head is made of large lateral lobes, each having an ellipse of up to six simple eyes.[23]:562–563
+The thorax is made of three fused segments, the prothorax, mesothorax, and metathorax, each with a pair of legs. The first segment contains the first pair of legs. In some males of the butterfly family Nymphalidae, the forelegs are greatly reduced and are not used for walking or perching.[23]:586 The three pairs of legs are covered with scales. Lepidoptera also have olfactory organs on their feet, which aid the butterfly in "tasting" or "smelling" out its food.[25] In the larval form there are 3 pairs of true legs, with up to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[12]
+The two pairs of wings are found on the middle and third segments, or mesothorax and metathorax, respectively. In the more recent genera, the wings of the second segment are much more pronounced, although some more primitive forms have similarly sized wings of both segments. The wings are covered in scales arranged like shingles, which form an extraordinary variety of colors and patterns. The mesothorax has more powerful muscles to propel the moth or butterfly through the air, with the wing of this segment (forewing) having a stronger vein structure.[23]:560 The largest superfamily, the Noctuidae, has their wings modified to act as tympanal or hearing organs.[26]
+The caterpillar has an elongated, soft body that may have hair-like or other projections, three pairs of true legs, with none to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[12] The thorax usually has a pair of legs on each segment. The thorax is also lined with many spiracles on both the mesothorax and metathorax, except for a few aquatic species, which instead have a form of gills.[23]:563
+The abdomen, which is less sclerotized than the thorax, consists of 10 segments with membranes in between, allowing for articulated movement. The sternum, on the first segment, is small in some families and is completely absent in others. The last two or three segments form the external parts of the species' sex organs. The genitalia of Lepidoptera are highly varied and are often the only means of differentiating between species. Male genitals include a valva, which is usually large, as it is used to grasp the female during mating. Female genitalia include three distinct sections.
+The females of basal moths have only one sex organ, which is used for copulation and as an ovipositor, or egg-laying organ. About 98% of moth species have a separate organ for mating, and an external duct that carries the sperm from the male.[23]:561
+The abdomen of the caterpillar has four pairs of prolegs, normally located on the third to sixth segments of the abdomen, and a separate pair of prolegs by the anus, which have a pair of tiny hooks called crotchets. These aid in gripping and walking, especially in species that lack many prolegs (e. g. larvae of Geometridae). In some basal moths, these prolegs may be on every segment of the body, while prolegs may be completely absent in other groups, which are more adapted to boring and living in sand (e. g., Prodoxidae and Nepticulidae, respectively).[23]:563
+The wings, head, and parts of the thorax and abdomen of Lepidoptera are covered with minute scales, a feature from which the order derives its name. Most scales are lamellar, or blade-like, and attached with a pedicel, while other forms may be hair-like or specialized as secondary sexual characteristics.[27]
+The lumen or surface of the lamella has a complex structure. It gives color either by colored pigments it contains, or through structural coloration with mechanisms that include photonic crystals and diffraction gratings.[28]
+Scales function in insulation, thermoregulation, producing pheromones (in males only),[29] and aiding gliding flight, but the most important is the large diversity of vivid or indistinct patterns they provide, which help the organism protect itself by camouflage or mimicry, and which act as signals to other animals including rivals and potential mates.[27]
+In the reproductive system of butterflies and moths, the male genitalia are complex and unclear. In females the three types of genitalia are based on the relating taxa: 'monotrysian', 'exoporian', and 'ditrysian'. In the monotrysian type is an opening on the fused segments of the sterna 9 and 10, which act as insemination and oviposition. In the exoporian type (in Hepaloidae and Mnesarchaeoidea) are two separate places for insemination and oviposition, both occurring on the same sterna as the monotrysian type, i.e. 9 and 10.[21] The ditrysian groups have an internal duct that carries sperm, with separate openings for copulation and egg-laying.[4] In most species, the genitalia are flanked by two soft lobes, although they may be specialized and sclerotized in some species for ovipositing in area such as crevices and inside plant tissue.[21] Hormones and the glands that produce them run the development of butterflies and moths as they go through their life cycles, called the endocrine system. The first insect hormone prothoracicotropic hormone (PTTH) operates the species life cycle and diapause.[30] This hormone is produced by corpora allata and corpora cardiaca, where it is also stored. Some glands are specialized to perform certain task such as producing silk or producing saliva in the palpi.[31]:65, 75 While the corpora cardiaca produce PTTH, the corpora allata also produces juvenile hormones, and the prothorocic glands produce moulting hormones.
+In the digestive system, the anterior region of the foregut has been modified to form a pharyngeal  sucking pump as they need it for the food they eat, which are for the most part liquids. An esophagus follows and leads to the posterior of the pharynx and in some species forms a form of crop. The midgut is short and straight, with the hindgut being longer and coiled.[21] Ancestors of lepidopteran species, stemming from Hymenoptera, had midgut ceca, although this is lost in current butterflies and moths. Instead, all the digestive enzymes, other than initial digestion, are immobilized at the surface of the midgut cells. In larvae, long-necked and stalked goblet cells are found in the anterior and posterior midgut regions, respectively. In insects, the goblet cells excrete positive potassium ions, which are absorbed from leaves ingested by the larvae. Most butterflies and moths display the usual digestive cycle, but species with different diets require adaptations to meet these new demands.[23]:279
+In the circulatory system, hemolymph, or insect blood, is used to circulate heat in a form of thermoregulation, where muscles contraction produces heat, which is transferred to the rest of the body when conditions are unfavorable.[32]
+In lepidopteran species, hemolymph is circulated through the veins in the wings by some form of pulsating organ, either by the heart or by the intake of air into the trachea.[31]:69
+Air is taken in through spiracles along the sides of the abdomen and thorax supplying the trachea with oxygen as it goes through the lepidopteran's respiratory system. Three different tracheaes supply and diffuse oxygen throughout the species' bodies. The dorsal tracheae supply oxygen to the dorsal musculature and vessels, while the ventral tracheae supply the ventral musculature and nerve cord, and the visceral tracheae supply the guts, fat bodies, and gonads.[31]:71, 72
+Polymorphism is the appearance of forms or "morphs", which differ in color and number of attributes within a single species.[12]:163[33] In Lepidoptera, polymorphism can be seen not only between individuals in a population, but also between the sexes as sexual dimorphism, between geographically separated populations in geographical polymorphism, and between generations flying at different seasons of the year (seasonal polymorphism or polyphenism). In some species, the polymorphism is limited to one sex, typically the female. This often includes the phenomenon of mimicry when mimetic morphs fly alongside nonmimetic morphs in a population of a particular species. Polymorphism occurs both at specific level with heritable variation in the overall morphological adaptations of individuals, as well as in certain specific morphological or physiological traits within a species.[12]
+Environmental polymorphism, in which traits are not inherited, is often termed as polyphenism, which in Lepidoptera is commonly seen in the form of seasonal morphs, especially in the butterfly families of Nymphalidae and Pieridae. An Old World pierid butterfly, the common grass yellow (Eurema hecabe) has a darker summer adult morph, triggered by a long day exceeding 13 hours in duration, while the shorter diurnal period of 12 hours or less induces a paler morph in the postmonsoon period.[34] Polyphenism also occurs in caterpillars, an example being the peppered moth, Biston betularia.[35]
+Geographical isolation causes a divergence of a species into different morphs. A good example is the Indian white admiral Limenitis procris, which has five forms, each geographically separated from the other by large mountain ranges.[36]:26 An even more dramatic showcase of geographical polymorphism is the Apollo butterfly (Parnassius apollo). Because the Apollos live in small local populations, thus having no contact with each other, coupled with their strong stenotopic nature and weak migration ability, interbreeding between populations of one species practically does not occur; by this, they form over 600 different morphs, with the size of spots on the wings of which varies greatly.[37]
+Sexual dimorphism is the occurrence of differences between males and females in a species. In Lepidoptera, it is widespread and almost completely set by genetic determination.[34] Sexual dimorphism is present in all families of the Papilionoidea and more prominent in the Lycaenidae, Pieridae, and certain taxa of the Nymphalidae. Apart from color variation, which may differ from slight to completely different color-pattern combinations, secondary sexual characteristics may also be present.[36]:25 Different genotypes maintained by natural selection may also be expressed at the same time.[34] Polymorphic and/or mimetic females occur in the case of some taxa in the Papilionidae primarily to obtain a level of protection not available to the male of their species. The most distinct case of sexual dimorphism is that of adult females of many Psychidae species which have only vestigial wings, legs, and mouthparts as compared to the adult males that are strong fliers with well-developed wings and feathery antennae.[38]
+Species of Lepidoptera undergo holometabolism or "complete metamorphosis". Their life cycle normally consists of an egg, a larva, a pupa, and an imago or adult.[12] The larvae are commonly called caterpillars, and the pupae of moths encapsulated in silk are called cocoons, while the uncovered pupae of butterflies are called chrysalides.
+Unless the species reproduces year-round, a butterfly or moth may enter diapause, a state of dormancy that allows the insect to survive unfavorable environmental conditions.
+Males usually start eclosion (emergence) earlier than females and peak in numbers before females. Both of the sexes are sexually mature by the time of eclosion.[23]:564 Butterflies and moths normally do not associate with each other, except for migrating species, staying relatively asocial. Mating begins with an adult (female or male) attracting a mate, normally using visual stimuli, especially in diurnal species like most butterflies. However, the females of most nocturnal species, including almost all moth species, use pheromones to attract males, sometimes from long distances.[12] Some species engage in a form of acoustic courtship, or attract mates using sound or vibration such as the polka-dot wasp moth, Syntomeida epilais.[39]
+Adaptations include undergoing one seasonal generation, two or even more, called voltinism (Univoltism, bivoltism, and multivism, respectively). Most lepidopterans in temperate climates are univoltine, while in tropical climates most have two seasonal broods. Some others may take advantage of any opportunity they can get, and mate continuously throughout the year. These seasonal adaptations are controlled by hormones, and these delays in reproduction are called diapause.[23]:567 Many lepidopteran species, after mating and laying their eggs, die shortly afterwards, having only lived for a few days after eclosion. Others may still be active for several weeks and then overwinter and become sexually active again when the weather becomes more favorable, or diapause. The sperm of the male that mated most recently with the female is most likely to have fertilized the eggs, but the sperm from a prior mating may still prevail.[23]:564
+Lepidoptera usually reproduce sexually and are oviparous (egg-laying), though some species exhibit live birth in a process called ovoviviparity. A variety of differences in egg-laying and the number of eggs laid occur. Some species simply drop their eggs in flight (these species normally have polyphagous larvae, meaning they eat a variety of plants e. g., hepialids and some nymphalids)[40] while most lay their eggs near or on the host plant on which the larvae feed. The number of eggs laid may vary from only a few to several thousand.[12] The females of both butterflies and moths select the host plant instinctively, and primarily, by chemical cues.[23]:564
+The eggs are derived from materials ingested as a larva and in some species, from the spermatophores received from males during mating.[41] An egg can only be 1/1000 the mass of the female, yet she may lay up to her own mass in eggs. Females lay smaller eggs as they age. Larger females lay larger eggs.[42] The egg is covered by a hard-ridged protective outer layer of shell, called the chorion. It is lined with a thin coating of wax, which prevents the egg from drying out. Each egg contains a number of micropyles, or tiny funnel-shaped openings at one end, the purpose of which is to allow sperm to enter and fertilize the egg. Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate.
+The egg stage lasts a few weeks in most butterflies, but eggs laid prior to winter, especially in temperate regions, go through diapause, and hatching may be delayed until spring. Other butterflies may lay their eggs in the spring and have them hatch in the summer. These butterflies are usually temperate species (e. g. Nymphalis antiopa).
+The larvae or caterpillars are the first stage in the life cycle after hatching. Caterpillars, are "characteristic polypod larvae with cylindrical bodies, short thoracic legs, and abdominal prolegs (pseudopods)".[43] They have a toughened (sclerotised) head capsule with an adfrontal suture formed by medial fusion of the sclerites, mandibles (mouthparts) for chewing, and a soft tubular, segmented body, that may have hair-like or other projections, three pairs of true legs, and additional prolegs (up to five pairs).[44] The body consists of thirteen segments, of which three are thoracic and ten are abdominal.[45] Most larvae are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores.[44]
+Different herbivorous species have adapted to feed on every part of the plant and are normally considered pests to their host plants; some species have been found to lay their eggs on the fruit and other species lay their eggs on clothing or fur (e. g., Tineola bisselliella, the common clothes moth). Some species are carnivorous and others are even parasitic. Some lycaenid species such as Phengaris rebeli are social parasites of Myrmica ants nests.[46] A species of Geometridae from Hawaii has carnivorous larvae that catch and eat flies.[47] Some pyralid caterpillars are aquatic.[48]
+The larvae develop rapidly with several generations in a year; however, some species may take up to 3 years to develop, and exceptional examples like Gynaephora groenlandica take as long as seven years.[12] The larval stage is where the feeding and growing stages occur, and the larvae periodically undergo hormone-induced ecdysis, developing further with each instar, until they undergo the final larval-pupal molt.
+The larvae of both butterflies and moths exhibit mimicry to deter potential predators. Some caterpillars have the ability to inflate parts of their heads to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called osmeteria (family Papilionidae), which are exposed to produce smelly chemicals used in defense. Host plants often have toxic substances in them and caterpillars are able to sequester these substances and retain them into the adult stage. This helps make them unpalatable to birds and other predators. Such unpalatability is advertised using bright red, orange, black, or white warning colors. The toxic chemicals in plants are often evolved specifically to prevent them from being eaten by insects. Insects, in turn, develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants.[49]
+No form of wing is externally visible on the larva, but when larvae are dissected, developing wings can be seen as disks, which can be found on the second and third thoracic segments, in place of the spiracles that are apparent on abdominal segments. Wing disks develop in association with a trachea that runs along the base of the wing, and are surrounded by a thin peripodial membrane, which is linked to the outer epidermis of the larva by a tiny duct. Wing disks are very small until the last larval instar, when they increase dramatically in size, are invaded by branching tracheae from the wing base that precede the formation of the wing veins, and begin to develop patterns associated with several landmarks of the wing.[50]
+Near pupation, the wings are forced outside the epidermis under pressure from the hemolymph, and although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval cuticle, they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled without damage to the wings.[50]
+After about five to seven instars,[51]:26–28 or molts, certain hormones, like PTTH, stimulate the production of ecdysone, which initiates insect molting. Then, the larva puparium, a sclerotized or hardened cuticle of the last larval instar, develops into the pupa. Depending on the species, the pupa may be covered in a silk cocoon, attached to different types of substrates, buried in the ground, or may not be covered at all. Features of the imago are externally recognizable in the pupa. All the appendages on the adult head and thorax are found cased inside the cuticle (antennae, mouthparts, etc.), with the wings wrapped around, adjacent to the antennae.[23]:564 The pupae of some species have functional mandibles, while the pupal mandibles are not functional in others.[22]
+While encased, some of the lower segments are not fused, and are able to move using small muscles found in between the membrane. Moving may help the pupa, for example, escape the sun, which would otherwise kill it. The pupa of the Mexican jumping bean moth (Cydia deshaisiana) does this. The larvae cut a trapdoor in the bean (species of Sebastiania) and use the bean as a shelter. With a sudden rise in temperature, the pupa inside twitches and jerks, pulling on the threads inside. Wiggling may also help to deter parasitoid wasps from laying eggs on the pupa. Other species of moths are able to make clicks to deter predators.[23]:564, 566
+The length of time before the pupa ecloses (emerges) varies greatly. The monarch butterfly may stay in its chrysalis for two weeks, while other species may need to stay for more than 10 months in diapause. The adult emerges from the pupa either by using abdominal hooks or from projections located on the head. The mandibles found in the most primitive moth families are used to escape from their cocoon (e. g., Micropterigoidea).[12][23]:564
+Most lepidopteran species do not live long after eclosion, only needing a few days to find a mate and then lay their eggs. Others may remain active for a longer period (from one to several weeks), or go through diapause and overwintering as monarch butterflies do, or waiting out environmental stress. Some adult species of microlepidoptera go through a stage where no reproductive-related activity occurs, lasting through summer and winter, followed by mating and oviposition in the early spring.[23]:564
+While most butterflies and moths are terrestrial, many species of Pyralidae are truly aquatic with all stages except the adult occurring in water. Many species from other families such as Erebidae, Nepticulidae, Cosmopterygidae, Tortricidae, Olethreutidae, Noctuidae, Cossidae, and Sphingidae are aquatic or semiaquatic.[52]:22
+Flight is an important aspect of the lives of butterflies and moths, and is used for evading predators, searching for food, and finding mates in a timely manner, as most lepidopteran species do not live long after eclosion. It is the main form of locomotion in most species. In Lepidoptera, the forewings and hindwings are mechanically coupled and flap in synchrony. Flight is anteromotoric, or being driven primarily by action of the forewings. Although lepidopteran species reportedly can still fly when their hindwings are cut off, it reduces their linear flight and turning capabilities.[53]
+Lepidopteran species have to be warm, about 77 to 79 °F (25 to 26 °C), to fly. They depend on their body temperature being sufficiently high and since they cannot regulate it themselves, this is dependent on their environment. Butterflies living in cooler climates may use their wings to warm their bodies. They will bask in the sun, spreading out their wings so that they get maximum exposure to the sunlight. In hotter climates butterflies can easily overheat, so they are usually active only during the cooler parts of the day, early morning, late afternoon or early evening. During the heat of the day, they rest in the shade. Some larger thick-bodied moths (e.g. Sphingidae) can generate their own heat to a limited degree by vibrating their wings. The heat generated by the flight muscles warms the thorax while the temperature of the abdomen is unimportant for flight. To avoid overheating, some moths rely on hairy scales, internal air sacs, and other structures to separate the thorax and abdomen and keep the abdomen cooler.[54]
+Some species of butterflies can reach fast speeds, such as the southern dart, which can go as fast as 48.4 km/h. Sphingids are some of the fastest flying insects, some are capable of flying at over 50 km/h (30 mi/h), having a wingspan of 35–150 mm.[1][55] In some species, sometimes a gliding component to their flight exists. Flight occurs either as hovering, or as forward or backward motion.[56] In butterfly and moth species, such as hawk moths, hovering is important as they need to maintain a certain stability over flowers when feeding on the nectar.[1]
+Navigation is important to Lepidoptera species, especially for those that migrate. Butterflies, which have more species that migrate, have been shown to navigate using time-compensated sun compasses. They can see polarized light, so can orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is suggested to be particularly important.[57] Most migratory butterflies are those that live in semiarid areas where breeding seasons are short.[58] The life histories of their host plants also influence the strategies of the butterflies.[59] Other theories include the use of landscapes. Lepidoptera may use coastal lines, mountains, and even roads to orient themselves. Above sea, the flight direction is much more accurate if the coast is still visible.[60]
+Many studies have also shown that moths navigate. One study showed that many moths may use the Earth's magnetic field to navigate, as a study of the moth heart and dart suggests.[61] Another study, of the migratory behavior of the silver Y, showed, even at high altitudes, the species can correct its course with changing winds, and prefers flying with favourable winds, suggesting a great sense of direction.[62][63] Aphrissa statira in Panama loses its navigational capacity when exposed to a magnetic field, suggesting it uses the Earth's magnetic field.[64]
+Moths exhibit a tendency to circle artificial lights repeatedly. This suggests they use a technique of celestial navigation called transverse orientation. By maintaining a constant angular relationship to a bright celestial light, such as the Moon, they can fly in a straight line. Celestial objects are so far away, even after traveling great distances, the change in angle between the moth and the light source is negligible; further, the moon will always be in the upper part of the visual field or on the horizon. When a moth encounters a much closer artificial light and uses it for navigation, the angle changes noticeably after only a short distance, in addition to being often below the horizon. The moth instinctively attempts to correct by turning toward the light, causing airborne moths to come plummeting downwards, and at close range, which results in a spiral flight path that gets closer and closer to the light source.[65] Other explanations have been suggested, such as the idea that moths may be impaired with a visual distortion called a Mach band by Henry Hsiao in 1972. He stated that they fly towards the darkest part of the sky in pursuit of safety, thus are inclined to circle ambient objects in the Mach band region.[66]
+Lepidopteran migration is typically seasonal, as the insects moving to escape dry seasons or other disadvantageous conditions. Most lepidopterans that migrate are butterflies, and the distance travelled varies. Some butterflies that migrate include the mourning cloak, painted lady, American lady, red admiral, and the common buckeye.[51]:29–30 A notable species of moth that migrates long distances is the bogong moth.[67] The most well-known migrations are those of the eastern population of the monarch butterfly from Mexico to northern United States and southern Canada, a distance of about 4,000–4,800 km (2,500–3,000 mi). Other well-known migratory species include the painted lady and several of the danaine butterflies. Spectacular and large-scale migrations associated with the monsoons are seen in peninsular India.[68] Migrations have been studied in more recent times using wing tags and stable hydrogen isotopes.[69][70]
+Moths also undertake migrations, an example being the uraniids. Urania fulgens undergoes population explosions and massive migrations that may be not surpassed by any other insect in the Neotropics. In Costa Rica and Panama, the first population movements may begin in July and early August and depending on the year, may be very massive, continuing unabated for as long as five months.[71]
+Pheromones are commonly involved in mating rituals among species, especially moths, but they are also an important aspect of other forms of communication. Usually, the pheromones are produced by either the male or the female and detected by members of the opposite sex with their antennae.[72] In many species, a gland between the eighth and ninth segments under the abdomen in the female produces the pheromones.[12] Communication can also occur through stridulation, or producing sounds by rubbing various parts of the body together.[63]
+Moths are known to engage in acoustic forms of communication, most often as courtship, attracting mates using sound or vibration. Like most other insects, moths pick up these sounds using tympanic membranes in their abdomens.[73] An example is that of the polka-dot wasp moth (Syntomeida epilais), which produces sounds with a frequency above that normally detectable by humans (about 20 kHz). These sounds also function as tactile communication, or communication through touch, as they stridulate, or vibrate a substrate like leaves and stems.[39]
+Most moths lack bright colors, as many species use coloration as camouflage, but butterflies engage in visual communication. Female cabbage butterflies, for example, use ultraviolet light to communicate, with scales colored in this range on the dorsal wing surface. When they fly, each down stroke of the wing creates a brief flash of ultraviolet light which the males apparently recognize as the flight signature of a potential mate. These flashes from the wings may attract several males that engage in aerial courtship displays.[73]
+Moths and butterflies are important in the natural ecosystem. They are integral participants in the food chain; having co-evolved with flowering plants and predators, lepidopteran species have formed a network of trophic relationships between autotrophs and heterotrophs, which are included in the stages of Lepidoptera larvae, pupae, and adults. Larvae and pupae are links in the diets of birds and parasitic entomophagous insects. The adults are included in food webs in a much broader range of consumers (including birds, small mammals, reptiles, etc.).[23]:567
+Lepidopteran species are soft bodied, fragile, and almost defenseless, while the immature stages move slowly or are immobile, hence all stages are exposed to predation. Adult butterflies and moths are preyed upon by birds, bats, lizards, amphibians, dragonflies, and spiders. Caterpillars and pupae fall prey not only to birds, but also to invertebrate predators and small mammals, as well as fungi and bacteria. Parasitoid and parasitic wasps and flies may lay eggs in the caterpillar, which eventually kill it as they hatch inside its body and eat its tissues. Insect-eating birds are probably the largest predators. Lepidoptera, especially the immature stages, are an ecologically important food to many insectivorous birds, such as the great tit in Europe.
+An "evolutionary arms race" can be seen between predator and prey species. The Lepidoptera have developed a number of strategies for defense and protection, including evolution of morphological characters and changes in ecological lifestyles and behaviors. These include aposematism, mimicry, camouflage, and development of threat patterns and displays.[74] Only a few birds, such as the nightjars, hunt nocturnal lepidopterans. Their main predators are bats. Again, an "evolutionary race" exists, which has led to numerous evolutionary adaptations of moths to escape from their main predators, such as the ability to hear ultrasonic sounds, or even to emit sounds in some cases. Lepidopteran eggs are also preyed upon. Some caterpillars, such as the zebra swallowtail butterfly larvae, are cannibalistic.
+Some species of Lepidoptera are poisonous to predators, such as the monarch butterfly in the Americas, Atrophaneura species (roses, windmills, etc.) in Asia, as well as Papilio antimachus, and the birdwings, the largest butterflies in Africa and Asia, respectively. They obtain their toxicity by sequestering the chemicals from the plants they eat into their own tissues. Some Lepidoptera manufacture their own toxins. Predators that eat poisonous butterflies and moths may become sick and vomit violently, learning not to eat those species. A predator which has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future, thus saving many other species, as well.[74][75] Toxic butterflies and larvae tend to develop bright colors and striking patterns as an indicator to predators about their toxicity. This phenomenon is known as aposematism.[76] Some caterpillars, especially members of Papilionidae, contain an osmeterium, a Y-shaped protrusible gland found in the prothoracic segment of the larvae.[74] When threatened, the caterpillar emits unpleasant smells from the organ to ward off the predators.[77][78]
+Camouflage is also an important defense strategy, which involves the use of coloration or shape to blend into the surrounding environment. Some lepidopteran species blend with their surroundings, making them difficult to spot by predators. Caterpillars can exhibit shades of green that match its host plant. Others look like inedible objects, such as twigs or leaves. For instance, the mourning cloak fades into the backdrop of trees when it folds its wings back. The larvae of some species, such as the common Mormon (Papilio polytes) and the western tiger swallowtail look like bird droppings.[74][79] For example, adult Sesiidae species (also known as clearwing moths) have a general appearance sufficiently similar to a wasp or hornet to make it likely the moths gain a reduction in predation by Batesian mimicry.[80] Eyespots are a type of automimicry used by some butterflies and moths. In butterflies, the spots are composed of concentric rings of scales in different colors. The proposed role of the eyespots is to deflect attention of predators. Their resemblance to eyes provokes the predator's instinct to attack these wing patterns.[81]
+Batesian and Müllerian mimicry complexes are commonly found in Lepidoptera. Genetic polymorphism and natural selection give rise to otherwise edible species (the mimic) gaining a survival advantage by resembling inedible species (the model). Such a mimicry complex is referred to as Batesian and is most commonly known in the example between the limenitidine viceroy butterfly in relation to the inedible danaine monarch. The viceroy is, in fact, more toxic than the monarch and this resemblance should be considered as a case of Müllerian mimicry.[82] In Müllerian mimicry, inedible species, usually within a taxonomic order, find it advantageous to resemble each other so as to reduce the sampling rate by predators that need to learn about the insects' inedibility. Taxa from the toxic genus Heliconius form one of the most well-known Müllerian complexes.[83] The adults of the various species now resemble each other so well, the species cannot be distinguished without close morphological observation and, in some cases, dissection or genetic analysis.
+Moths evidently are able to hear the range emitted by bats, which in effect causes flying moths to make evasive maneuvers because bats are a main predator of moths. Ultrasonic frequencies trigger a reflex action in the noctuid moth that cause it to drop a few inches in its flight to evade attack.[84] Tiger moths in a defense emit clicks within the same range of the bats, which interfere with the bats and foil their attempts to echolocate it.[85]
+Most species of Lepidoptera engage in some form of entomophily (more specifically psychophily and phalaenophily for butterflies and moths, respectively), or the pollination of flowers.[86] Most adult butterflies and moths feed on the nectar inside flowers, using their probosces to reach the nectar hidden at the base of the petals. In the process, the adults brush against the flowers' stamens, on which the reproductive pollen is made and stored. The pollen is transferred on appendages on the adults, which fly to the next flower to feed and unwittingly deposit the pollen on the stigma of the next flower, where the pollen germinates and fertilizes the seeds.[23]:813–814
+Flowers pollinated by butterflies tend to be large and flamboyant, pink or lavender in color, frequently having a landing area, and usually scented, as butterflies are typically day-flying. Since butterflies do not digest pollen (except for heliconid species,[86]) more nectar is offered than pollen. The flowers have simple nectar guides, with the nectaries usually hidden in narrow tubes or spurs, reached by the long "tongue" of the butterflies. Butterflies such as Thymelicus flavus have been observed to engage in flower constancy, which means they are more likely to transfer pollen to other conspecific plants. This can be beneficial for the plants being pollinated, as flower constancy prevents the loss of pollen during different flights and the pollinators from clogging stigmas with pollen of other flower species.[87]
+Among the more important moth pollinator groups are the hawk moths of the family Sphingidae. Their behavior is similar to hummingbirds, i.e., using rapid wing beats to hover in front of flowers. Most hawk moths are nocturnal or crepuscular, so moth-pollinated flowers (e.g., Silene latifolia ) tend to be white, night-opening, large, and showy with tubular corollae and a strong, sweet scent produced in the evening, night, or early morning. A lot of nectar is produced to fuel the high metabolic rates needed to power their flight.[88] Other moths (e.g., noctuids, geometrids, pyralids) fly slowly and settle on the flower. They do not require as much nectar as the fast-flying hawk moths, and the flowers tend to be small (though they may be aggregated in heads).[89]
+Mutualism is a form of biological interaction wherein each individual involved benefits in some way. An example of a mutualistic relationship would be that shared by yucca moths (Tegeculidae) and their host, yucca flowers (Liliaceae). Female yucca moths enter the host flowers, collect the pollen into a ball using specialized maxillary palps, then move to the apex of the pistil, where pollen is deposited on the stigma, and lay eggs into the base of the pistil where seeds will develop. The larvae develop in the fruit pod and feed on a portion of the seeds. Thus, both insect and plant benefit, forming a highly mutualistic relationship.[23]:814 Another form of mutualism occurs between some larvae of butterflies and certain species of ants (e.g. Lycaenidae). The larvae communicate with the ants using vibrations transmitted through a substrate, such as the wood of a tree or stems, as well as using chemical signals.[90] The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions.[91]
+Only 42 species of parasitoid lepidopterans are known (1 Pyralidae; 40 Epipyropidae).[23]:748 The larvae of the greater and lesser wax moths feed on the honeycomb inside bee nests and may become pests; they are also found in bumblebee and wasp nests, albeit to a lesser extent. In northern Europe, the wax moth is regarded as the most serious parasitoid of the bumblebee, and is found only in bumblebee nests. In some areas in southern England, as much as 80% of nests can be destroyed.[92] Other parasitic larvae are known to prey upon cicadas and leaf hoppers.[93]
+In reverse, moths and butterflies may be subject to parasitic wasps and flies, which may lay eggs on the caterpillars, which hatch and feed inside its body, resulting in death. Although, in a form of parasitism called idiobiont, the adult paralyzes the host, so as not to kill it but for it to live as long as possible, for the parasitic larvae to benefit the most. In another form of parasitism, koinobiont, the species live off their hosts while inside (endoparasitic). These parasites live inside the host caterpillar throughout its life cycle, or may affect it later on as an adult. In other orders, koinobionts include flies, a majority of coleopteran, and many hymenopteran parasitoids.[23]:748–749 Some species may be subject to a variety of parasites, such as the gypsy moth (Lymantaria dispar), which is attacked by a series of 13 species, in six different taxa throughout its life cycle.[23]:750
+In response to a parasitoid egg or larva in the caterpillar's body, the plasmatocytes, or simply the host's cells can form a multilayered capsule that eventually causes the endoparasite to asphyxiate. The process, called encapsulation, is one of the caterpillar's only means of defense against parasitoids.[23]:748
+A few species of Lepidoptera are secondary consumers, or predators. These species typically prey upon the eggs of other insects, aphids, scale insects, or ant larvae.[23]:567 Some caterpillars are cannibals, and others prey on caterpillars of other species (e.g. Hawaiian Eupithecia ). Those of the 15 species in Eupithecia that mirror inchworms, are the only known species of butterflies and moths that are ambush predators.[94] Four species are known to eat snails. For example, the Hawaiian caterpillar (Hyposmocoma molluscivora) uses silk traps, in a manner similar to that of spiders, to capture certain species of snails (typically Tornatellides).[93]
+Larvae of some species of moths in the Tineidae, Gelechioidae, and Noctuidae (family/superfamily/families, respectively), besides others, feed on detritus, or dead organic material, such as fallen leaves and fruit, fungi, and animal products, and turn it into humus.[23]:567 Well-known species include the cloth moths (Tineola bisselliella, Tinea pellionella, and Trichophaga tapetzella), which feed on detritus containing keratin, including hair, feathers, cobwebs, bird nests (particularly of domestic pigeons, Columba livia domestica) and fruits or vegetables. These species are important to ecosystems as they remove substances that would otherwise take a long time to decompose.[95]
+In 2015 it was reported that wasp bracovirus DNA was present in Lepidoptera such as monarch butterflies, silkworms and moths.[96] These were described in some newspaper articles as examples of a naturally occurring genetically engineered insects.[97]
+Linnaeus in Systema Naturae (1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx and Phalaena, with seven subgroups in Phalaena.[98] These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Michael Denis & Ignaz Schiffermüller (1775), Johan Christian Fabricius (1775) and Pierre André Latreille (1796). Jacob Hübner described many genera, and the lepidopteran genera were catalogued by Ferdinand Ochsenheimer and Georg Friedrich Treitschke in a series of volumes on the lepidopteran fauna of Europe published between 1807 and 1835.[98] Gottlieb August Wilhelm Herrich-Schäffer (several volumes, 1843–1856), and Edward Meyrick (1895) based their classifications primarily on wing venation. Sir George Francis Hampson worked on the microlepidoptera during this period and Philipp Christoph Zeller published The Natural History of the Tineinae also on microlepidoptera (1855).
+Among the first entomologists to study fossil insects and their evolution was Samuel Hubbard Scudder (1837–1911), who worked on butterflies.[99] He published a study of the Florissant deposits of Colorado, including the exceptionally preserved Prodryas persephone. Andreas V. Martynov (1879–1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.[99]
+Major contributions in the 20th century included the creation of the monotrysia and ditrysia (based on female genital structure) by Borner in 1925 and 1939.[98] Willi Hennig (1913–1976) developed the cladistic methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among monotrysian families and Kristensen worked more generally on insect phylogeny and higher Lepidoptera too.[98][99] While it is often found that DNA-based phylogenies differ from those based on morphology, this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.[99]
+Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not monophyletic: Microlepidoptera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.[98]
+The fossil record for Lepidoptera is lacking in comparison to other winged species, and tends not to be as common as some other insects in habitats that are most conducive to fossilization, such as lakes and ponds; their juvenile stage has only the head capsule as a hard part that might be preserved. The location and abundance of the most common moth species are indicative that mass migrations of moths occurred over the Palaeogene North Sea, which is why there is a serious lack of moth fossils.[100] Yet there are fossils, some preserved in amber and some in very fine sediments. Leaf mines are also seen in fossil leaves, although the interpretation of them is tricky.[99]
+Putative fossil stem group representatives of Amphiesmenoptera (the clade comprising Trichoptera and Lepidoptera) are known from the Triassic.[23]:567 The earliest known lepidopteran fossils are fossilized scales from the Triassic-Jurassic boundary. They were found as rare palynological elements in the sediments of the Triassic-Jurassic boundary from the cored Schandelah-1 well, drilled near Braunschweig in northern Germany. This pushes back the fossil record and origin of glossatan lepidopterans by about 70 million years, supporting molecular estimates of a Norian (ca 212 million years) divergence of glossatan and non-glossatan lepidopterans. The findings were reported in 2018 in the journal Science Advances. The authors of the study proposed that lepidopterans evolved a proboscis as an adaptation to drink from droplets and thin films of water for maintaining their fluid balance in the hot and arid climate of the Triassic.[101]
+The earliest described lepidopteran taxon is Archaeolepis mane, a primitive moth-like species from the Jurassic, dated back to around 190 million years ago, and known only from three wings found in Dorset, UK. The wings show scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared with Trichoptera (caddisflies).[102][103] Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from the Cretaceous, which all belong to primitive moth-like families.[99]
+Many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber. The oldest genuine butterflies of the superfamily Papilionoidea have been found in the Paleocene MoClay or Fur Formation of Denmark. The best preserved fossil lepidopteran is the Eocene Prodryas persephone from the Florissant Fossil Beds.
+Lepidoptera and Trichoptera (caddisflies) are sister groups, sharing many similarities that are lacking in others; for example the females of both orders are heterogametic, meaning they have two different sex chromosomes, whereas in most species the males are heterogametic and the females have two identical sex chromosomes. The adults in both orders display a particular wing venation pattern on their forewings. The larvae in the two orders have mouth structures and glands with which they make and manipulate silk. Willi Hennig grouped the two orders into the superorder Amphiesmenoptera; together they are sister to the extinct order Tarachoptera.[104] Lepidoptera descend from a diurnal moth-like common ancestor that either fed on dead or living plants.[105]
+The cladogram, based on a 2008 DNA and protein analysis, shows the order as a clade, sister to the Trichoptera, and more distantly related to the Diptera (true flies) and Mecoptera (scorpionflies).[106][107][108][109]
+Diptera (true flies)
+Mecoptera (scorpionflies)
+Boreidae (snow scorpionflies)
+Siphonaptera (fleas)
+Trichoptera (caddisflies)
+Lepidoptera (butterflies and moths)
+Hymenoptera (sawflies, wasps, ants, bees)
+Micropterigidae, Agathiphagidae and Heterobathmiidae are the oldest and most basal lineages of Lepidoptera. The adults of these families do not have the curled tongue or proboscis, that are found in most members of the order, but instead have chewing mandibles adapted for a special diet. Micropterigidae larvae feed on leaves, fungi, or liverworts (much like the Trichoptera).[98] Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live inside kauri pines and feed on seeds. In Heterobathmiidae the larvae feed on the leaves of Nothofagus, the southern beech tree. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon after metamorphosis.[98]
+The Eriocraniidae have a short coiled proboscis in the adult stage, and though they retain their pupal mandibles with which they escaped the cocoon, their mandibles are non-functional thereafter.[98] Most of these non-ditrysian families, are primarily leaf miners in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.[99]
+With the evolution of the Ditrysia in the mid-Cretaceous, there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is one cloaca, or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, Gracillarioidea and Gelechioidea are mostly leaf miners, but more recent lineages feed externally. In the Tineoidea, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.[99]
+The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.[99] They evolved about the time that flowering plants underwent an expansive adaptive radiation in the mid-Cretaceous, and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved coevolution or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.
+In the so-called "macrolepidoptera", which constitutes about 60% of lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant.[99] Many also have tympanal organs, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.[99]
+The main lineages in the macrolepidoptera are the Noctuoidea, Bombycoidea, Lasiocampidae, Mimallonoidea, Geometroidea and Rhopalocera. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a monophyletic group.[99] The Rhopalocera, comprising the Papilionoidea (butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved.[98] There is quite a good fossil record for this group, with the oldest skipper dating from 56 million years ago.[99]
+Taxonomy is the classification of species in selected taxa, the process of naming being called nomenclature. There are over 120 families in Lepidoptera, in 45 to 48 superfamilies. Lepidoptera have always been, historically, classified in five suborders, one of which is of primitive moths that never lost the morphological features of their ancestors. The rest of the moths and butterflies make up ninety-eight percent of the other taxa, making Ditrysia. More recently, findings of new taxa, larvae and pupa have aided in detailing the relationships of primitive taxa, phylogenetic analysis showing the primitive lineages to be paraphyletic compared to the rest of Lepidoptera lineages. Recently lepidopterists have abandoned clades like suborders, and those between orders and superfamilies.[23]:569
+Artistic depictions of butterflies have been used in many cultures including as early as 3500 years ago, in Egyptian hieroglyphs.[114] Today, butterflies are widely used in various objects of art and jewelry: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and in some mixed media artworks and furnishings.[115] Butterflies have also inspired the "butterfly fairy" as an art and fictional character.
+In many cultures the soul of a dead person is associated with the butterfly, for example in Ancient Greece, where the word for butterfly ψυχή (psyche) also means soul and breath. In Latin, as in Ancient Greece, the word for "butterfly" papilio was associated with the soul of the dead.[116] The skull-like marking on the thorax of the death's-head hawkmoth has helped these moths, particularly A. atropos, earn a negative reputation, such as associations with the supernatural and evil. The moth has been prominently featured in art and movies such as Un Chien Andalou (by Buñuel and Dalí) and The Silence of the Lambs, and in the artwork of the Japanese metal band Sigh's album Hail Horror Hail. According to Kwaidan: Stories and Studies of Strange Things, by Lafcadio Hearn, a butterfly was seen in Japan as the personification of a person's soul; whether they be living, dying, or already dead. One Japanese superstition says that if a butterfly enters your guestroom and perches behind the bamboo screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad omens. When Taira no Masakado was secretly preparing for his famous revolt, there appeared in Kyoto so vast a swarm of butterflies that the people were frightened—thinking the apparition to be a portent of coming evil.[117]
+In the ancient Mesoamerican city of Teotihuacan, the brilliantly colored image of the butterfly was carved into many temples, buildings, jewelry, and emblazoned on incense burners in particular. The butterfly was sometimes depicted with the maw of a jaguar and some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies to fire and warfare persisted through to the Aztec civilization and evidence of similar jaguar-butterfly images has been found among the Zapotec, and Maya civilizations.[118]
+The larvae of many lepidopteran species are major pests in agriculture. Some of the major pests include Tortricidae, Noctuidae, and Pyralidae. The larvae of the Noctuidae genus Spodoptera (armyworms), Helicoverpa (corn earworm), or Pieris brassicae can cause extensive damage to certain crops.[98] Helicoverpa zea larvae (cotton bollworms or tomato fruitworms) are polyphagous, meaning they eat a variety of crops, including tomatoes and cotton.[119] Peridroma saucia (variegated cutworms) are described as one of the most damaging pests to gardens, with the ability to destroy entire gardens and fields in a matter of days.[120]
+Butterflies and moths are one of the largest taxa to solely feed and be dependent on living plants, in terms of the number of species, and they are in many ecosystems, making up the largest biomass to do so. In many species, the female may produce anywhere from 200 to 600 eggs, while in some others it may go as high as 30,000 eggs in one day. This can create many problems for agriculture, where many caterpillars can affect acres of vegetation. Some reports estimate that there have been over 80,000 caterpillars of several different taxa feeding on a single oak tree. In some cases, phytophagous larvae can lead to the destruction of entire trees in relatively short periods of time.[23]:567
+Ecological ways of removing pest Lepidoptera species are becoming more economically viable, as research has shown ways like introducing parasitic wasps and flies. For example, Sarcophaga aldrichi, a fly which deposited larvae feed upon the pupae of the forest tent caterpillar moth. Pesticides can affect other species other than the species they are targeted to eliminate, damaging the natural ecosystem.[121] Another good biological pest control method is the use of pheromone traps. A pheromone trap is a type of insect trap that uses pheromones to lure insects. Sex pheromones and aggregating pheromones are the most common types used. A pheromone-impregnated lure is encased in a conventional trap such as a Delta trap, water-pan trap, or funnel trap.[122]
+Species of moths that are detritivores would naturally eat detritus containing keratin, such as hairs or feathers. Well known species are cloth moths (T. bisselliella, T. pellionella, and T. tapetzella), feeding on foodstuffs that people find economically important, such as cotton, linen, silk and wool fabrics as well as furs; furthermore they have been found on shed feathers and hair, bran, semolina and flour (possibly preferring wheat flour), biscuits, casein, and insect specimens in museums.[95]
+Even though most butterflies and moths affect the economy negatively, some species are a valuable economic resource. The most prominent example is that of the domesticated silkworm moth (Bombyx mori), the larvae of which make their cocoons out of silk, which can be spun into cloth. Silk is and has been an important economic resource throughout history. The species Bombyx mori has been domesticated to the point where it is completely dependent on mankind for survival.[123] A number of wild moths such as Bombyx mandarina, and Antheraea species, besides others, provide commercially important silks.[124]
+The preference of the larvae of most lepidopteran species to feed on a single species or limited range of plants is used as a mechanism for biological control of weeds in place of herbicides. The pyralid cactus moth was introduced from Argentina to Australia, where it successfully suppressed millions of acres of prickly pear cactus.[23]:567 Another species of the Pyralidae, called the alligator weed stem borer (Arcola malloi), was used to control the aquatic plant known as alligator weed (Alternanthera philoxeroides) in conjunction with the alligator weed flea beetle; in this case, the two insects work in synergy and the weed rarely recovers.[125]
+Breeding butterflies and moths, or butterfly gardening/rearing, has become an ecologically viable process of introducing species into the ecosystem to benefit it. Butterfly ranching in Papua New Guinea permits nationals of that country to "farm" economically valuable insect species for the collectors market in an ecologically sustainable manner.[126]
+Lepidoptera feature prominently in entomophagy as food items on almost every continent. While in most cases, adults, larvae or pupae are eaten as staples by indigenous people, beondegi or silkworm pupae are eaten as a snack in Korean cuisine[127] while Maguey worm is considered a delicacy in Mexico.[128] In some parts of Huasteca, the silk nests of the Madrone butterfly are maintained on the edge of roof tops of houses for consumption.[129] In the Carnia region of Italy, children catch and eat ingluvies of the toxic Zygaena moths in early summer. The ingluvies, despite having a very low cyanogenic content, serve as a convenient, supplementary source of sugar to the children who can include this resource as a seasonal delicacy at minimum risk.[130]
+Some larvae of both moths and butterflies have a form of hair that has been known to be a cause of human health problems. Caterpillar hairs sometimes have toxins in them and species from approximately 12 families of moths or butterflies worldwide can inflict serious human injuries (urticarial dermatitis and atopic asthma to osteochondritis, consumption coagulopathy, renal failure, and intracerebral hemorrhage).[131] Skin rashes are the most common, but there have been fatalities.[132] Lonomia is a frequent cause of envenomation in humans in Brazil, with 354 cases reported between 1989 and 2005. Lethality ranging up to 20% with death caused most often by intracranial hemorrhage.[133]
+These hairs have also been known to cause keratoconjunctivitis. The sharp barbs on the end of caterpillar hairs can get lodged in soft tissues and mucous membranes such as the eyes. Once they enter such tissues, they can be difficult to extract, often exacerbating the problem as they migrate across the membrane.[134] This becomes a particular problem in an indoor setting. The hairs easily enter buildings through ventilation systems and accumulate in indoor environments because of their small size, which makes it difficult for them to be vented out. This accumulation increases the risk of human contact in indoor environments.[135]

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+Aglossata
+Glossata
+Heterobathmiina
+Zeugloptera
+Lepidoptera (/ˌlɛpɪˈdɒptərə/ LEP-i-DOP-tər-ə, from Ancient Greek lepís “scale” + pterón “wing”) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families[1] and 46 superfamilies,[2] 10 per cent of the total described species of living organisms.[2][3] It is one of the most widespread and widely recognizable insect orders in the world.[4] The Lepidoptera show many variations of the basic body structure that have evolved to gain advantages in lifestyle and distribution. Recent estimates suggest the order may have more species than earlier thought,[5] and is among the four most speciose orders, along with the Hymenoptera, Diptera, and Coleoptera.[4]
+Lepidopteran species are characterized by more than three derived features. The most apparent is the presence of scales that cover the bodies, wings, and a proboscis. The scales are modified, flattened "hairs", and give butterflies and moths their wide variety of colors and patterns. Almost all species have some form of membranous wings, except for a few that have reduced wings or are wingless. Mating and the laying of eggs are carried out by adults, normally near or on host plants for the larvae.  Like most other insects, butterflies and moths are holometabolous, meaning they undergo complete metamorphosis.  The larvae are commonly called caterpillars, and are completely different from their adult moth or butterfly forms, having a cylindrical body with a well-developed head, mandible mouth parts, three pairs of thoracic legs and from none up to five pairs of prolegs. As they grow, these larvae change in appearance, going through a series of stages called instars. Once fully matured, the larva develops into a pupa. A few butterflies and many moth species spin a silk case or cocoon prior to pupating, while others do not, instead going underground.[4] A butterfly pupa, called a chrysalis, has a hard skin, usually with no cocoon. Once the pupa has completed its metamorphosis, a sexually mature adult emerges.
+The Lepidoptera have, over millions of years, evolved a wide range of wing patterns and coloration ranging from drab moths akin to the related order Trichoptera, to the brightly colored and complex-patterned butterflies.[1] Accordingly, this is the most recognized and popular of insect orders with many people involved in the observation, study, collection, rearing of, and commerce in these insects. A person who collects or studies this order is referred to as a lepidopterist.
+Butterflies and moths play an important role in the natural ecosystem as pollinators and as food in the food chain; conversely, their larvae are considered very problematic to vegetation in agriculture, as their main source of food is often live plant matter. In many species, the female may produce from 200 to 600 eggs, while in others, the number may approach 30,000 eggs in one day. The caterpillars hatching from these eggs can cause damage to large quantities of crops. Many moth and butterfly species are of economic interest by virtue of their role as pollinators, the silk they produce, or as pest species.
+The term was coined by Linnaeus in 1735 and is derived from Greek λεπίς, gen. λεπίδος ("scale") and πτερόν ("wing").[6] Sometimes, the term Rhopalocera is used for the clade of all butterfly species, derived from the Ancient Greek ῥόπαλον (rhopalon)[7]:4150 and κέρας (keras)[7]:3993 meaning "club" and "horn", respectively, coming from the shape of the antennae of butterflies.
+The origins of the common names "butterfly" and "moth" are varied and often obscure. The English word butterfly is from Old English buttorfleoge, with many variations in spelling. Other than that, the origin is unknown, although it could be derived from the pale yellow color of many species' wings suggesting the color of butter.[8][9] The species of Heterocera are commonly called moths. The origins of the English word moth are more clear, deriving from the Old English moððe" (cf. Northumbrian dialect mohðe) from Common Germanic (compare Old Norse motti, Dutch mot and German Motte all meaning "moth"). Perhaps its origins are related to Old English maða meaning "maggot" or from the root of "midge", which until the 16th century was used mostly to indicate the larva, usually in reference to devouring clothes.[10]
+The etymological origins of the word "caterpillar", the larval form of butterflies and moths, are from the early 16th century, from Middle English catirpel, catirpeller, probably an alteration of Old North French catepelose: cate, cat (from Latin cattus) + pelose, hairy (from Latin pilōsus).[11]
+The Lepidoptera are among the most successful groups of insects. They are found on all continents, except Antarctica, and inhabit all terrestrial habitats ranging from desert to rainforest, from lowland grasslands to mountain plateaus, but almost always associated with higher plants, especially angiosperms (flowering plants).[12] Among the most northern dwelling species of butterflies and moths is the Arctic Apollo (Parnassius arcticus), which is found in the Arctic Circle in northeastern Yakutia, at an altitude of 1500 m above sea level.[13] In the Himalayas, various Apollo species such as Parnassius epaphus  have been recorded to occur up to an altitude of 6,000 m above sea level.[14]:221
+Some lepidopteran species exhibit symbiotic, phoretic, or parasitic lifestyles, inhabiting the bodies of organisms rather than the environment. Coprophagous pyralid moth species, called sloth moths, such as Bradipodicola hahneli and Cryptoses choloepi, are unusual in that they are exclusively found inhabiting the fur of sloths, mammals found in Central and South America.[15][16]
+Two species of Tinea moths have been recorded as feeding on horny tissue and have been bred from the horns of cattle. The larva of Zenodochium coccivorella is an internal parasite of the coccid Kermes species. Many species have been recorded as breeding in natural materials or refuse such as owl pellets, bat caves, honeycombs or diseased fruit.[16]
+As of 2007, there was roughly 174,250 lepidopteran species described, with butterflies and skippers estimated to comprise around 17,950, and moths making up the rest.[2][17] The vast majority of Lepidoptera are to be found in the tropics, but substantial diversity exists on most continents. North America has over 700 species of butterflies and over 11,000 species of moths,[18][19] while about 400 species of butterflies and 14,000 species of moths are reported from Australia.[20] The diversity of Lepidoptera in each faunal region has been estimated by John Heppner in 1991 based partly on actual counts from the literature, partly on the card indices in the Natural History Museum (London) and the National Museum of Natural History (Washington), and partly on estimates:[5]
+Lepidoptera are morphologically distinguished from other orders principally by the presence of scales on the external parts of the body and appendages, especially the wings. Butterflies and moths vary in size from microlepidoptera only a few millimeters long, to conspicuous animals with a wingspan greater than 25 centimetres, such as the Queen Alexandra's birdwing and Atlas moth.[21]:246
+Lepidopterans undergo a four-stage life cycle: egg; larva or caterpillar; pupa or chrysalis; and imago (plural: imagines) / adult and show many variations of the basic body structure, which give these animals advantages for diverse lifestyles and environments.
+The head is where many sensing organs and the mouth parts are found. Like the adult, the larva also has a toughened, or sclerotized head capsule.[22] Here, two compound eyes, and chaetosema, raised spots or clusters of sensory bristles unique to Lepidoptera, occur, though many taxa have lost one or both of these spots. The antennae have a wide variation in form among species and even between different sexes. The antennae of butterflies are usually filiform and shaped like clubs, those of the skippers are hooked, while those of moths have flagellar segments variously enlarged or branched. Some moths have enlarged antennae or ones that are tapered and hooked at the ends.[23]:559–560
+The maxillary galeae are modified and form an elongated proboscis. The proboscis consists of one to five segments, usually kept coiled up under the head by small muscles when it is not being used to suck up nectar from flowers or other liquids. Some basal moths still have mandibles, or separate moving jaws, like their ancestors, and these form the family Micropterigidae.[22][23]:560[24]
+The larvae, called caterpillars, have a toughened head capsule. Caterpillars lack the proboscis and have separate chewing mouthparts.[22] These mouthparts, called mandibles, are used to chew up the plant matter that the larvae eat. The lower jaw, or labium, is weak, but may carry a spinneret, an organ used to create silk. The head is made of large lateral lobes, each having an ellipse of up to six simple eyes.[23]:562–563
+The thorax is made of three fused segments, the prothorax, mesothorax, and metathorax, each with a pair of legs. The first segment contains the first pair of legs. In some males of the butterfly family Nymphalidae, the forelegs are greatly reduced and are not used for walking or perching.[23]:586 The three pairs of legs are covered with scales. Lepidoptera also have olfactory organs on their feet, which aid the butterfly in "tasting" or "smelling" out its food.[25] In the larval form there are 3 pairs of true legs, with up to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[12]
+The two pairs of wings are found on the middle and third segments, or mesothorax and metathorax, respectively. In the more recent genera, the wings of the second segment are much more pronounced, although some more primitive forms have similarly sized wings of both segments. The wings are covered in scales arranged like shingles, which form an extraordinary variety of colors and patterns. The mesothorax has more powerful muscles to propel the moth or butterfly through the air, with the wing of this segment (forewing) having a stronger vein structure.[23]:560 The largest superfamily, the Noctuidae, has their wings modified to act as tympanal or hearing organs.[26]
+The caterpillar has an elongated, soft body that may have hair-like or other projections, three pairs of true legs, with none to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[12] The thorax usually has a pair of legs on each segment. The thorax is also lined with many spiracles on both the mesothorax and metathorax, except for a few aquatic species, which instead have a form of gills.[23]:563
+The abdomen, which is less sclerotized than the thorax, consists of 10 segments with membranes in between, allowing for articulated movement. The sternum, on the first segment, is small in some families and is completely absent in others. The last two or three segments form the external parts of the species' sex organs. The genitalia of Lepidoptera are highly varied and are often the only means of differentiating between species. Male genitals include a valva, which is usually large, as it is used to grasp the female during mating. Female genitalia include three distinct sections.
+The females of basal moths have only one sex organ, which is used for copulation and as an ovipositor, or egg-laying organ. About 98% of moth species have a separate organ for mating, and an external duct that carries the sperm from the male.[23]:561
+The abdomen of the caterpillar has four pairs of prolegs, normally located on the third to sixth segments of the abdomen, and a separate pair of prolegs by the anus, which have a pair of tiny hooks called crotchets. These aid in gripping and walking, especially in species that lack many prolegs (e. g. larvae of Geometridae). In some basal moths, these prolegs may be on every segment of the body, while prolegs may be completely absent in other groups, which are more adapted to boring and living in sand (e. g., Prodoxidae and Nepticulidae, respectively).[23]:563
+The wings, head, and parts of the thorax and abdomen of Lepidoptera are covered with minute scales, a feature from which the order derives its name. Most scales are lamellar, or blade-like, and attached with a pedicel, while other forms may be hair-like or specialized as secondary sexual characteristics.[27]
+The lumen or surface of the lamella has a complex structure. It gives color either by colored pigments it contains, or through structural coloration with mechanisms that include photonic crystals and diffraction gratings.[28]
+Scales function in insulation, thermoregulation, producing pheromones (in males only),[29] and aiding gliding flight, but the most important is the large diversity of vivid or indistinct patterns they provide, which help the organism protect itself by camouflage or mimicry, and which act as signals to other animals including rivals and potential mates.[27]
+In the reproductive system of butterflies and moths, the male genitalia are complex and unclear. In females the three types of genitalia are based on the relating taxa: 'monotrysian', 'exoporian', and 'ditrysian'. In the monotrysian type is an opening on the fused segments of the sterna 9 and 10, which act as insemination and oviposition. In the exoporian type (in Hepaloidae and Mnesarchaeoidea) are two separate places for insemination and oviposition, both occurring on the same sterna as the monotrysian type, i.e. 9 and 10.[21] The ditrysian groups have an internal duct that carries sperm, with separate openings for copulation and egg-laying.[4] In most species, the genitalia are flanked by two soft lobes, although they may be specialized and sclerotized in some species for ovipositing in area such as crevices and inside plant tissue.[21] Hormones and the glands that produce them run the development of butterflies and moths as they go through their life cycles, called the endocrine system. The first insect hormone prothoracicotropic hormone (PTTH) operates the species life cycle and diapause.[30] This hormone is produced by corpora allata and corpora cardiaca, where it is also stored. Some glands are specialized to perform certain task such as producing silk or producing saliva in the palpi.[31]:65, 75 While the corpora cardiaca produce PTTH, the corpora allata also produces juvenile hormones, and the prothorocic glands produce moulting hormones.
+In the digestive system, the anterior region of the foregut has been modified to form a pharyngeal  sucking pump as they need it for the food they eat, which are for the most part liquids. An esophagus follows and leads to the posterior of the pharynx and in some species forms a form of crop. The midgut is short and straight, with the hindgut being longer and coiled.[21] Ancestors of lepidopteran species, stemming from Hymenoptera, had midgut ceca, although this is lost in current butterflies and moths. Instead, all the digestive enzymes, other than initial digestion, are immobilized at the surface of the midgut cells. In larvae, long-necked and stalked goblet cells are found in the anterior and posterior midgut regions, respectively. In insects, the goblet cells excrete positive potassium ions, which are absorbed from leaves ingested by the larvae. Most butterflies and moths display the usual digestive cycle, but species with different diets require adaptations to meet these new demands.[23]:279
+In the circulatory system, hemolymph, or insect blood, is used to circulate heat in a form of thermoregulation, where muscles contraction produces heat, which is transferred to the rest of the body when conditions are unfavorable.[32]
+In lepidopteran species, hemolymph is circulated through the veins in the wings by some form of pulsating organ, either by the heart or by the intake of air into the trachea.[31]:69
+Air is taken in through spiracles along the sides of the abdomen and thorax supplying the trachea with oxygen as it goes through the lepidopteran's respiratory system. Three different tracheaes supply and diffuse oxygen throughout the species' bodies. The dorsal tracheae supply oxygen to the dorsal musculature and vessels, while the ventral tracheae supply the ventral musculature and nerve cord, and the visceral tracheae supply the guts, fat bodies, and gonads.[31]:71, 72
+Polymorphism is the appearance of forms or "morphs", which differ in color and number of attributes within a single species.[12]:163[33] In Lepidoptera, polymorphism can be seen not only between individuals in a population, but also between the sexes as sexual dimorphism, between geographically separated populations in geographical polymorphism, and between generations flying at different seasons of the year (seasonal polymorphism or polyphenism). In some species, the polymorphism is limited to one sex, typically the female. This often includes the phenomenon of mimicry when mimetic morphs fly alongside nonmimetic morphs in a population of a particular species. Polymorphism occurs both at specific level with heritable variation in the overall morphological adaptations of individuals, as well as in certain specific morphological or physiological traits within a species.[12]
+Environmental polymorphism, in which traits are not inherited, is often termed as polyphenism, which in Lepidoptera is commonly seen in the form of seasonal morphs, especially in the butterfly families of Nymphalidae and Pieridae. An Old World pierid butterfly, the common grass yellow (Eurema hecabe) has a darker summer adult morph, triggered by a long day exceeding 13 hours in duration, while the shorter diurnal period of 12 hours or less induces a paler morph in the postmonsoon period.[34] Polyphenism also occurs in caterpillars, an example being the peppered moth, Biston betularia.[35]
+Geographical isolation causes a divergence of a species into different morphs. A good example is the Indian white admiral Limenitis procris, which has five forms, each geographically separated from the other by large mountain ranges.[36]:26 An even more dramatic showcase of geographical polymorphism is the Apollo butterfly (Parnassius apollo). Because the Apollos live in small local populations, thus having no contact with each other, coupled with their strong stenotopic nature and weak migration ability, interbreeding between populations of one species practically does not occur; by this, they form over 600 different morphs, with the size of spots on the wings of which varies greatly.[37]
+Sexual dimorphism is the occurrence of differences between males and females in a species. In Lepidoptera, it is widespread and almost completely set by genetic determination.[34] Sexual dimorphism is present in all families of the Papilionoidea and more prominent in the Lycaenidae, Pieridae, and certain taxa of the Nymphalidae. Apart from color variation, which may differ from slight to completely different color-pattern combinations, secondary sexual characteristics may also be present.[36]:25 Different genotypes maintained by natural selection may also be expressed at the same time.[34] Polymorphic and/or mimetic females occur in the case of some taxa in the Papilionidae primarily to obtain a level of protection not available to the male of their species. The most distinct case of sexual dimorphism is that of adult females of many Psychidae species which have only vestigial wings, legs, and mouthparts as compared to the adult males that are strong fliers with well-developed wings and feathery antennae.[38]
+Species of Lepidoptera undergo holometabolism or "complete metamorphosis". Their life cycle normally consists of an egg, a larva, a pupa, and an imago or adult.[12] The larvae are commonly called caterpillars, and the pupae of moths encapsulated in silk are called cocoons, while the uncovered pupae of butterflies are called chrysalides.
+Unless the species reproduces year-round, a butterfly or moth may enter diapause, a state of dormancy that allows the insect to survive unfavorable environmental conditions.
+Males usually start eclosion (emergence) earlier than females and peak in numbers before females. Both of the sexes are sexually mature by the time of eclosion.[23]:564 Butterflies and moths normally do not associate with each other, except for migrating species, staying relatively asocial. Mating begins with an adult (female or male) attracting a mate, normally using visual stimuli, especially in diurnal species like most butterflies. However, the females of most nocturnal species, including almost all moth species, use pheromones to attract males, sometimes from long distances.[12] Some species engage in a form of acoustic courtship, or attract mates using sound or vibration such as the polka-dot wasp moth, Syntomeida epilais.[39]
+Adaptations include undergoing one seasonal generation, two or even more, called voltinism (Univoltism, bivoltism, and multivism, respectively). Most lepidopterans in temperate climates are univoltine, while in tropical climates most have two seasonal broods. Some others may take advantage of any opportunity they can get, and mate continuously throughout the year. These seasonal adaptations are controlled by hormones, and these delays in reproduction are called diapause.[23]:567 Many lepidopteran species, after mating and laying their eggs, die shortly afterwards, having only lived for a few days after eclosion. Others may still be active for several weeks and then overwinter and become sexually active again when the weather becomes more favorable, or diapause. The sperm of the male that mated most recently with the female is most likely to have fertilized the eggs, but the sperm from a prior mating may still prevail.[23]:564
+Lepidoptera usually reproduce sexually and are oviparous (egg-laying), though some species exhibit live birth in a process called ovoviviparity. A variety of differences in egg-laying and the number of eggs laid occur. Some species simply drop their eggs in flight (these species normally have polyphagous larvae, meaning they eat a variety of plants e. g., hepialids and some nymphalids)[40] while most lay their eggs near or on the host plant on which the larvae feed. The number of eggs laid may vary from only a few to several thousand.[12] The females of both butterflies and moths select the host plant instinctively, and primarily, by chemical cues.[23]:564
+The eggs are derived from materials ingested as a larva and in some species, from the spermatophores received from males during mating.[41] An egg can only be 1/1000 the mass of the female, yet she may lay up to her own mass in eggs. Females lay smaller eggs as they age. Larger females lay larger eggs.[42] The egg is covered by a hard-ridged protective outer layer of shell, called the chorion. It is lined with a thin coating of wax, which prevents the egg from drying out. Each egg contains a number of micropyles, or tiny funnel-shaped openings at one end, the purpose of which is to allow sperm to enter and fertilize the egg. Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate.
+The egg stage lasts a few weeks in most butterflies, but eggs laid prior to winter, especially in temperate regions, go through diapause, and hatching may be delayed until spring. Other butterflies may lay their eggs in the spring and have them hatch in the summer. These butterflies are usually temperate species (e. g. Nymphalis antiopa).
+The larvae or caterpillars are the first stage in the life cycle after hatching. Caterpillars, are "characteristic polypod larvae with cylindrical bodies, short thoracic legs, and abdominal prolegs (pseudopods)".[43] They have a toughened (sclerotised) head capsule with an adfrontal suture formed by medial fusion of the sclerites, mandibles (mouthparts) for chewing, and a soft tubular, segmented body, that may have hair-like or other projections, three pairs of true legs, and additional prolegs (up to five pairs).[44] The body consists of thirteen segments, of which three are thoracic and ten are abdominal.[45] Most larvae are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores.[44]
+Different herbivorous species have adapted to feed on every part of the plant and are normally considered pests to their host plants; some species have been found to lay their eggs on the fruit and other species lay their eggs on clothing or fur (e. g., Tineola bisselliella, the common clothes moth). Some species are carnivorous and others are even parasitic. Some lycaenid species such as Phengaris rebeli are social parasites of Myrmica ants nests.[46] A species of Geometridae from Hawaii has carnivorous larvae that catch and eat flies.[47] Some pyralid caterpillars are aquatic.[48]
+The larvae develop rapidly with several generations in a year; however, some species may take up to 3 years to develop, and exceptional examples like Gynaephora groenlandica take as long as seven years.[12] The larval stage is where the feeding and growing stages occur, and the larvae periodically undergo hormone-induced ecdysis, developing further with each instar, until they undergo the final larval-pupal molt.
+The larvae of both butterflies and moths exhibit mimicry to deter potential predators. Some caterpillars have the ability to inflate parts of their heads to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called osmeteria (family Papilionidae), which are exposed to produce smelly chemicals used in defense. Host plants often have toxic substances in them and caterpillars are able to sequester these substances and retain them into the adult stage. This helps make them unpalatable to birds and other predators. Such unpalatability is advertised using bright red, orange, black, or white warning colors. The toxic chemicals in plants are often evolved specifically to prevent them from being eaten by insects. Insects, in turn, develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants.[49]
+No form of wing is externally visible on the larva, but when larvae are dissected, developing wings can be seen as disks, which can be found on the second and third thoracic segments, in place of the spiracles that are apparent on abdominal segments. Wing disks develop in association with a trachea that runs along the base of the wing, and are surrounded by a thin peripodial membrane, which is linked to the outer epidermis of the larva by a tiny duct. Wing disks are very small until the last larval instar, when they increase dramatically in size, are invaded by branching tracheae from the wing base that precede the formation of the wing veins, and begin to develop patterns associated with several landmarks of the wing.[50]
+Near pupation, the wings are forced outside the epidermis under pressure from the hemolymph, and although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval cuticle, they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled without damage to the wings.[50]
+After about five to seven instars,[51]:26–28 or molts, certain hormones, like PTTH, stimulate the production of ecdysone, which initiates insect molting. Then, the larva puparium, a sclerotized or hardened cuticle of the last larval instar, develops into the pupa. Depending on the species, the pupa may be covered in a silk cocoon, attached to different types of substrates, buried in the ground, or may not be covered at all. Features of the imago are externally recognizable in the pupa. All the appendages on the adult head and thorax are found cased inside the cuticle (antennae, mouthparts, etc.), with the wings wrapped around, adjacent to the antennae.[23]:564 The pupae of some species have functional mandibles, while the pupal mandibles are not functional in others.[22]
+While encased, some of the lower segments are not fused, and are able to move using small muscles found in between the membrane. Moving may help the pupa, for example, escape the sun, which would otherwise kill it. The pupa of the Mexican jumping bean moth (Cydia deshaisiana) does this. The larvae cut a trapdoor in the bean (species of Sebastiania) and use the bean as a shelter. With a sudden rise in temperature, the pupa inside twitches and jerks, pulling on the threads inside. Wiggling may also help to deter parasitoid wasps from laying eggs on the pupa. Other species of moths are able to make clicks to deter predators.[23]:564, 566
+The length of time before the pupa ecloses (emerges) varies greatly. The monarch butterfly may stay in its chrysalis for two weeks, while other species may need to stay for more than 10 months in diapause. The adult emerges from the pupa either by using abdominal hooks or from projections located on the head. The mandibles found in the most primitive moth families are used to escape from their cocoon (e. g., Micropterigoidea).[12][23]:564
+Most lepidopteran species do not live long after eclosion, only needing a few days to find a mate and then lay their eggs. Others may remain active for a longer period (from one to several weeks), or go through diapause and overwintering as monarch butterflies do, or waiting out environmental stress. Some adult species of microlepidoptera go through a stage where no reproductive-related activity occurs, lasting through summer and winter, followed by mating and oviposition in the early spring.[23]:564
+While most butterflies and moths are terrestrial, many species of Pyralidae are truly aquatic with all stages except the adult occurring in water. Many species from other families such as Erebidae, Nepticulidae, Cosmopterygidae, Tortricidae, Olethreutidae, Noctuidae, Cossidae, and Sphingidae are aquatic or semiaquatic.[52]:22
+Flight is an important aspect of the lives of butterflies and moths, and is used for evading predators, searching for food, and finding mates in a timely manner, as most lepidopteran species do not live long after eclosion. It is the main form of locomotion in most species. In Lepidoptera, the forewings and hindwings are mechanically coupled and flap in synchrony. Flight is anteromotoric, or being driven primarily by action of the forewings. Although lepidopteran species reportedly can still fly when their hindwings are cut off, it reduces their linear flight and turning capabilities.[53]
+Lepidopteran species have to be warm, about 77 to 79 °F (25 to 26 °C), to fly. They depend on their body temperature being sufficiently high and since they cannot regulate it themselves, this is dependent on their environment. Butterflies living in cooler climates may use their wings to warm their bodies. They will bask in the sun, spreading out their wings so that they get maximum exposure to the sunlight. In hotter climates butterflies can easily overheat, so they are usually active only during the cooler parts of the day, early morning, late afternoon or early evening. During the heat of the day, they rest in the shade. Some larger thick-bodied moths (e.g. Sphingidae) can generate their own heat to a limited degree by vibrating their wings. The heat generated by the flight muscles warms the thorax while the temperature of the abdomen is unimportant for flight. To avoid overheating, some moths rely on hairy scales, internal air sacs, and other structures to separate the thorax and abdomen and keep the abdomen cooler.[54]
+Some species of butterflies can reach fast speeds, such as the southern dart, which can go as fast as 48.4 km/h. Sphingids are some of the fastest flying insects, some are capable of flying at over 50 km/h (30 mi/h), having a wingspan of 35–150 mm.[1][55] In some species, sometimes a gliding component to their flight exists. Flight occurs either as hovering, or as forward or backward motion.[56] In butterfly and moth species, such as hawk moths, hovering is important as they need to maintain a certain stability over flowers when feeding on the nectar.[1]
+Navigation is important to Lepidoptera species, especially for those that migrate. Butterflies, which have more species that migrate, have been shown to navigate using time-compensated sun compasses. They can see polarized light, so can orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is suggested to be particularly important.[57] Most migratory butterflies are those that live in semiarid areas where breeding seasons are short.[58] The life histories of their host plants also influence the strategies of the butterflies.[59] Other theories include the use of landscapes. Lepidoptera may use coastal lines, mountains, and even roads to orient themselves. Above sea, the flight direction is much more accurate if the coast is still visible.[60]
+Many studies have also shown that moths navigate. One study showed that many moths may use the Earth's magnetic field to navigate, as a study of the moth heart and dart suggests.[61] Another study, of the migratory behavior of the silver Y, showed, even at high altitudes, the species can correct its course with changing winds, and prefers flying with favourable winds, suggesting a great sense of direction.[62][63] Aphrissa statira in Panama loses its navigational capacity when exposed to a magnetic field, suggesting it uses the Earth's magnetic field.[64]
+Moths exhibit a tendency to circle artificial lights repeatedly. This suggests they use a technique of celestial navigation called transverse orientation. By maintaining a constant angular relationship to a bright celestial light, such as the Moon, they can fly in a straight line. Celestial objects are so far away, even after traveling great distances, the change in angle between the moth and the light source is negligible; further, the moon will always be in the upper part of the visual field or on the horizon. When a moth encounters a much closer artificial light and uses it for navigation, the angle changes noticeably after only a short distance, in addition to being often below the horizon. The moth instinctively attempts to correct by turning toward the light, causing airborne moths to come plummeting downwards, and at close range, which results in a spiral flight path that gets closer and closer to the light source.[65] Other explanations have been suggested, such as the idea that moths may be impaired with a visual distortion called a Mach band by Henry Hsiao in 1972. He stated that they fly towards the darkest part of the sky in pursuit of safety, thus are inclined to circle ambient objects in the Mach band region.[66]
+Lepidopteran migration is typically seasonal, as the insects moving to escape dry seasons or other disadvantageous conditions. Most lepidopterans that migrate are butterflies, and the distance travelled varies. Some butterflies that migrate include the mourning cloak, painted lady, American lady, red admiral, and the common buckeye.[51]:29–30 A notable species of moth that migrates long distances is the bogong moth.[67] The most well-known migrations are those of the eastern population of the monarch butterfly from Mexico to northern United States and southern Canada, a distance of about 4,000–4,800 km (2,500–3,000 mi). Other well-known migratory species include the painted lady and several of the danaine butterflies. Spectacular and large-scale migrations associated with the monsoons are seen in peninsular India.[68] Migrations have been studied in more recent times using wing tags and stable hydrogen isotopes.[69][70]
+Moths also undertake migrations, an example being the uraniids. Urania fulgens undergoes population explosions and massive migrations that may be not surpassed by any other insect in the Neotropics. In Costa Rica and Panama, the first population movements may begin in July and early August and depending on the year, may be very massive, continuing unabated for as long as five months.[71]
+Pheromones are commonly involved in mating rituals among species, especially moths, but they are also an important aspect of other forms of communication. Usually, the pheromones are produced by either the male or the female and detected by members of the opposite sex with their antennae.[72] In many species, a gland between the eighth and ninth segments under the abdomen in the female produces the pheromones.[12] Communication can also occur through stridulation, or producing sounds by rubbing various parts of the body together.[63]
+Moths are known to engage in acoustic forms of communication, most often as courtship, attracting mates using sound or vibration. Like most other insects, moths pick up these sounds using tympanic membranes in their abdomens.[73] An example is that of the polka-dot wasp moth (Syntomeida epilais), which produces sounds with a frequency above that normally detectable by humans (about 20 kHz). These sounds also function as tactile communication, or communication through touch, as they stridulate, or vibrate a substrate like leaves and stems.[39]
+Most moths lack bright colors, as many species use coloration as camouflage, but butterflies engage in visual communication. Female cabbage butterflies, for example, use ultraviolet light to communicate, with scales colored in this range on the dorsal wing surface. When they fly, each down stroke of the wing creates a brief flash of ultraviolet light which the males apparently recognize as the flight signature of a potential mate. These flashes from the wings may attract several males that engage in aerial courtship displays.[73]
+Moths and butterflies are important in the natural ecosystem. They are integral participants in the food chain; having co-evolved with flowering plants and predators, lepidopteran species have formed a network of trophic relationships between autotrophs and heterotrophs, which are included in the stages of Lepidoptera larvae, pupae, and adults. Larvae and pupae are links in the diets of birds and parasitic entomophagous insects. The adults are included in food webs in a much broader range of consumers (including birds, small mammals, reptiles, etc.).[23]:567
+Lepidopteran species are soft bodied, fragile, and almost defenseless, while the immature stages move slowly or are immobile, hence all stages are exposed to predation. Adult butterflies and moths are preyed upon by birds, bats, lizards, amphibians, dragonflies, and spiders. Caterpillars and pupae fall prey not only to birds, but also to invertebrate predators and small mammals, as well as fungi and bacteria. Parasitoid and parasitic wasps and flies may lay eggs in the caterpillar, which eventually kill it as they hatch inside its body and eat its tissues. Insect-eating birds are probably the largest predators. Lepidoptera, especially the immature stages, are an ecologically important food to many insectivorous birds, such as the great tit in Europe.
+An "evolutionary arms race" can be seen between predator and prey species. The Lepidoptera have developed a number of strategies for defense and protection, including evolution of morphological characters and changes in ecological lifestyles and behaviors. These include aposematism, mimicry, camouflage, and development of threat patterns and displays.[74] Only a few birds, such as the nightjars, hunt nocturnal lepidopterans. Their main predators are bats. Again, an "evolutionary race" exists, which has led to numerous evolutionary adaptations of moths to escape from their main predators, such as the ability to hear ultrasonic sounds, or even to emit sounds in some cases. Lepidopteran eggs are also preyed upon. Some caterpillars, such as the zebra swallowtail butterfly larvae, are cannibalistic.
+Some species of Lepidoptera are poisonous to predators, such as the monarch butterfly in the Americas, Atrophaneura species (roses, windmills, etc.) in Asia, as well as Papilio antimachus, and the birdwings, the largest butterflies in Africa and Asia, respectively. They obtain their toxicity by sequestering the chemicals from the plants they eat into their own tissues. Some Lepidoptera manufacture their own toxins. Predators that eat poisonous butterflies and moths may become sick and vomit violently, learning not to eat those species. A predator which has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future, thus saving many other species, as well.[74][75] Toxic butterflies and larvae tend to develop bright colors and striking patterns as an indicator to predators about their toxicity. This phenomenon is known as aposematism.[76] Some caterpillars, especially members of Papilionidae, contain an osmeterium, a Y-shaped protrusible gland found in the prothoracic segment of the larvae.[74] When threatened, the caterpillar emits unpleasant smells from the organ to ward off the predators.[77][78]
+Camouflage is also an important defense strategy, which involves the use of coloration or shape to blend into the surrounding environment. Some lepidopteran species blend with their surroundings, making them difficult to spot by predators. Caterpillars can exhibit shades of green that match its host plant. Others look like inedible objects, such as twigs or leaves. For instance, the mourning cloak fades into the backdrop of trees when it folds its wings back. The larvae of some species, such as the common Mormon (Papilio polytes) and the western tiger swallowtail look like bird droppings.[74][79] For example, adult Sesiidae species (also known as clearwing moths) have a general appearance sufficiently similar to a wasp or hornet to make it likely the moths gain a reduction in predation by Batesian mimicry.[80] Eyespots are a type of automimicry used by some butterflies and moths. In butterflies, the spots are composed of concentric rings of scales in different colors. The proposed role of the eyespots is to deflect attention of predators. Their resemblance to eyes provokes the predator's instinct to attack these wing patterns.[81]
+Batesian and Müllerian mimicry complexes are commonly found in Lepidoptera. Genetic polymorphism and natural selection give rise to otherwise edible species (the mimic) gaining a survival advantage by resembling inedible species (the model). Such a mimicry complex is referred to as Batesian and is most commonly known in the example between the limenitidine viceroy butterfly in relation to the inedible danaine monarch. The viceroy is, in fact, more toxic than the monarch and this resemblance should be considered as a case of Müllerian mimicry.[82] In Müllerian mimicry, inedible species, usually within a taxonomic order, find it advantageous to resemble each other so as to reduce the sampling rate by predators that need to learn about the insects' inedibility. Taxa from the toxic genus Heliconius form one of the most well-known Müllerian complexes.[83] The adults of the various species now resemble each other so well, the species cannot be distinguished without close morphological observation and, in some cases, dissection or genetic analysis.
+Moths evidently are able to hear the range emitted by bats, which in effect causes flying moths to make evasive maneuvers because bats are a main predator of moths. Ultrasonic frequencies trigger a reflex action in the noctuid moth that cause it to drop a few inches in its flight to evade attack.[84] Tiger moths in a defense emit clicks within the same range of the bats, which interfere with the bats and foil their attempts to echolocate it.[85]
+Most species of Lepidoptera engage in some form of entomophily (more specifically psychophily and phalaenophily for butterflies and moths, respectively), or the pollination of flowers.[86] Most adult butterflies and moths feed on the nectar inside flowers, using their probosces to reach the nectar hidden at the base of the petals. In the process, the adults brush against the flowers' stamens, on which the reproductive pollen is made and stored. The pollen is transferred on appendages on the adults, which fly to the next flower to feed and unwittingly deposit the pollen on the stigma of the next flower, where the pollen germinates and fertilizes the seeds.[23]:813–814
+Flowers pollinated by butterflies tend to be large and flamboyant, pink or lavender in color, frequently having a landing area, and usually scented, as butterflies are typically day-flying. Since butterflies do not digest pollen (except for heliconid species,[86]) more nectar is offered than pollen. The flowers have simple nectar guides, with the nectaries usually hidden in narrow tubes or spurs, reached by the long "tongue" of the butterflies. Butterflies such as Thymelicus flavus have been observed to engage in flower constancy, which means they are more likely to transfer pollen to other conspecific plants. This can be beneficial for the plants being pollinated, as flower constancy prevents the loss of pollen during different flights and the pollinators from clogging stigmas with pollen of other flower species.[87]
+Among the more important moth pollinator groups are the hawk moths of the family Sphingidae. Their behavior is similar to hummingbirds, i.e., using rapid wing beats to hover in front of flowers. Most hawk moths are nocturnal or crepuscular, so moth-pollinated flowers (e.g., Silene latifolia ) tend to be white, night-opening, large, and showy with tubular corollae and a strong, sweet scent produced in the evening, night, or early morning. A lot of nectar is produced to fuel the high metabolic rates needed to power their flight.[88] Other moths (e.g., noctuids, geometrids, pyralids) fly slowly and settle on the flower. They do not require as much nectar as the fast-flying hawk moths, and the flowers tend to be small (though they may be aggregated in heads).[89]
+Mutualism is a form of biological interaction wherein each individual involved benefits in some way. An example of a mutualistic relationship would be that shared by yucca moths (Tegeculidae) and their host, yucca flowers (Liliaceae). Female yucca moths enter the host flowers, collect the pollen into a ball using specialized maxillary palps, then move to the apex of the pistil, where pollen is deposited on the stigma, and lay eggs into the base of the pistil where seeds will develop. The larvae develop in the fruit pod and feed on a portion of the seeds. Thus, both insect and plant benefit, forming a highly mutualistic relationship.[23]:814 Another form of mutualism occurs between some larvae of butterflies and certain species of ants (e.g. Lycaenidae). The larvae communicate with the ants using vibrations transmitted through a substrate, such as the wood of a tree or stems, as well as using chemical signals.[90] The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions.[91]
+Only 42 species of parasitoid lepidopterans are known (1 Pyralidae; 40 Epipyropidae).[23]:748 The larvae of the greater and lesser wax moths feed on the honeycomb inside bee nests and may become pests; they are also found in bumblebee and wasp nests, albeit to a lesser extent. In northern Europe, the wax moth is regarded as the most serious parasitoid of the bumblebee, and is found only in bumblebee nests. In some areas in southern England, as much as 80% of nests can be destroyed.[92] Other parasitic larvae are known to prey upon cicadas and leaf hoppers.[93]
+In reverse, moths and butterflies may be subject to parasitic wasps and flies, which may lay eggs on the caterpillars, which hatch and feed inside its body, resulting in death. Although, in a form of parasitism called idiobiont, the adult paralyzes the host, so as not to kill it but for it to live as long as possible, for the parasitic larvae to benefit the most. In another form of parasitism, koinobiont, the species live off their hosts while inside (endoparasitic). These parasites live inside the host caterpillar throughout its life cycle, or may affect it later on as an adult. In other orders, koinobionts include flies, a majority of coleopteran, and many hymenopteran parasitoids.[23]:748–749 Some species may be subject to a variety of parasites, such as the gypsy moth (Lymantaria dispar), which is attacked by a series of 13 species, in six different taxa throughout its life cycle.[23]:750
+In response to a parasitoid egg or larva in the caterpillar's body, the plasmatocytes, or simply the host's cells can form a multilayered capsule that eventually causes the endoparasite to asphyxiate. The process, called encapsulation, is one of the caterpillar's only means of defense against parasitoids.[23]:748
+A few species of Lepidoptera are secondary consumers, or predators. These species typically prey upon the eggs of other insects, aphids, scale insects, or ant larvae.[23]:567 Some caterpillars are cannibals, and others prey on caterpillars of other species (e.g. Hawaiian Eupithecia ). Those of the 15 species in Eupithecia that mirror inchworms, are the only known species of butterflies and moths that are ambush predators.[94] Four species are known to eat snails. For example, the Hawaiian caterpillar (Hyposmocoma molluscivora) uses silk traps, in a manner similar to that of spiders, to capture certain species of snails (typically Tornatellides).[93]
+Larvae of some species of moths in the Tineidae, Gelechioidae, and Noctuidae (family/superfamily/families, respectively), besides others, feed on detritus, or dead organic material, such as fallen leaves and fruit, fungi, and animal products, and turn it into humus.[23]:567 Well-known species include the cloth moths (Tineola bisselliella, Tinea pellionella, and Trichophaga tapetzella), which feed on detritus containing keratin, including hair, feathers, cobwebs, bird nests (particularly of domestic pigeons, Columba livia domestica) and fruits or vegetables. These species are important to ecosystems as they remove substances that would otherwise take a long time to decompose.[95]
+In 2015 it was reported that wasp bracovirus DNA was present in Lepidoptera such as monarch butterflies, silkworms and moths.[96] These were described in some newspaper articles as examples of a naturally occurring genetically engineered insects.[97]
+Linnaeus in Systema Naturae (1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx and Phalaena, with seven subgroups in Phalaena.[98] These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Michael Denis & Ignaz Schiffermüller (1775), Johan Christian Fabricius (1775) and Pierre André Latreille (1796). Jacob Hübner described many genera, and the lepidopteran genera were catalogued by Ferdinand Ochsenheimer and Georg Friedrich Treitschke in a series of volumes on the lepidopteran fauna of Europe published between 1807 and 1835.[98] Gottlieb August Wilhelm Herrich-Schäffer (several volumes, 1843–1856), and Edward Meyrick (1895) based their classifications primarily on wing venation. Sir George Francis Hampson worked on the microlepidoptera during this period and Philipp Christoph Zeller published The Natural History of the Tineinae also on microlepidoptera (1855).
+Among the first entomologists to study fossil insects and their evolution was Samuel Hubbard Scudder (1837–1911), who worked on butterflies.[99] He published a study of the Florissant deposits of Colorado, including the exceptionally preserved Prodryas persephone. Andreas V. Martynov (1879–1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.[99]
+Major contributions in the 20th century included the creation of the monotrysia and ditrysia (based on female genital structure) by Borner in 1925 and 1939.[98] Willi Hennig (1913–1976) developed the cladistic methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among monotrysian families and Kristensen worked more generally on insect phylogeny and higher Lepidoptera too.[98][99] While it is often found that DNA-based phylogenies differ from those based on morphology, this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.[99]
+Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not monophyletic: Microlepidoptera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.[98]
+The fossil record for Lepidoptera is lacking in comparison to other winged species, and tends not to be as common as some other insects in habitats that are most conducive to fossilization, such as lakes and ponds; their juvenile stage has only the head capsule as a hard part that might be preserved. The location and abundance of the most common moth species are indicative that mass migrations of moths occurred over the Palaeogene North Sea, which is why there is a serious lack of moth fossils.[100] Yet there are fossils, some preserved in amber and some in very fine sediments. Leaf mines are also seen in fossil leaves, although the interpretation of them is tricky.[99]
+Putative fossil stem group representatives of Amphiesmenoptera (the clade comprising Trichoptera and Lepidoptera) are known from the Triassic.[23]:567 The earliest known lepidopteran fossils are fossilized scales from the Triassic-Jurassic boundary. They were found as rare palynological elements in the sediments of the Triassic-Jurassic boundary from the cored Schandelah-1 well, drilled near Braunschweig in northern Germany. This pushes back the fossil record and origin of glossatan lepidopterans by about 70 million years, supporting molecular estimates of a Norian (ca 212 million years) divergence of glossatan and non-glossatan lepidopterans. The findings were reported in 2018 in the journal Science Advances. The authors of the study proposed that lepidopterans evolved a proboscis as an adaptation to drink from droplets and thin films of water for maintaining their fluid balance in the hot and arid climate of the Triassic.[101]
+The earliest described lepidopteran taxon is Archaeolepis mane, a primitive moth-like species from the Jurassic, dated back to around 190 million years ago, and known only from three wings found in Dorset, UK. The wings show scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared with Trichoptera (caddisflies).[102][103] Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from the Cretaceous, which all belong to primitive moth-like families.[99]
+Many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber. The oldest genuine butterflies of the superfamily Papilionoidea have been found in the Paleocene MoClay or Fur Formation of Denmark. The best preserved fossil lepidopteran is the Eocene Prodryas persephone from the Florissant Fossil Beds.
+Lepidoptera and Trichoptera (caddisflies) are sister groups, sharing many similarities that are lacking in others; for example the females of both orders are heterogametic, meaning they have two different sex chromosomes, whereas in most species the males are heterogametic and the females have two identical sex chromosomes. The adults in both orders display a particular wing venation pattern on their forewings. The larvae in the two orders have mouth structures and glands with which they make and manipulate silk. Willi Hennig grouped the two orders into the superorder Amphiesmenoptera; together they are sister to the extinct order Tarachoptera.[104] Lepidoptera descend from a diurnal moth-like common ancestor that either fed on dead or living plants.[105]
+The cladogram, based on a 2008 DNA and protein analysis, shows the order as a clade, sister to the Trichoptera, and more distantly related to the Diptera (true flies) and Mecoptera (scorpionflies).[106][107][108][109]
+Diptera (true flies)
+Mecoptera (scorpionflies)
+Boreidae (snow scorpionflies)
+Siphonaptera (fleas)
+Trichoptera (caddisflies)
+Lepidoptera (butterflies and moths)
+Hymenoptera (sawflies, wasps, ants, bees)
+Micropterigidae, Agathiphagidae and Heterobathmiidae are the oldest and most basal lineages of Lepidoptera. The adults of these families do not have the curled tongue or proboscis, that are found in most members of the order, but instead have chewing mandibles adapted for a special diet. Micropterigidae larvae feed on leaves, fungi, or liverworts (much like the Trichoptera).[98] Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live inside kauri pines and feed on seeds. In Heterobathmiidae the larvae feed on the leaves of Nothofagus, the southern beech tree. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon after metamorphosis.[98]
+The Eriocraniidae have a short coiled proboscis in the adult stage, and though they retain their pupal mandibles with which they escaped the cocoon, their mandibles are non-functional thereafter.[98] Most of these non-ditrysian families, are primarily leaf miners in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.[99]
+With the evolution of the Ditrysia in the mid-Cretaceous, there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is one cloaca, or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, Gracillarioidea and Gelechioidea are mostly leaf miners, but more recent lineages feed externally. In the Tineoidea, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.[99]
+The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.[99] They evolved about the time that flowering plants underwent an expansive adaptive radiation in the mid-Cretaceous, and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved coevolution or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.
+In the so-called "macrolepidoptera", which constitutes about 60% of lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant.[99] Many also have tympanal organs, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.[99]
+The main lineages in the macrolepidoptera are the Noctuoidea, Bombycoidea, Lasiocampidae, Mimallonoidea, Geometroidea and Rhopalocera. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a monophyletic group.[99] The Rhopalocera, comprising the Papilionoidea (butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved.[98] There is quite a good fossil record for this group, with the oldest skipper dating from 56 million years ago.[99]
+Taxonomy is the classification of species in selected taxa, the process of naming being called nomenclature. There are over 120 families in Lepidoptera, in 45 to 48 superfamilies. Lepidoptera have always been, historically, classified in five suborders, one of which is of primitive moths that never lost the morphological features of their ancestors. The rest of the moths and butterflies make up ninety-eight percent of the other taxa, making Ditrysia. More recently, findings of new taxa, larvae and pupa have aided in detailing the relationships of primitive taxa, phylogenetic analysis showing the primitive lineages to be paraphyletic compared to the rest of Lepidoptera lineages. Recently lepidopterists have abandoned clades like suborders, and those between orders and superfamilies.[23]:569
+Artistic depictions of butterflies have been used in many cultures including as early as 3500 years ago, in Egyptian hieroglyphs.[114] Today, butterflies are widely used in various objects of art and jewelry: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and in some mixed media artworks and furnishings.[115] Butterflies have also inspired the "butterfly fairy" as an art and fictional character.
+In many cultures the soul of a dead person is associated with the butterfly, for example in Ancient Greece, where the word for butterfly ψυχή (psyche) also means soul and breath. In Latin, as in Ancient Greece, the word for "butterfly" papilio was associated with the soul of the dead.[116] The skull-like marking on the thorax of the death's-head hawkmoth has helped these moths, particularly A. atropos, earn a negative reputation, such as associations with the supernatural and evil. The moth has been prominently featured in art and movies such as Un Chien Andalou (by Buñuel and Dalí) and The Silence of the Lambs, and in the artwork of the Japanese metal band Sigh's album Hail Horror Hail. According to Kwaidan: Stories and Studies of Strange Things, by Lafcadio Hearn, a butterfly was seen in Japan as the personification of a person's soul; whether they be living, dying, or already dead. One Japanese superstition says that if a butterfly enters your guestroom and perches behind the bamboo screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad omens. When Taira no Masakado was secretly preparing for his famous revolt, there appeared in Kyoto so vast a swarm of butterflies that the people were frightened—thinking the apparition to be a portent of coming evil.[117]
+In the ancient Mesoamerican city of Teotihuacan, the brilliantly colored image of the butterfly was carved into many temples, buildings, jewelry, and emblazoned on incense burners in particular. The butterfly was sometimes depicted with the maw of a jaguar and some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies to fire and warfare persisted through to the Aztec civilization and evidence of similar jaguar-butterfly images has been found among the Zapotec, and Maya civilizations.[118]
+The larvae of many lepidopteran species are major pests in agriculture. Some of the major pests include Tortricidae, Noctuidae, and Pyralidae. The larvae of the Noctuidae genus Spodoptera (armyworms), Helicoverpa (corn earworm), or Pieris brassicae can cause extensive damage to certain crops.[98] Helicoverpa zea larvae (cotton bollworms or tomato fruitworms) are polyphagous, meaning they eat a variety of crops, including tomatoes and cotton.[119] Peridroma saucia (variegated cutworms) are described as one of the most damaging pests to gardens, with the ability to destroy entire gardens and fields in a matter of days.[120]
+Butterflies and moths are one of the largest taxa to solely feed and be dependent on living plants, in terms of the number of species, and they are in many ecosystems, making up the largest biomass to do so. In many species, the female may produce anywhere from 200 to 600 eggs, while in some others it may go as high as 30,000 eggs in one day. This can create many problems for agriculture, where many caterpillars can affect acres of vegetation. Some reports estimate that there have been over 80,000 caterpillars of several different taxa feeding on a single oak tree. In some cases, phytophagous larvae can lead to the destruction of entire trees in relatively short periods of time.[23]:567
+Ecological ways of removing pest Lepidoptera species are becoming more economically viable, as research has shown ways like introducing parasitic wasps and flies. For example, Sarcophaga aldrichi, a fly which deposited larvae feed upon the pupae of the forest tent caterpillar moth. Pesticides can affect other species other than the species they are targeted to eliminate, damaging the natural ecosystem.[121] Another good biological pest control method is the use of pheromone traps. A pheromone trap is a type of insect trap that uses pheromones to lure insects. Sex pheromones and aggregating pheromones are the most common types used. A pheromone-impregnated lure is encased in a conventional trap such as a Delta trap, water-pan trap, or funnel trap.[122]
+Species of moths that are detritivores would naturally eat detritus containing keratin, such as hairs or feathers. Well known species are cloth moths (T. bisselliella, T. pellionella, and T. tapetzella), feeding on foodstuffs that people find economically important, such as cotton, linen, silk and wool fabrics as well as furs; furthermore they have been found on shed feathers and hair, bran, semolina and flour (possibly preferring wheat flour), biscuits, casein, and insect specimens in museums.[95]
+Even though most butterflies and moths affect the economy negatively, some species are a valuable economic resource. The most prominent example is that of the domesticated silkworm moth (Bombyx mori), the larvae of which make their cocoons out of silk, which can be spun into cloth. Silk is and has been an important economic resource throughout history. The species Bombyx mori has been domesticated to the point where it is completely dependent on mankind for survival.[123] A number of wild moths such as Bombyx mandarina, and Antheraea species, besides others, provide commercially important silks.[124]
+The preference of the larvae of most lepidopteran species to feed on a single species or limited range of plants is used as a mechanism for biological control of weeds in place of herbicides. The pyralid cactus moth was introduced from Argentina to Australia, where it successfully suppressed millions of acres of prickly pear cactus.[23]:567 Another species of the Pyralidae, called the alligator weed stem borer (Arcola malloi), was used to control the aquatic plant known as alligator weed (Alternanthera philoxeroides) in conjunction with the alligator weed flea beetle; in this case, the two insects work in synergy and the weed rarely recovers.[125]
+Breeding butterflies and moths, or butterfly gardening/rearing, has become an ecologically viable process of introducing species into the ecosystem to benefit it. Butterfly ranching in Papua New Guinea permits nationals of that country to "farm" economically valuable insect species for the collectors market in an ecologically sustainable manner.[126]
+Lepidoptera feature prominently in entomophagy as food items on almost every continent. While in most cases, adults, larvae or pupae are eaten as staples by indigenous people, beondegi or silkworm pupae are eaten as a snack in Korean cuisine[127] while Maguey worm is considered a delicacy in Mexico.[128] In some parts of Huasteca, the silk nests of the Madrone butterfly are maintained on the edge of roof tops of houses for consumption.[129] In the Carnia region of Italy, children catch and eat ingluvies of the toxic Zygaena moths in early summer. The ingluvies, despite having a very low cyanogenic content, serve as a convenient, supplementary source of sugar to the children who can include this resource as a seasonal delicacy at minimum risk.[130]
+Some larvae of both moths and butterflies have a form of hair that has been known to be a cause of human health problems. Caterpillar hairs sometimes have toxins in them and species from approximately 12 families of moths or butterflies worldwide can inflict serious human injuries (urticarial dermatitis and atopic asthma to osteochondritis, consumption coagulopathy, renal failure, and intracerebral hemorrhage).[131] Skin rashes are the most common, but there have been fatalities.[132] Lonomia is a frequent cause of envenomation in humans in Brazil, with 354 cases reported between 1989 and 2005. Lethality ranging up to 20% with death caused most often by intracranial hemorrhage.[133]
+These hairs have also been known to cause keratoconjunctivitis. The sharp barbs on the end of caterpillar hairs can get lodged in soft tissues and mucous membranes such as the eyes. Once they enter such tissues, they can be difficult to extract, often exacerbating the problem as they migrate across the membrane.[134] This becomes a particular problem in an indoor setting. The hairs easily enter buildings through ventilation systems and accumulate in indoor environments because of their small size, which makes it difficult for them to be vented out. This accumulation increases the risk of human contact in indoor environments.[135]

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+Aglossata
+Glossata
+Heterobathmiina
+Zeugloptera
+Lepidoptera (/ˌlɛpɪˈdɒptərə/ LEP-i-DOP-tər-ə, from Ancient Greek lepís “scale” + pterón “wing”) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families[1] and 46 superfamilies,[2] 10 per cent of the total described species of living organisms.[2][3] It is one of the most widespread and widely recognizable insect orders in the world.[4] The Lepidoptera show many variations of the basic body structure that have evolved to gain advantages in lifestyle and distribution. Recent estimates suggest the order may have more species than earlier thought,[5] and is among the four most speciose orders, along with the Hymenoptera, Diptera, and Coleoptera.[4]
+Lepidopteran species are characterized by more than three derived features. The most apparent is the presence of scales that cover the bodies, wings, and a proboscis. The scales are modified, flattened "hairs", and give butterflies and moths their wide variety of colors and patterns. Almost all species have some form of membranous wings, except for a few that have reduced wings or are wingless. Mating and the laying of eggs are carried out by adults, normally near or on host plants for the larvae.  Like most other insects, butterflies and moths are holometabolous, meaning they undergo complete metamorphosis.  The larvae are commonly called caterpillars, and are completely different from their adult moth or butterfly forms, having a cylindrical body with a well-developed head, mandible mouth parts, three pairs of thoracic legs and from none up to five pairs of prolegs. As they grow, these larvae change in appearance, going through a series of stages called instars. Once fully matured, the larva develops into a pupa. A few butterflies and many moth species spin a silk case or cocoon prior to pupating, while others do not, instead going underground.[4] A butterfly pupa, called a chrysalis, has a hard skin, usually with no cocoon. Once the pupa has completed its metamorphosis, a sexually mature adult emerges.
+The Lepidoptera have, over millions of years, evolved a wide range of wing patterns and coloration ranging from drab moths akin to the related order Trichoptera, to the brightly colored and complex-patterned butterflies.[1] Accordingly, this is the most recognized and popular of insect orders with many people involved in the observation, study, collection, rearing of, and commerce in these insects. A person who collects or studies this order is referred to as a lepidopterist.
+Butterflies and moths play an important role in the natural ecosystem as pollinators and as food in the food chain; conversely, their larvae are considered very problematic to vegetation in agriculture, as their main source of food is often live plant matter. In many species, the female may produce from 200 to 600 eggs, while in others, the number may approach 30,000 eggs in one day. The caterpillars hatching from these eggs can cause damage to large quantities of crops. Many moth and butterfly species are of economic interest by virtue of their role as pollinators, the silk they produce, or as pest species.
+The term was coined by Linnaeus in 1735 and is derived from Greek λεπίς, gen. λεπίδος ("scale") and πτερόν ("wing").[6] Sometimes, the term Rhopalocera is used for the clade of all butterfly species, derived from the Ancient Greek ῥόπαλον (rhopalon)[7]:4150 and κέρας (keras)[7]:3993 meaning "club" and "horn", respectively, coming from the shape of the antennae of butterflies.
+The origins of the common names "butterfly" and "moth" are varied and often obscure. The English word butterfly is from Old English buttorfleoge, with many variations in spelling. Other than that, the origin is unknown, although it could be derived from the pale yellow color of many species' wings suggesting the color of butter.[8][9] The species of Heterocera are commonly called moths. The origins of the English word moth are more clear, deriving from the Old English moððe" (cf. Northumbrian dialect mohðe) from Common Germanic (compare Old Norse motti, Dutch mot and German Motte all meaning "moth"). Perhaps its origins are related to Old English maða meaning "maggot" or from the root of "midge", which until the 16th century was used mostly to indicate the larva, usually in reference to devouring clothes.[10]
+The etymological origins of the word "caterpillar", the larval form of butterflies and moths, are from the early 16th century, from Middle English catirpel, catirpeller, probably an alteration of Old North French catepelose: cate, cat (from Latin cattus) + pelose, hairy (from Latin pilōsus).[11]
+The Lepidoptera are among the most successful groups of insects. They are found on all continents, except Antarctica, and inhabit all terrestrial habitats ranging from desert to rainforest, from lowland grasslands to mountain plateaus, but almost always associated with higher plants, especially angiosperms (flowering plants).[12] Among the most northern dwelling species of butterflies and moths is the Arctic Apollo (Parnassius arcticus), which is found in the Arctic Circle in northeastern Yakutia, at an altitude of 1500 m above sea level.[13] In the Himalayas, various Apollo species such as Parnassius epaphus  have been recorded to occur up to an altitude of 6,000 m above sea level.[14]:221
+Some lepidopteran species exhibit symbiotic, phoretic, or parasitic lifestyles, inhabiting the bodies of organisms rather than the environment. Coprophagous pyralid moth species, called sloth moths, such as Bradipodicola hahneli and Cryptoses choloepi, are unusual in that they are exclusively found inhabiting the fur of sloths, mammals found in Central and South America.[15][16]
+Two species of Tinea moths have been recorded as feeding on horny tissue and have been bred from the horns of cattle. The larva of Zenodochium coccivorella is an internal parasite of the coccid Kermes species. Many species have been recorded as breeding in natural materials or refuse such as owl pellets, bat caves, honeycombs or diseased fruit.[16]
+As of 2007, there was roughly 174,250 lepidopteran species described, with butterflies and skippers estimated to comprise around 17,950, and moths making up the rest.[2][17] The vast majority of Lepidoptera are to be found in the tropics, but substantial diversity exists on most continents. North America has over 700 species of butterflies and over 11,000 species of moths,[18][19] while about 400 species of butterflies and 14,000 species of moths are reported from Australia.[20] The diversity of Lepidoptera in each faunal region has been estimated by John Heppner in 1991 based partly on actual counts from the literature, partly on the card indices in the Natural History Museum (London) and the National Museum of Natural History (Washington), and partly on estimates:[5]
+Lepidoptera are morphologically distinguished from other orders principally by the presence of scales on the external parts of the body and appendages, especially the wings. Butterflies and moths vary in size from microlepidoptera only a few millimeters long, to conspicuous animals with a wingspan greater than 25 centimetres, such as the Queen Alexandra's birdwing and Atlas moth.[21]:246
+Lepidopterans undergo a four-stage life cycle: egg; larva or caterpillar; pupa or chrysalis; and imago (plural: imagines) / adult and show many variations of the basic body structure, which give these animals advantages for diverse lifestyles and environments.
+The head is where many sensing organs and the mouth parts are found. Like the adult, the larva also has a toughened, or sclerotized head capsule.[22] Here, two compound eyes, and chaetosema, raised spots or clusters of sensory bristles unique to Lepidoptera, occur, though many taxa have lost one or both of these spots. The antennae have a wide variation in form among species and even between different sexes. The antennae of butterflies are usually filiform and shaped like clubs, those of the skippers are hooked, while those of moths have flagellar segments variously enlarged or branched. Some moths have enlarged antennae or ones that are tapered and hooked at the ends.[23]:559–560
+The maxillary galeae are modified and form an elongated proboscis. The proboscis consists of one to five segments, usually kept coiled up under the head by small muscles when it is not being used to suck up nectar from flowers or other liquids. Some basal moths still have mandibles, or separate moving jaws, like their ancestors, and these form the family Micropterigidae.[22][23]:560[24]
+The larvae, called caterpillars, have a toughened head capsule. Caterpillars lack the proboscis and have separate chewing mouthparts.[22] These mouthparts, called mandibles, are used to chew up the plant matter that the larvae eat. The lower jaw, or labium, is weak, but may carry a spinneret, an organ used to create silk. The head is made of large lateral lobes, each having an ellipse of up to six simple eyes.[23]:562–563
+The thorax is made of three fused segments, the prothorax, mesothorax, and metathorax, each with a pair of legs. The first segment contains the first pair of legs. In some males of the butterfly family Nymphalidae, the forelegs are greatly reduced and are not used for walking or perching.[23]:586 The three pairs of legs are covered with scales. Lepidoptera also have olfactory organs on their feet, which aid the butterfly in "tasting" or "smelling" out its food.[25] In the larval form there are 3 pairs of true legs, with up to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[12]
+The two pairs of wings are found on the middle and third segments, or mesothorax and metathorax, respectively. In the more recent genera, the wings of the second segment are much more pronounced, although some more primitive forms have similarly sized wings of both segments. The wings are covered in scales arranged like shingles, which form an extraordinary variety of colors and patterns. The mesothorax has more powerful muscles to propel the moth or butterfly through the air, with the wing of this segment (forewing) having a stronger vein structure.[23]:560 The largest superfamily, the Noctuidae, has their wings modified to act as tympanal or hearing organs.[26]
+The caterpillar has an elongated, soft body that may have hair-like or other projections, three pairs of true legs, with none to 11 pairs of abdominal legs (usually eight) and hooklets, called apical crochets.[12] The thorax usually has a pair of legs on each segment. The thorax is also lined with many spiracles on both the mesothorax and metathorax, except for a few aquatic species, which instead have a form of gills.[23]:563
+The abdomen, which is less sclerotized than the thorax, consists of 10 segments with membranes in between, allowing for articulated movement. The sternum, on the first segment, is small in some families and is completely absent in others. The last two or three segments form the external parts of the species' sex organs. The genitalia of Lepidoptera are highly varied and are often the only means of differentiating between species. Male genitals include a valva, which is usually large, as it is used to grasp the female during mating. Female genitalia include three distinct sections.
+The females of basal moths have only one sex organ, which is used for copulation and as an ovipositor, or egg-laying organ. About 98% of moth species have a separate organ for mating, and an external duct that carries the sperm from the male.[23]:561
+The abdomen of the caterpillar has four pairs of prolegs, normally located on the third to sixth segments of the abdomen, and a separate pair of prolegs by the anus, which have a pair of tiny hooks called crotchets. These aid in gripping and walking, especially in species that lack many prolegs (e. g. larvae of Geometridae). In some basal moths, these prolegs may be on every segment of the body, while prolegs may be completely absent in other groups, which are more adapted to boring and living in sand (e. g., Prodoxidae and Nepticulidae, respectively).[23]:563
+The wings, head, and parts of the thorax and abdomen of Lepidoptera are covered with minute scales, a feature from which the order derives its name. Most scales are lamellar, or blade-like, and attached with a pedicel, while other forms may be hair-like or specialized as secondary sexual characteristics.[27]
+The lumen or surface of the lamella has a complex structure. It gives color either by colored pigments it contains, or through structural coloration with mechanisms that include photonic crystals and diffraction gratings.[28]
+Scales function in insulation, thermoregulation, producing pheromones (in males only),[29] and aiding gliding flight, but the most important is the large diversity of vivid or indistinct patterns they provide, which help the organism protect itself by camouflage or mimicry, and which act as signals to other animals including rivals and potential mates.[27]
+In the reproductive system of butterflies and moths, the male genitalia are complex and unclear. In females the three types of genitalia are based on the relating taxa: 'monotrysian', 'exoporian', and 'ditrysian'. In the monotrysian type is an opening on the fused segments of the sterna 9 and 10, which act as insemination and oviposition. In the exoporian type (in Hepaloidae and Mnesarchaeoidea) are two separate places for insemination and oviposition, both occurring on the same sterna as the monotrysian type, i.e. 9 and 10.[21] The ditrysian groups have an internal duct that carries sperm, with separate openings for copulation and egg-laying.[4] In most species, the genitalia are flanked by two soft lobes, although they may be specialized and sclerotized in some species for ovipositing in area such as crevices and inside plant tissue.[21] Hormones and the glands that produce them run the development of butterflies and moths as they go through their life cycles, called the endocrine system. The first insect hormone prothoracicotropic hormone (PTTH) operates the species life cycle and diapause.[30] This hormone is produced by corpora allata and corpora cardiaca, where it is also stored. Some glands are specialized to perform certain task such as producing silk or producing saliva in the palpi.[31]:65, 75 While the corpora cardiaca produce PTTH, the corpora allata also produces juvenile hormones, and the prothorocic glands produce moulting hormones.
+In the digestive system, the anterior region of the foregut has been modified to form a pharyngeal  sucking pump as they need it for the food they eat, which are for the most part liquids. An esophagus follows and leads to the posterior of the pharynx and in some species forms a form of crop. The midgut is short and straight, with the hindgut being longer and coiled.[21] Ancestors of lepidopteran species, stemming from Hymenoptera, had midgut ceca, although this is lost in current butterflies and moths. Instead, all the digestive enzymes, other than initial digestion, are immobilized at the surface of the midgut cells. In larvae, long-necked and stalked goblet cells are found in the anterior and posterior midgut regions, respectively. In insects, the goblet cells excrete positive potassium ions, which are absorbed from leaves ingested by the larvae. Most butterflies and moths display the usual digestive cycle, but species with different diets require adaptations to meet these new demands.[23]:279
+In the circulatory system, hemolymph, or insect blood, is used to circulate heat in a form of thermoregulation, where muscles contraction produces heat, which is transferred to the rest of the body when conditions are unfavorable.[32]
+In lepidopteran species, hemolymph is circulated through the veins in the wings by some form of pulsating organ, either by the heart or by the intake of air into the trachea.[31]:69
+Air is taken in through spiracles along the sides of the abdomen and thorax supplying the trachea with oxygen as it goes through the lepidopteran's respiratory system. Three different tracheaes supply and diffuse oxygen throughout the species' bodies. The dorsal tracheae supply oxygen to the dorsal musculature and vessels, while the ventral tracheae supply the ventral musculature and nerve cord, and the visceral tracheae supply the guts, fat bodies, and gonads.[31]:71, 72
+Polymorphism is the appearance of forms or "morphs", which differ in color and number of attributes within a single species.[12]:163[33] In Lepidoptera, polymorphism can be seen not only between individuals in a population, but also between the sexes as sexual dimorphism, between geographically separated populations in geographical polymorphism, and between generations flying at different seasons of the year (seasonal polymorphism or polyphenism). In some species, the polymorphism is limited to one sex, typically the female. This often includes the phenomenon of mimicry when mimetic morphs fly alongside nonmimetic morphs in a population of a particular species. Polymorphism occurs both at specific level with heritable variation in the overall morphological adaptations of individuals, as well as in certain specific morphological or physiological traits within a species.[12]
+Environmental polymorphism, in which traits are not inherited, is often termed as polyphenism, which in Lepidoptera is commonly seen in the form of seasonal morphs, especially in the butterfly families of Nymphalidae and Pieridae. An Old World pierid butterfly, the common grass yellow (Eurema hecabe) has a darker summer adult morph, triggered by a long day exceeding 13 hours in duration, while the shorter diurnal period of 12 hours or less induces a paler morph in the postmonsoon period.[34] Polyphenism also occurs in caterpillars, an example being the peppered moth, Biston betularia.[35]
+Geographical isolation causes a divergence of a species into different morphs. A good example is the Indian white admiral Limenitis procris, which has five forms, each geographically separated from the other by large mountain ranges.[36]:26 An even more dramatic showcase of geographical polymorphism is the Apollo butterfly (Parnassius apollo). Because the Apollos live in small local populations, thus having no contact with each other, coupled with their strong stenotopic nature and weak migration ability, interbreeding between populations of one species practically does not occur; by this, they form over 600 different morphs, with the size of spots on the wings of which varies greatly.[37]
+Sexual dimorphism is the occurrence of differences between males and females in a species. In Lepidoptera, it is widespread and almost completely set by genetic determination.[34] Sexual dimorphism is present in all families of the Papilionoidea and more prominent in the Lycaenidae, Pieridae, and certain taxa of the Nymphalidae. Apart from color variation, which may differ from slight to completely different color-pattern combinations, secondary sexual characteristics may also be present.[36]:25 Different genotypes maintained by natural selection may also be expressed at the same time.[34] Polymorphic and/or mimetic females occur in the case of some taxa in the Papilionidae primarily to obtain a level of protection not available to the male of their species. The most distinct case of sexual dimorphism is that of adult females of many Psychidae species which have only vestigial wings, legs, and mouthparts as compared to the adult males that are strong fliers with well-developed wings and feathery antennae.[38]
+Species of Lepidoptera undergo holometabolism or "complete metamorphosis". Their life cycle normally consists of an egg, a larva, a pupa, and an imago or adult.[12] The larvae are commonly called caterpillars, and the pupae of moths encapsulated in silk are called cocoons, while the uncovered pupae of butterflies are called chrysalides.
+Unless the species reproduces year-round, a butterfly or moth may enter diapause, a state of dormancy that allows the insect to survive unfavorable environmental conditions.
+Males usually start eclosion (emergence) earlier than females and peak in numbers before females. Both of the sexes are sexually mature by the time of eclosion.[23]:564 Butterflies and moths normally do not associate with each other, except for migrating species, staying relatively asocial. Mating begins with an adult (female or male) attracting a mate, normally using visual stimuli, especially in diurnal species like most butterflies. However, the females of most nocturnal species, including almost all moth species, use pheromones to attract males, sometimes from long distances.[12] Some species engage in a form of acoustic courtship, or attract mates using sound or vibration such as the polka-dot wasp moth, Syntomeida epilais.[39]
+Adaptations include undergoing one seasonal generation, two or even more, called voltinism (Univoltism, bivoltism, and multivism, respectively). Most lepidopterans in temperate climates are univoltine, while in tropical climates most have two seasonal broods. Some others may take advantage of any opportunity they can get, and mate continuously throughout the year. These seasonal adaptations are controlled by hormones, and these delays in reproduction are called diapause.[23]:567 Many lepidopteran species, after mating and laying their eggs, die shortly afterwards, having only lived for a few days after eclosion. Others may still be active for several weeks and then overwinter and become sexually active again when the weather becomes more favorable, or diapause. The sperm of the male that mated most recently with the female is most likely to have fertilized the eggs, but the sperm from a prior mating may still prevail.[23]:564
+Lepidoptera usually reproduce sexually and are oviparous (egg-laying), though some species exhibit live birth in a process called ovoviviparity. A variety of differences in egg-laying and the number of eggs laid occur. Some species simply drop their eggs in flight (these species normally have polyphagous larvae, meaning they eat a variety of plants e. g., hepialids and some nymphalids)[40] while most lay their eggs near or on the host plant on which the larvae feed. The number of eggs laid may vary from only a few to several thousand.[12] The females of both butterflies and moths select the host plant instinctively, and primarily, by chemical cues.[23]:564
+The eggs are derived from materials ingested as a larva and in some species, from the spermatophores received from males during mating.[41] An egg can only be 1/1000 the mass of the female, yet she may lay up to her own mass in eggs. Females lay smaller eggs as they age. Larger females lay larger eggs.[42] The egg is covered by a hard-ridged protective outer layer of shell, called the chorion. It is lined with a thin coating of wax, which prevents the egg from drying out. Each egg contains a number of micropyles, or tiny funnel-shaped openings at one end, the purpose of which is to allow sperm to enter and fertilize the egg. Butterfly and moth eggs vary greatly in size between species, but they are all either spherical or ovate.
+The egg stage lasts a few weeks in most butterflies, but eggs laid prior to winter, especially in temperate regions, go through diapause, and hatching may be delayed until spring. Other butterflies may lay their eggs in the spring and have them hatch in the summer. These butterflies are usually temperate species (e. g. Nymphalis antiopa).
+The larvae or caterpillars are the first stage in the life cycle after hatching. Caterpillars, are "characteristic polypod larvae with cylindrical bodies, short thoracic legs, and abdominal prolegs (pseudopods)".[43] They have a toughened (sclerotised) head capsule with an adfrontal suture formed by medial fusion of the sclerites, mandibles (mouthparts) for chewing, and a soft tubular, segmented body, that may have hair-like or other projections, three pairs of true legs, and additional prolegs (up to five pairs).[44] The body consists of thirteen segments, of which three are thoracic and ten are abdominal.[45] Most larvae are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores.[44]
+Different herbivorous species have adapted to feed on every part of the plant and are normally considered pests to their host plants; some species have been found to lay their eggs on the fruit and other species lay their eggs on clothing or fur (e. g., Tineola bisselliella, the common clothes moth). Some species are carnivorous and others are even parasitic. Some lycaenid species such as Phengaris rebeli are social parasites of Myrmica ants nests.[46] A species of Geometridae from Hawaii has carnivorous larvae that catch and eat flies.[47] Some pyralid caterpillars are aquatic.[48]
+The larvae develop rapidly with several generations in a year; however, some species may take up to 3 years to develop, and exceptional examples like Gynaephora groenlandica take as long as seven years.[12] The larval stage is where the feeding and growing stages occur, and the larvae periodically undergo hormone-induced ecdysis, developing further with each instar, until they undergo the final larval-pupal molt.
+The larvae of both butterflies and moths exhibit mimicry to deter potential predators. Some caterpillars have the ability to inflate parts of their heads to appear snake-like. Many have false eye-spots to enhance this effect. Some caterpillars have special structures called osmeteria (family Papilionidae), which are exposed to produce smelly chemicals used in defense. Host plants often have toxic substances in them and caterpillars are able to sequester these substances and retain them into the adult stage. This helps make them unpalatable to birds and other predators. Such unpalatability is advertised using bright red, orange, black, or white warning colors. The toxic chemicals in plants are often evolved specifically to prevent them from being eaten by insects. Insects, in turn, develop countermeasures or make use of these toxins for their own survival. This "arms race" has led to the coevolution of insects and their host plants.[49]
+No form of wing is externally visible on the larva, but when larvae are dissected, developing wings can be seen as disks, which can be found on the second and third thoracic segments, in place of the spiracles that are apparent on abdominal segments. Wing disks develop in association with a trachea that runs along the base of the wing, and are surrounded by a thin peripodial membrane, which is linked to the outer epidermis of the larva by a tiny duct. Wing disks are very small until the last larval instar, when they increase dramatically in size, are invaded by branching tracheae from the wing base that precede the formation of the wing veins, and begin to develop patterns associated with several landmarks of the wing.[50]
+Near pupation, the wings are forced outside the epidermis under pressure from the hemolymph, and although they are initially quite flexible and fragile, by the time the pupa breaks free of the larval cuticle, they have adhered tightly to the outer cuticle of the pupa (in obtect pupae). Within hours, the wings form a cuticle so hard and well-joined to the body that pupae can be picked up and handled without damage to the wings.[50]
+After about five to seven instars,[51]:26–28 or molts, certain hormones, like PTTH, stimulate the production of ecdysone, which initiates insect molting. Then, the larva puparium, a sclerotized or hardened cuticle of the last larval instar, develops into the pupa. Depending on the species, the pupa may be covered in a silk cocoon, attached to different types of substrates, buried in the ground, or may not be covered at all. Features of the imago are externally recognizable in the pupa. All the appendages on the adult head and thorax are found cased inside the cuticle (antennae, mouthparts, etc.), with the wings wrapped around, adjacent to the antennae.[23]:564 The pupae of some species have functional mandibles, while the pupal mandibles are not functional in others.[22]
+While encased, some of the lower segments are not fused, and are able to move using small muscles found in between the membrane. Moving may help the pupa, for example, escape the sun, which would otherwise kill it. The pupa of the Mexican jumping bean moth (Cydia deshaisiana) does this. The larvae cut a trapdoor in the bean (species of Sebastiania) and use the bean as a shelter. With a sudden rise in temperature, the pupa inside twitches and jerks, pulling on the threads inside. Wiggling may also help to deter parasitoid wasps from laying eggs on the pupa. Other species of moths are able to make clicks to deter predators.[23]:564, 566
+The length of time before the pupa ecloses (emerges) varies greatly. The monarch butterfly may stay in its chrysalis for two weeks, while other species may need to stay for more than 10 months in diapause. The adult emerges from the pupa either by using abdominal hooks or from projections located on the head. The mandibles found in the most primitive moth families are used to escape from their cocoon (e. g., Micropterigoidea).[12][23]:564
+Most lepidopteran species do not live long after eclosion, only needing a few days to find a mate and then lay their eggs. Others may remain active for a longer period (from one to several weeks), or go through diapause and overwintering as monarch butterflies do, or waiting out environmental stress. Some adult species of microlepidoptera go through a stage where no reproductive-related activity occurs, lasting through summer and winter, followed by mating and oviposition in the early spring.[23]:564
+While most butterflies and moths are terrestrial, many species of Pyralidae are truly aquatic with all stages except the adult occurring in water. Many species from other families such as Erebidae, Nepticulidae, Cosmopterygidae, Tortricidae, Olethreutidae, Noctuidae, Cossidae, and Sphingidae are aquatic or semiaquatic.[52]:22
+Flight is an important aspect of the lives of butterflies and moths, and is used for evading predators, searching for food, and finding mates in a timely manner, as most lepidopteran species do not live long after eclosion. It is the main form of locomotion in most species. In Lepidoptera, the forewings and hindwings are mechanically coupled and flap in synchrony. Flight is anteromotoric, or being driven primarily by action of the forewings. Although lepidopteran species reportedly can still fly when their hindwings are cut off, it reduces their linear flight and turning capabilities.[53]
+Lepidopteran species have to be warm, about 77 to 79 °F (25 to 26 °C), to fly. They depend on their body temperature being sufficiently high and since they cannot regulate it themselves, this is dependent on their environment. Butterflies living in cooler climates may use their wings to warm their bodies. They will bask in the sun, spreading out their wings so that they get maximum exposure to the sunlight. In hotter climates butterflies can easily overheat, so they are usually active only during the cooler parts of the day, early morning, late afternoon or early evening. During the heat of the day, they rest in the shade. Some larger thick-bodied moths (e.g. Sphingidae) can generate their own heat to a limited degree by vibrating their wings. The heat generated by the flight muscles warms the thorax while the temperature of the abdomen is unimportant for flight. To avoid overheating, some moths rely on hairy scales, internal air sacs, and other structures to separate the thorax and abdomen and keep the abdomen cooler.[54]
+Some species of butterflies can reach fast speeds, such as the southern dart, which can go as fast as 48.4 km/h. Sphingids are some of the fastest flying insects, some are capable of flying at over 50 km/h (30 mi/h), having a wingspan of 35–150 mm.[1][55] In some species, sometimes a gliding component to their flight exists. Flight occurs either as hovering, or as forward or backward motion.[56] In butterfly and moth species, such as hawk moths, hovering is important as they need to maintain a certain stability over flowers when feeding on the nectar.[1]
+Navigation is important to Lepidoptera species, especially for those that migrate. Butterflies, which have more species that migrate, have been shown to navigate using time-compensated sun compasses. They can see polarized light, so can orient even in cloudy conditions. The polarized light in the region close to the ultraviolet spectrum is suggested to be particularly important.[57] Most migratory butterflies are those that live in semiarid areas where breeding seasons are short.[58] The life histories of their host plants also influence the strategies of the butterflies.[59] Other theories include the use of landscapes. Lepidoptera may use coastal lines, mountains, and even roads to orient themselves. Above sea, the flight direction is much more accurate if the coast is still visible.[60]
+Many studies have also shown that moths navigate. One study showed that many moths may use the Earth's magnetic field to navigate, as a study of the moth heart and dart suggests.[61] Another study, of the migratory behavior of the silver Y, showed, even at high altitudes, the species can correct its course with changing winds, and prefers flying with favourable winds, suggesting a great sense of direction.[62][63] Aphrissa statira in Panama loses its navigational capacity when exposed to a magnetic field, suggesting it uses the Earth's magnetic field.[64]
+Moths exhibit a tendency to circle artificial lights repeatedly. This suggests they use a technique of celestial navigation called transverse orientation. By maintaining a constant angular relationship to a bright celestial light, such as the Moon, they can fly in a straight line. Celestial objects are so far away, even after traveling great distances, the change in angle between the moth and the light source is negligible; further, the moon will always be in the upper part of the visual field or on the horizon. When a moth encounters a much closer artificial light and uses it for navigation, the angle changes noticeably after only a short distance, in addition to being often below the horizon. The moth instinctively attempts to correct by turning toward the light, causing airborne moths to come plummeting downwards, and at close range, which results in a spiral flight path that gets closer and closer to the light source.[65] Other explanations have been suggested, such as the idea that moths may be impaired with a visual distortion called a Mach band by Henry Hsiao in 1972. He stated that they fly towards the darkest part of the sky in pursuit of safety, thus are inclined to circle ambient objects in the Mach band region.[66]
+Lepidopteran migration is typically seasonal, as the insects moving to escape dry seasons or other disadvantageous conditions. Most lepidopterans that migrate are butterflies, and the distance travelled varies. Some butterflies that migrate include the mourning cloak, painted lady, American lady, red admiral, and the common buckeye.[51]:29–30 A notable species of moth that migrates long distances is the bogong moth.[67] The most well-known migrations are those of the eastern population of the monarch butterfly from Mexico to northern United States and southern Canada, a distance of about 4,000–4,800 km (2,500–3,000 mi). Other well-known migratory species include the painted lady and several of the danaine butterflies. Spectacular and large-scale migrations associated with the monsoons are seen in peninsular India.[68] Migrations have been studied in more recent times using wing tags and stable hydrogen isotopes.[69][70]
+Moths also undertake migrations, an example being the uraniids. Urania fulgens undergoes population explosions and massive migrations that may be not surpassed by any other insect in the Neotropics. In Costa Rica and Panama, the first population movements may begin in July and early August and depending on the year, may be very massive, continuing unabated for as long as five months.[71]
+Pheromones are commonly involved in mating rituals among species, especially moths, but they are also an important aspect of other forms of communication. Usually, the pheromones are produced by either the male or the female and detected by members of the opposite sex with their antennae.[72] In many species, a gland between the eighth and ninth segments under the abdomen in the female produces the pheromones.[12] Communication can also occur through stridulation, or producing sounds by rubbing various parts of the body together.[63]
+Moths are known to engage in acoustic forms of communication, most often as courtship, attracting mates using sound or vibration. Like most other insects, moths pick up these sounds using tympanic membranes in their abdomens.[73] An example is that of the polka-dot wasp moth (Syntomeida epilais), which produces sounds with a frequency above that normally detectable by humans (about 20 kHz). These sounds also function as tactile communication, or communication through touch, as they stridulate, or vibrate a substrate like leaves and stems.[39]
+Most moths lack bright colors, as many species use coloration as camouflage, but butterflies engage in visual communication. Female cabbage butterflies, for example, use ultraviolet light to communicate, with scales colored in this range on the dorsal wing surface. When they fly, each down stroke of the wing creates a brief flash of ultraviolet light which the males apparently recognize as the flight signature of a potential mate. These flashes from the wings may attract several males that engage in aerial courtship displays.[73]
+Moths and butterflies are important in the natural ecosystem. They are integral participants in the food chain; having co-evolved with flowering plants and predators, lepidopteran species have formed a network of trophic relationships between autotrophs and heterotrophs, which are included in the stages of Lepidoptera larvae, pupae, and adults. Larvae and pupae are links in the diets of birds and parasitic entomophagous insects. The adults are included in food webs in a much broader range of consumers (including birds, small mammals, reptiles, etc.).[23]:567
+Lepidopteran species are soft bodied, fragile, and almost defenseless, while the immature stages move slowly or are immobile, hence all stages are exposed to predation. Adult butterflies and moths are preyed upon by birds, bats, lizards, amphibians, dragonflies, and spiders. Caterpillars and pupae fall prey not only to birds, but also to invertebrate predators and small mammals, as well as fungi and bacteria. Parasitoid and parasitic wasps and flies may lay eggs in the caterpillar, which eventually kill it as they hatch inside its body and eat its tissues. Insect-eating birds are probably the largest predators. Lepidoptera, especially the immature stages, are an ecologically important food to many insectivorous birds, such as the great tit in Europe.
+An "evolutionary arms race" can be seen between predator and prey species. The Lepidoptera have developed a number of strategies for defense and protection, including evolution of morphological characters and changes in ecological lifestyles and behaviors. These include aposematism, mimicry, camouflage, and development of threat patterns and displays.[74] Only a few birds, such as the nightjars, hunt nocturnal lepidopterans. Their main predators are bats. Again, an "evolutionary race" exists, which has led to numerous evolutionary adaptations of moths to escape from their main predators, such as the ability to hear ultrasonic sounds, or even to emit sounds in some cases. Lepidopteran eggs are also preyed upon. Some caterpillars, such as the zebra swallowtail butterfly larvae, are cannibalistic.
+Some species of Lepidoptera are poisonous to predators, such as the monarch butterfly in the Americas, Atrophaneura species (roses, windmills, etc.) in Asia, as well as Papilio antimachus, and the birdwings, the largest butterflies in Africa and Asia, respectively. They obtain their toxicity by sequestering the chemicals from the plants they eat into their own tissues. Some Lepidoptera manufacture their own toxins. Predators that eat poisonous butterflies and moths may become sick and vomit violently, learning not to eat those species. A predator which has previously eaten a poisonous lepidopteran may avoid other species with similar markings in the future, thus saving many other species, as well.[74][75] Toxic butterflies and larvae tend to develop bright colors and striking patterns as an indicator to predators about their toxicity. This phenomenon is known as aposematism.[76] Some caterpillars, especially members of Papilionidae, contain an osmeterium, a Y-shaped protrusible gland found in the prothoracic segment of the larvae.[74] When threatened, the caterpillar emits unpleasant smells from the organ to ward off the predators.[77][78]
+Camouflage is also an important defense strategy, which involves the use of coloration or shape to blend into the surrounding environment. Some lepidopteran species blend with their surroundings, making them difficult to spot by predators. Caterpillars can exhibit shades of green that match its host plant. Others look like inedible objects, such as twigs or leaves. For instance, the mourning cloak fades into the backdrop of trees when it folds its wings back. The larvae of some species, such as the common Mormon (Papilio polytes) and the western tiger swallowtail look like bird droppings.[74][79] For example, adult Sesiidae species (also known as clearwing moths) have a general appearance sufficiently similar to a wasp or hornet to make it likely the moths gain a reduction in predation by Batesian mimicry.[80] Eyespots are a type of automimicry used by some butterflies and moths. In butterflies, the spots are composed of concentric rings of scales in different colors. The proposed role of the eyespots is to deflect attention of predators. Their resemblance to eyes provokes the predator's instinct to attack these wing patterns.[81]
+Batesian and Müllerian mimicry complexes are commonly found in Lepidoptera. Genetic polymorphism and natural selection give rise to otherwise edible species (the mimic) gaining a survival advantage by resembling inedible species (the model). Such a mimicry complex is referred to as Batesian and is most commonly known in the example between the limenitidine viceroy butterfly in relation to the inedible danaine monarch. The viceroy is, in fact, more toxic than the monarch and this resemblance should be considered as a case of Müllerian mimicry.[82] In Müllerian mimicry, inedible species, usually within a taxonomic order, find it advantageous to resemble each other so as to reduce the sampling rate by predators that need to learn about the insects' inedibility. Taxa from the toxic genus Heliconius form one of the most well-known Müllerian complexes.[83] The adults of the various species now resemble each other so well, the species cannot be distinguished without close morphological observation and, in some cases, dissection or genetic analysis.
+Moths evidently are able to hear the range emitted by bats, which in effect causes flying moths to make evasive maneuvers because bats are a main predator of moths. Ultrasonic frequencies trigger a reflex action in the noctuid moth that cause it to drop a few inches in its flight to evade attack.[84] Tiger moths in a defense emit clicks within the same range of the bats, which interfere with the bats and foil their attempts to echolocate it.[85]
+Most species of Lepidoptera engage in some form of entomophily (more specifically psychophily and phalaenophily for butterflies and moths, respectively), or the pollination of flowers.[86] Most adult butterflies and moths feed on the nectar inside flowers, using their probosces to reach the nectar hidden at the base of the petals. In the process, the adults brush against the flowers' stamens, on which the reproductive pollen is made and stored. The pollen is transferred on appendages on the adults, which fly to the next flower to feed and unwittingly deposit the pollen on the stigma of the next flower, where the pollen germinates and fertilizes the seeds.[23]:813–814
+Flowers pollinated by butterflies tend to be large and flamboyant, pink or lavender in color, frequently having a landing area, and usually scented, as butterflies are typically day-flying. Since butterflies do not digest pollen (except for heliconid species,[86]) more nectar is offered than pollen. The flowers have simple nectar guides, with the nectaries usually hidden in narrow tubes or spurs, reached by the long "tongue" of the butterflies. Butterflies such as Thymelicus flavus have been observed to engage in flower constancy, which means they are more likely to transfer pollen to other conspecific plants. This can be beneficial for the plants being pollinated, as flower constancy prevents the loss of pollen during different flights and the pollinators from clogging stigmas with pollen of other flower species.[87]
+Among the more important moth pollinator groups are the hawk moths of the family Sphingidae. Their behavior is similar to hummingbirds, i.e., using rapid wing beats to hover in front of flowers. Most hawk moths are nocturnal or crepuscular, so moth-pollinated flowers (e.g., Silene latifolia ) tend to be white, night-opening, large, and showy with tubular corollae and a strong, sweet scent produced in the evening, night, or early morning. A lot of nectar is produced to fuel the high metabolic rates needed to power their flight.[88] Other moths (e.g., noctuids, geometrids, pyralids) fly slowly and settle on the flower. They do not require as much nectar as the fast-flying hawk moths, and the flowers tend to be small (though they may be aggregated in heads).[89]
+Mutualism is a form of biological interaction wherein each individual involved benefits in some way. An example of a mutualistic relationship would be that shared by yucca moths (Tegeculidae) and their host, yucca flowers (Liliaceae). Female yucca moths enter the host flowers, collect the pollen into a ball using specialized maxillary palps, then move to the apex of the pistil, where pollen is deposited on the stigma, and lay eggs into the base of the pistil where seeds will develop. The larvae develop in the fruit pod and feed on a portion of the seeds. Thus, both insect and plant benefit, forming a highly mutualistic relationship.[23]:814 Another form of mutualism occurs between some larvae of butterflies and certain species of ants (e.g. Lycaenidae). The larvae communicate with the ants using vibrations transmitted through a substrate, such as the wood of a tree or stems, as well as using chemical signals.[90] The ants provide some degree of protection to these larvae and they in turn gather honeydew secretions.[91]
+Only 42 species of parasitoid lepidopterans are known (1 Pyralidae; 40 Epipyropidae).[23]:748 The larvae of the greater and lesser wax moths feed on the honeycomb inside bee nests and may become pests; they are also found in bumblebee and wasp nests, albeit to a lesser extent. In northern Europe, the wax moth is regarded as the most serious parasitoid of the bumblebee, and is found only in bumblebee nests. In some areas in southern England, as much as 80% of nests can be destroyed.[92] Other parasitic larvae are known to prey upon cicadas and leaf hoppers.[93]
+In reverse, moths and butterflies may be subject to parasitic wasps and flies, which may lay eggs on the caterpillars, which hatch and feed inside its body, resulting in death. Although, in a form of parasitism called idiobiont, the adult paralyzes the host, so as not to kill it but for it to live as long as possible, for the parasitic larvae to benefit the most. In another form of parasitism, koinobiont, the species live off their hosts while inside (endoparasitic). These parasites live inside the host caterpillar throughout its life cycle, or may affect it later on as an adult. In other orders, koinobionts include flies, a majority of coleopteran, and many hymenopteran parasitoids.[23]:748–749 Some species may be subject to a variety of parasites, such as the gypsy moth (Lymantaria dispar), which is attacked by a series of 13 species, in six different taxa throughout its life cycle.[23]:750
+In response to a parasitoid egg or larva in the caterpillar's body, the plasmatocytes, or simply the host's cells can form a multilayered capsule that eventually causes the endoparasite to asphyxiate. The process, called encapsulation, is one of the caterpillar's only means of defense against parasitoids.[23]:748
+A few species of Lepidoptera are secondary consumers, or predators. These species typically prey upon the eggs of other insects, aphids, scale insects, or ant larvae.[23]:567 Some caterpillars are cannibals, and others prey on caterpillars of other species (e.g. Hawaiian Eupithecia ). Those of the 15 species in Eupithecia that mirror inchworms, are the only known species of butterflies and moths that are ambush predators.[94] Four species are known to eat snails. For example, the Hawaiian caterpillar (Hyposmocoma molluscivora) uses silk traps, in a manner similar to that of spiders, to capture certain species of snails (typically Tornatellides).[93]
+Larvae of some species of moths in the Tineidae, Gelechioidae, and Noctuidae (family/superfamily/families, respectively), besides others, feed on detritus, or dead organic material, such as fallen leaves and fruit, fungi, and animal products, and turn it into humus.[23]:567 Well-known species include the cloth moths (Tineola bisselliella, Tinea pellionella, and Trichophaga tapetzella), which feed on detritus containing keratin, including hair, feathers, cobwebs, bird nests (particularly of domestic pigeons, Columba livia domestica) and fruits or vegetables. These species are important to ecosystems as they remove substances that would otherwise take a long time to decompose.[95]
+In 2015 it was reported that wasp bracovirus DNA was present in Lepidoptera such as monarch butterflies, silkworms and moths.[96] These were described in some newspaper articles as examples of a naturally occurring genetically engineered insects.[97]
+Linnaeus in Systema Naturae (1758) recognized three divisions of the Lepidoptera: Papilio, Sphinx and Phalaena, with seven subgroups in Phalaena.[98] These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Michael Denis & Ignaz Schiffermüller (1775), Johan Christian Fabricius (1775) and Pierre André Latreille (1796). Jacob Hübner described many genera, and the lepidopteran genera were catalogued by Ferdinand Ochsenheimer and Georg Friedrich Treitschke in a series of volumes on the lepidopteran fauna of Europe published between 1807 and 1835.[98] Gottlieb August Wilhelm Herrich-Schäffer (several volumes, 1843–1856), and Edward Meyrick (1895) based their classifications primarily on wing venation. Sir George Francis Hampson worked on the microlepidoptera during this period and Philipp Christoph Zeller published The Natural History of the Tineinae also on microlepidoptera (1855).
+Among the first entomologists to study fossil insects and their evolution was Samuel Hubbard Scudder (1837–1911), who worked on butterflies.[99] He published a study of the Florissant deposits of Colorado, including the exceptionally preserved Prodryas persephone. Andreas V. Martynov (1879–1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.[99]
+Major contributions in the 20th century included the creation of the monotrysia and ditrysia (based on female genital structure) by Borner in 1925 and 1939.[98] Willi Hennig (1913–1976) developed the cladistic methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among monotrysian families and Kristensen worked more generally on insect phylogeny and higher Lepidoptera too.[98][99] While it is often found that DNA-based phylogenies differ from those based on morphology, this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.[99]
+Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not monophyletic: Microlepidoptera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.[98]
+The fossil record for Lepidoptera is lacking in comparison to other winged species, and tends not to be as common as some other insects in habitats that are most conducive to fossilization, such as lakes and ponds; their juvenile stage has only the head capsule as a hard part that might be preserved. The location and abundance of the most common moth species are indicative that mass migrations of moths occurred over the Palaeogene North Sea, which is why there is a serious lack of moth fossils.[100] Yet there are fossils, some preserved in amber and some in very fine sediments. Leaf mines are also seen in fossil leaves, although the interpretation of them is tricky.[99]
+Putative fossil stem group representatives of Amphiesmenoptera (the clade comprising Trichoptera and Lepidoptera) are known from the Triassic.[23]:567 The earliest known lepidopteran fossils are fossilized scales from the Triassic-Jurassic boundary. They were found as rare palynological elements in the sediments of the Triassic-Jurassic boundary from the cored Schandelah-1 well, drilled near Braunschweig in northern Germany. This pushes back the fossil record and origin of glossatan lepidopterans by about 70 million years, supporting molecular estimates of a Norian (ca 212 million years) divergence of glossatan and non-glossatan lepidopterans. The findings were reported in 2018 in the journal Science Advances. The authors of the study proposed that lepidopterans evolved a proboscis as an adaptation to drink from droplets and thin films of water for maintaining their fluid balance in the hot and arid climate of the Triassic.[101]
+The earliest described lepidopteran taxon is Archaeolepis mane, a primitive moth-like species from the Jurassic, dated back to around 190 million years ago, and known only from three wings found in Dorset, UK. The wings show scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared with Trichoptera (caddisflies).[102][103] Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from the Cretaceous, which all belong to primitive moth-like families.[99]
+Many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber. The oldest genuine butterflies of the superfamily Papilionoidea have been found in the Paleocene MoClay or Fur Formation of Denmark. The best preserved fossil lepidopteran is the Eocene Prodryas persephone from the Florissant Fossil Beds.
+Lepidoptera and Trichoptera (caddisflies) are sister groups, sharing many similarities that are lacking in others; for example the females of both orders are heterogametic, meaning they have two different sex chromosomes, whereas in most species the males are heterogametic and the females have two identical sex chromosomes. The adults in both orders display a particular wing venation pattern on their forewings. The larvae in the two orders have mouth structures and glands with which they make and manipulate silk. Willi Hennig grouped the two orders into the superorder Amphiesmenoptera; together they are sister to the extinct order Tarachoptera.[104] Lepidoptera descend from a diurnal moth-like common ancestor that either fed on dead or living plants.[105]
+The cladogram, based on a 2008 DNA and protein analysis, shows the order as a clade, sister to the Trichoptera, and more distantly related to the Diptera (true flies) and Mecoptera (scorpionflies).[106][107][108][109]
+Diptera (true flies)
+Mecoptera (scorpionflies)
+Boreidae (snow scorpionflies)
+Siphonaptera (fleas)
+Trichoptera (caddisflies)
+Lepidoptera (butterflies and moths)
+Hymenoptera (sawflies, wasps, ants, bees)
+Micropterigidae, Agathiphagidae and Heterobathmiidae are the oldest and most basal lineages of Lepidoptera. The adults of these families do not have the curled tongue or proboscis, that are found in most members of the order, but instead have chewing mandibles adapted for a special diet. Micropterigidae larvae feed on leaves, fungi, or liverworts (much like the Trichoptera).[98] Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live inside kauri pines and feed on seeds. In Heterobathmiidae the larvae feed on the leaves of Nothofagus, the southern beech tree. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon after metamorphosis.[98]
+The Eriocraniidae have a short coiled proboscis in the adult stage, and though they retain their pupal mandibles with which they escaped the cocoon, their mandibles are non-functional thereafter.[98] Most of these non-ditrysian families, are primarily leaf miners in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.[99]
+With the evolution of the Ditrysia in the mid-Cretaceous, there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is one cloaca, or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, Gracillarioidea and Gelechioidea are mostly leaf miners, but more recent lineages feed externally. In the Tineoidea, most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.[99]
+The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.[99] They evolved about the time that flowering plants underwent an expansive adaptive radiation in the mid-Cretaceous, and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved coevolution or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.
+In the so-called "macrolepidoptera", which constitutes about 60% of lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant.[99] Many also have tympanal organs, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.[99]
+The main lineages in the macrolepidoptera are the Noctuoidea, Bombycoidea, Lasiocampidae, Mimallonoidea, Geometroidea and Rhopalocera. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a monophyletic group.[99] The Rhopalocera, comprising the Papilionoidea (butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved.[98] There is quite a good fossil record for this group, with the oldest skipper dating from 56 million years ago.[99]
+Taxonomy is the classification of species in selected taxa, the process of naming being called nomenclature. There are over 120 families in Lepidoptera, in 45 to 48 superfamilies. Lepidoptera have always been, historically, classified in five suborders, one of which is of primitive moths that never lost the morphological features of their ancestors. The rest of the moths and butterflies make up ninety-eight percent of the other taxa, making Ditrysia. More recently, findings of new taxa, larvae and pupa have aided in detailing the relationships of primitive taxa, phylogenetic analysis showing the primitive lineages to be paraphyletic compared to the rest of Lepidoptera lineages. Recently lepidopterists have abandoned clades like suborders, and those between orders and superfamilies.[23]:569
+Artistic depictions of butterflies have been used in many cultures including as early as 3500 years ago, in Egyptian hieroglyphs.[114] Today, butterflies are widely used in various objects of art and jewelry: mounted in frames, embedded in resin, displayed in bottles, laminated in paper, and in some mixed media artworks and furnishings.[115] Butterflies have also inspired the "butterfly fairy" as an art and fictional character.
+In many cultures the soul of a dead person is associated with the butterfly, for example in Ancient Greece, where the word for butterfly ψυχή (psyche) also means soul and breath. In Latin, as in Ancient Greece, the word for "butterfly" papilio was associated with the soul of the dead.[116] The skull-like marking on the thorax of the death's-head hawkmoth has helped these moths, particularly A. atropos, earn a negative reputation, such as associations with the supernatural and evil. The moth has been prominently featured in art and movies such as Un Chien Andalou (by Buñuel and Dalí) and The Silence of the Lambs, and in the artwork of the Japanese metal band Sigh's album Hail Horror Hail. According to Kwaidan: Stories and Studies of Strange Things, by Lafcadio Hearn, a butterfly was seen in Japan as the personification of a person's soul; whether they be living, dying, or already dead. One Japanese superstition says that if a butterfly enters your guestroom and perches behind the bamboo screen, the person whom you most love is coming to see you. However, large numbers of butterflies are viewed as bad omens. When Taira no Masakado was secretly preparing for his famous revolt, there appeared in Kyoto so vast a swarm of butterflies that the people were frightened—thinking the apparition to be a portent of coming evil.[117]
+In the ancient Mesoamerican city of Teotihuacan, the brilliantly colored image of the butterfly was carved into many temples, buildings, jewelry, and emblazoned on incense burners in particular. The butterfly was sometimes depicted with the maw of a jaguar and some species were considered to be the reincarnations of the souls of dead warriors. The close association of butterflies to fire and warfare persisted through to the Aztec civilization and evidence of similar jaguar-butterfly images has been found among the Zapotec, and Maya civilizations.[118]
+The larvae of many lepidopteran species are major pests in agriculture. Some of the major pests include Tortricidae, Noctuidae, and Pyralidae. The larvae of the Noctuidae genus Spodoptera (armyworms), Helicoverpa (corn earworm), or Pieris brassicae can cause extensive damage to certain crops.[98] Helicoverpa zea larvae (cotton bollworms or tomato fruitworms) are polyphagous, meaning they eat a variety of crops, including tomatoes and cotton.[119] Peridroma saucia (variegated cutworms) are described as one of the most damaging pests to gardens, with the ability to destroy entire gardens and fields in a matter of days.[120]
+Butterflies and moths are one of the largest taxa to solely feed and be dependent on living plants, in terms of the number of species, and they are in many ecosystems, making up the largest biomass to do so. In many species, the female may produce anywhere from 200 to 600 eggs, while in some others it may go as high as 30,000 eggs in one day. This can create many problems for agriculture, where many caterpillars can affect acres of vegetation. Some reports estimate that there have been over 80,000 caterpillars of several different taxa feeding on a single oak tree. In some cases, phytophagous larvae can lead to the destruction of entire trees in relatively short periods of time.[23]:567
+Ecological ways of removing pest Lepidoptera species are becoming more economically viable, as research has shown ways like introducing parasitic wasps and flies. For example, Sarcophaga aldrichi, a fly which deposited larvae feed upon the pupae of the forest tent caterpillar moth. Pesticides can affect other species other than the species they are targeted to eliminate, damaging the natural ecosystem.[121] Another good biological pest control method is the use of pheromone traps. A pheromone trap is a type of insect trap that uses pheromones to lure insects. Sex pheromones and aggregating pheromones are the most common types used. A pheromone-impregnated lure is encased in a conventional trap such as a Delta trap, water-pan trap, or funnel trap.[122]
+Species of moths that are detritivores would naturally eat detritus containing keratin, such as hairs or feathers. Well known species are cloth moths (T. bisselliella, T. pellionella, and T. tapetzella), feeding on foodstuffs that people find economically important, such as cotton, linen, silk and wool fabrics as well as furs; furthermore they have been found on shed feathers and hair, bran, semolina and flour (possibly preferring wheat flour), biscuits, casein, and insect specimens in museums.[95]
+Even though most butterflies and moths affect the economy negatively, some species are a valuable economic resource. The most prominent example is that of the domesticated silkworm moth (Bombyx mori), the larvae of which make their cocoons out of silk, which can be spun into cloth. Silk is and has been an important economic resource throughout history. The species Bombyx mori has been domesticated to the point where it is completely dependent on mankind for survival.[123] A number of wild moths such as Bombyx mandarina, and Antheraea species, besides others, provide commercially important silks.[124]
+The preference of the larvae of most lepidopteran species to feed on a single species or limited range of plants is used as a mechanism for biological control of weeds in place of herbicides. The pyralid cactus moth was introduced from Argentina to Australia, where it successfully suppressed millions of acres of prickly pear cactus.[23]:567 Another species of the Pyralidae, called the alligator weed stem borer (Arcola malloi), was used to control the aquatic plant known as alligator weed (Alternanthera philoxeroides) in conjunction with the alligator weed flea beetle; in this case, the two insects work in synergy and the weed rarely recovers.[125]
+Breeding butterflies and moths, or butterfly gardening/rearing, has become an ecologically viable process of introducing species into the ecosystem to benefit it. Butterfly ranching in Papua New Guinea permits nationals of that country to "farm" economically valuable insect species for the collectors market in an ecologically sustainable manner.[126]
+Lepidoptera feature prominently in entomophagy as food items on almost every continent. While in most cases, adults, larvae or pupae are eaten as staples by indigenous people, beondegi or silkworm pupae are eaten as a snack in Korean cuisine[127] while Maguey worm is considered a delicacy in Mexico.[128] In some parts of Huasteca, the silk nests of the Madrone butterfly are maintained on the edge of roof tops of houses for consumption.[129] In the Carnia region of Italy, children catch and eat ingluvies of the toxic Zygaena moths in early summer. The ingluvies, despite having a very low cyanogenic content, serve as a convenient, supplementary source of sugar to the children who can include this resource as a seasonal delicacy at minimum risk.[130]
+Some larvae of both moths and butterflies have a form of hair that has been known to be a cause of human health problems. Caterpillar hairs sometimes have toxins in them and species from approximately 12 families of moths or butterflies worldwide can inflict serious human injuries (urticarial dermatitis and atopic asthma to osteochondritis, consumption coagulopathy, renal failure, and intracerebral hemorrhage).[131] Skin rashes are the most common, but there have been fatalities.[132] Lonomia is a frequent cause of envenomation in humans in Brazil, with 354 cases reported between 1989 and 2005. Lethality ranging up to 20% with death caused most often by intracranial hemorrhage.[133]
+These hairs have also been known to cause keratoconjunctivitis. The sharp barbs on the end of caterpillar hairs can get lodged in soft tissues and mucous membranes such as the eyes. Once they enter such tissues, they can be difficult to extract, often exacerbating the problem as they migrate across the membrane.[134] This becomes a particular problem in an indoor setting. The hairs easily enter buildings through ventilation systems and accumulate in indoor environments because of their small size, which makes it difficult for them to be vented out. This accumulation increases the risk of human contact in indoor environments.[135]

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+The serval (Leptailurus serval) /ˈsɜːrvəl/ is a wild cat native to Africa. It is rare in North Africa and the Sahel, but widespread in sub-Saharan countries except rainforest regions. On the IUCN Red List it is listed as Least Concern.[1]
+It was first described by von Schreber in 1776.[3] It is the sole member of the genus Leptailurus. Three subspecies are recognised. The serval is a slender, medium-sized cat that stands 54–62 cm (21–24 in) at the shoulder and weighs 9–18 kg (20–40 lb). It is characterised by a small head, large ears, a golden-yellow to buff coat spotted and striped with black, and a short, black-tipped tail. The serval has the longest legs of any cat relative to its body size.
+Active in the day as well as at night, servals tend to be solitary with minimal social interaction. Both sexes establish highly overlapping home ranges of 10 to 32 km2 (4 to 12 sq mi), and mark them with feces and saliva. Servals are carnivores – they prey on rodents (particularly vlei rats), small birds, frogs, insects, and reptiles. The serval uses its sense of hearing to locate the prey; to kill small prey, it leaps over 2 m (6 ft 7 in) above the ground to land on the prey on its forefeet, and finally kills it with a bite on the neck or the head. Mating takes place at different times of the year in different parts of their range, but typically once or twice a year in an area. After a gestational period of two to three months, a litter of one to four is born. Weaning occurs at one month, and kittens begin hunting on their own at six months. The juveniles leave their mother at 12 months.
+It occurs in protected areas across its range, and hunting of servals is either prohibited or regulated in several countries.
+The name "serval" is a Portuguese name used by Georges-Louis Leclerc, Comte de Buffon in 1765 for a spotted cat that was kept at the time in the Royal Menagerie in Versailles.[4]
+The name Leptailurus may have been derived from the medieval Greek λεπταλέος or λεπτός meaning "fine, delicate".[5]
+Felis serval was first described by Johann Christian Daniel von Schreber in 1776.[3] In the 19th and 20th centuries, the following serval zoological specimens were described:
+The generic name Leptailurus was proposed by Nikolai Severtzov in 1858.[12] The serval is the sole member of this genus.[13]
+In 1944, Pocock recognised three serval races in North Africa.[14]
+Three subspecies are recognised as valid since 2017:[15]
+The phylogenetic relationships of the serval have remained in dispute; in 1997, palaeontologists M. C. McKenna and S. K. Bell classified Leptailurus as a subgenus of Felis, while others like O. R. P. Bininda-Edmonds (of the Technical University of Munich) have grouped it with Felis, Lynx and Caracal. Studies in the 2000s and the 2010s show that the serval, along with the caracal and the African golden cat, forms one of the eight lineages of Felidae. According to a 2006 genetic study, the Caracal lineage came into existence 8.5 million years ago, and the ancestor of this lineage arrived in Africa 8.5–5.6 mya.[2][16]
+The phylogenetic relationships of the serval are as follows:[2][16]
+Marbled cat (P. marmorata)
+Bay cat (Catopuma badia)
+Asian golden cat (Catopuma temminckii)
+Serval (L. serval)
+Caracal (Caracal caracal)
+African golden cat (Caracal aurata)
+Leopardus
+Lynx
+Acinonyx
+Puma
+Otocolobus
+Prionailurus
+Felis
+In April 1986, the first savannah cat, a hybrid between a male serval and a female domestic cat, was born; it was larger than a typical domestic kitten and resembled its father in its coat pattern. It appeared to have inherited a few domestic cat traits, such as tameness, from its mother. This cat breed may have a dog-like habit of following its owner about, and can be a good swimmer. Over the years it has gained popularity as a pet.[17]
+The serval is a slender, medium-sized cat; it stands 54 to 62 cm (21–24 in) at the shoulder and weighs 8 to 18 kg (18–40 lb), but females tend to be lighter. The head-and-body length is typically between 67 and 100 cm (26–39 in).[18] Males tend to be sturdier than females.[19] Prominent characteristics include the small head, large ears, spotted and striped coat, long legs and a black-tipped tail that is around 30 cm (12 in) long.[20][21] The serval has the longest legs of any cat relative to its body size, largely due to the greatly elongated metatarsal bones in the feet.[22][23] The toes are elongated as well, and unusually mobile.[22]
+The coat is basically golden-yellow to buff, and extensively marked with black spots and stripes.[19] The spots show great variation in size. Melanistic servals are also known.[22] Facial features include the brownish or greenish eyes, white whiskers on the snout and near the ears, ears as large as those of a domestic cat (but large relative to the size of the head) and black on the back with a white horizontal band in the middle, whitish chin, and spots and streaks on the cheeks and the forehead. Three to four black stripes run from the back of the head onto the shoulders, and then break into rows of spots. The white underbelly has dense and fluffy basal fur, and the soft guard hairs (the layer of fur protecting the basal fur) are 5–10 cm (2–4 in) long. Guard hairs are up to 3 cm (1 1⁄4 in) long on the neck, back and the flanks, and are merely 1 cm (1⁄2 in) long on the face.[21][24][19] The closely set ears are black on the back with a horizontal white band;[19] the ears can rotate up to 180 degrees independently of each other.[22] The serval has a good sense of smell, hearing and vision.[21]
+The serval is similar to the sympatric caracal, but has a narrower spoor, a rounder skull, and lacks its prominent ear tufts. The African golden cat is darker, with different cranial features.[19] It resembles the cheetah in its build and coat pattern, though not in size.[21] The serval shares its adaptations to its marshy habitat with the jungle cat; both cats have large and sharp ears that help in locating the prey efficiently, and their long legs raise them above muddy ground and water.[26]
+In North Africa, the serval is known only from Morocco and has been reintroduced in Tunisia, but is feared to be extinct in Algeria. It inhabits semi-arid areas and cork oak forests close to the Mediterranean Sea, but avoids rainforests and arid areas. It occurs in the Sahel, and is widespread in Southern Africa. It inhabits grasslands, moorlands and bamboo thickets at high altitudes up to 3,800 m (12,500 ft) on Mount Kilimanjaro. It prefers areas close to water bodies such as wetland and savanna, which provide cover such as reeds and tall grasses.[1][19]
+In 2014 and 2015, it was recorded in the floodplains and gallery forests of Benin’s Pendjari National Park by camera-traps.[27]
+In the East Sudanian Savanna, it was recorded in the transboundary Dinder–Alatash protected area complex during surveys between 2015 and 2018.[28]
+In Zambia's Luambe National Park, the population density was recorded as 0.1/km2 (0.26/sq mi) in 2011.[29]
+In South Africa, the serval was recorded in Free State, eastern Northern Cape, and southern North West.[30]
+In Namibia, it is present in Khaudum and Mudumu National Parks.[31]
+The serval is active in the day as well as at night; activity might peak in early morning, around twilight and at midnight. Servals might be active for a longer time on cool or rainy days. During the hot midday, they rest or groom themselves in the shade of bushes and grasses. Servals remain cautious of their vicinity, though they may be less alert when no large carnivores or prey animals are around. Servals walk as much as 2 to 4 kilometres (1 1⁄4 to 2 1⁄2 miles) every night.[20][18] Servals will often use special trails to reach certain hunting areas. A solitary animal, there is little social interaction among servals except in the mating season, when pairs of opposite sexes may stay together. The only long-lasting bond appears to be of the mother and her cubs, which leave their mother only when they are a year old.[19]
+Both males and females establish home ranges, and are most active only in certain regions ('core areas') within them. The area of these ranges can vary from 10 to 32 square kilometres (4 to 12 square miles); prey density, availability of cover and human interference could be significant factors in determining their size.[19][33] Home ranges might overlap extensively, but occupants show minimal interaction. Aggressive encounters are rare, as servals appear to mutually avoid one another rather than fight and defend their ranges. Agonistic behaviour involves vertical movement of the head (contrary to the horizontal movement observed in other cats), raising the hair and the tail, displaying the teeth and the white band on the ears, and yowling. Individuals mark their ranges and preferred paths by spraying urine on nearby vegetation, dropping scats along the way, and rubbing their mouth on grasses or the ground while releasing saliva. Servals tend to be sedentary, shifting only a few kilometres away even if they leave their range.[19][18]
+The serval is vulnerable to hyaenas and wild dogs. It will seek cover to escape their view, and, if the predator is very close, immediately flee in long leaps, changing its direction frequently and with the tail raised.[18] The serval is an efficient, though not frequent, climber; an individual was observed to have climbed a tree to a height of more than 9 metres (30 feet) to escape dogs.[22] Like many cats, the serval is able to purr;[34] it also has a high-pitched chirp, and can hiss, cackle, growl, grunt and meow.[22]
+The serval is a carnivore that preys on rodents, particularly vlei rats, small birds, frogs, insects and reptiles, and also feeds on grass that can facilitate digestion or act as an emetic. Up to 90% of the preyed animals weigh less than 200 g (7 oz); occasionally it also hunts larger prey such as duikers, hares, flamingoes and young antelopes.[22] The percentage of rodents in the diet has been estimated at 80-97%.[33][35][36] Apart from vlei rats, other rodents recorded frequently in the diet include the African grass rat, African pygmy mouse and multimammate mice.[19]
+Servals locate prey by their strong sense of hearing. To kill small prey, the serval will slowly stalk it, then pounce on it with the forefeet directed toward the chest, and finally land on it with its forelegs outstretched. The prey, receiving a blow from one or both of the serval's forepaws, is incapacitated, and the serval gives it a bite on the head or the neck and immediately swallows it. Snakes are dealt more blows and even bites, and may be consumed even as they are moving. Larger prey, such as larger birds, are killed by a sprint followed by a leap to catch them as they are trying to flee, and are eaten slowly. Servals have been observed caching large kills to be consumed later by concealing them in dead leaves and grasses. Servals typically get rid of the internal organs of rodents while eating, and pluck feathers from birds before consuming them. During a leap, a serval can reach more than 2 m (6 ft 7 in) above the ground and cover a horizontal distance of up to 3.6 m (11 ft 10 in).  Servals appear to be efficient hunters; a study in Ngorongoro showed that servals were successful in half of their hunting attempts, regardless of the time of hunting, and a mother serval was found to have a success rate of 62%. The number of kills in a 24-hour period averaged 15 to 16. Scavenging has been observed, but very rarely.[22][19]
+Both sexes become sexually mature when they are one to two years old. Oestrus in females lasts one to four days; it typically occurs once or twice a year, though it can occur three or four times a year if the mother loses her litters.[37] Observations of captive servals suggest that when a female enters oestrus, the rate of urine-marking increases in her as well as the males in her vicinity. Zoologist Jonathan Kingdon described the behaviour of a female serval in oestrus in his 1997 book East African Mammals. He noted that she would roam restlessly, spray urine frequently holding her vibrating tail in a vertical manner, rub her head near the place she has marked, salivate continuously, give out sharp and short "miaow"s that can be heard for quite a distance, and rub her mouth and cheeks against the face of an approaching male. The time when mating takes place varies geographically; births peak in winter in Botswana, and toward the end of the dry season in the Ngorongoro Crater. A trend generally observed across the range is that births precede the breeding season of murid rodents.[22]
+Gestation lasts for two to three months, following which a litter of one to four kittens is born. Births take place in secluded areas, for example in dense vegetation or burrows abandoned by aardvarks and porcupines. Blind at birth, newborn weigh nearly 250 g (9 oz) and have soft, woolly hair (greyer than in adults) and unclear markings. The eyes open after nine to thirteen days. Weaning begins a month after birth; the mother brings small kills to her kittens and calls out to them as she approaches the "den".[22] A mother with young kittens rests for a notably lesser time and has to spend almost twice the time and energy for hunting than do other servals.[33] If disturbed, the mother shifts her kittens one by one to a more secure place.[24] Kittens eventually start accompanying their mother to hunts. At around six months, they acquire their permanent canines and begin to hunt themselves; they leave their mother at about 12 months of age. They may reach sexual maturity from 12 to 25 months of age.[22] Life expectancy is about 10 years in the wild, and up to 20 years in captivity.[38]
+A major threat to the survival of the serval include the degradation of wetlands and grasslands. Trade of serval skins, though on the decline, still occurs in countries such as Benin and Senegal. In West Africa, the serval has significance in traditional medicine. Pastoralists often kill servals to protect their livestock, though servals generally do not prey on livestock.[1]
+The serval is listed as least concern on the IUCN Red List, and is included in CITES Appendix II. It occurs in several protected areas across its range. Hunting of servals is prohibited in Algeria, Botswana, Congo, Kenya, Liberia, Morocco, Mozambique, Nigeria, Rwanda, Tunisia and South Africa's Cape Province; hunting regulations apply in Angola, Burkina Faso, Central African Republic, the Democratic Republic of the Congo, Ghana, Malawi, Senegal, Sierra Leone, Somalia, Tanzania, Togo and Zambia.[1]
+The association of servals with human beings dates to the time of Ancient Egypt.[39] Servals are depicted as gifts or traded objects from Nubia in Egyptian art.[40]
+Like many other species of felid, servals are occasionally kept as pets, although their wild nature means that ownership of servals is regulated in most countries.[41][42][43][44][45][46][47][48]

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+The rising average temperature of Earth's climate system, called global warming, is driving changes in rainfall patterns, extreme weather, arrival of seasons, and more. Collectively, global warming and its effects are known as climate change.  While there have been prehistoric periods of global warming, observed changes since the mid-20th century have been unprecedented in rate and scale.[1]
+The Intergovernmental Panel on Climate Change (IPCC) concluded that "human influence on climate has been the dominant cause of observed warming since the mid-20th century". These findings have been recognized by the national science academies of major nations and are not disputed by any scientific body of national or international standing.[4] The largest human influence has been the emission of greenhouse gases, with over 90% of the impact from carbon dioxide and methane.[5] Fossil fuel burning is the principal source of these gases, with agricultural emissions and deforestation also playing significant roles. Temperature rise is enhanced by self-reinforcing climate feedbacks, such as loss of snow cover, increased water vapour, and melting permafrost.
+Land surfaces are heating faster than the ocean surface, leading to heat waves, wildfires, and the expansion of deserts.[6] Increasing atmospheric energy and rates of evaporation are causing more intense storms and weather extremes, damaging infrastructure and agriculture.[7] Surface temperature increases are greatest in the Arctic and have contributed to the retreat of glaciers, permafrost, and sea ice. Environmental impacts include the extinction or relocation of many species as their ecosystems change, most immediately in coral reefs, mountains, and the Arctic. Surface temperatures would stabilize and decline a little if emissions were cut off, but other impacts will continue for centuries, including rising sea levels from melting ice sheets, rising ocean temperatures, and ocean acidification from elevated levels of carbon dioxide.[8]
+Extreme weather. Drought and high temperatures worsened the 2020 bushfires in Australia.
+Habitat destruction. Many arctic animals rely on sea ice, which has been disappearing in a warming Arctic.
+Storm intensification. Bangladesh after Cyclone Sidr is an example of catastrophic flooding from increased rainfall.
+Heat wave intensification. Events like the June 2019 European heat wave are becoming more common.
+Farming. Droughts, rising temperatures, and extreme weather negatively impact agriculture.
+Tidal flooding. Sea level rise increases flooding in low lying coastal regions. Shown: Venice, Italy.
+Arctic warming. Permafrost thaws undermine infrastructure and release methane in a positive feedback loop.
+Ecological collapse possibilities. Bleaching has damaged the Great Barrier Reef and threatens reefs worldwide.
+Pest propagation. Mild winters allow more pine beetles to survive to kill large swaths of forest.
+Environmental migration. Sparser rainfall leads to desertification that harms agriculture and can displace populations.
+Mitigation efforts to address global warming include the development and deployment of low carbon energy technologies, policies to reduce fossil fuel emissions, reforestation, forest preservation, as well as the development of potential climate engineering technologies. Societies and governments are also working to adapt to current and future global warming impacts, including improved coastline protection, better disaster management, and the development of more resistant crops.
+Countries work together on climate change under the umbrella of the United Nations Framework Convention on Climate Change (UNFCCC), which has near-universal membership. The goal of the convention is to "prevent dangerous anthropogenic interference with the climate system". The IPCC has stressed the need to keep global warming below 1.5 °C (2.7 °F) compared to pre-industrial levels in order to avoid some irreversible impacts.[10] With current policies and pledges, global warming by the end of the century is expected to reach about 2.8 °C (5.0 °F).[11] At the current greenhouse gas (GHG) emission rate, the carbon budget for staying below 1.5 °C (2.7 °F) would be exhausted by 2028.[12]
+Multiple independently produced instrumental datasets show that the climate system is warming,[14] with the 2009–2018 decade being 0.93 ± 0.07 °C (1.67 ± 0.13 °F) warmer than the pre-industrial baseline (1850–1900).[15] Currently, surface temperatures are rising by about 0.2 °C (0.36 °F) per decade.[16] Since 1950, the number of cold days and nights has decreased, and the number of warm days and nights has increased.[17] Historical patterns of warming and cooling, like the Medieval Climate Anomaly and the Little Ice Age, were not as synchronous as current warming, but may have reached temperatures as high as those of the late-20th century in a limited set of regions.[18] There have been prehistorical episodes of global warming, such as the Paleocene–Eocene Thermal Maximum.[19] However, the observed rise in temperature and CO2 concentrations has been so rapid that even abrupt geophysical events that took place in Earth's history do not approach current rates.[20]
+Climate proxy records show that natural variations offset the early effects of the Industrial Revolution, so there was little net warming between the 18th century and the mid-19th century,[21] when thermometer records began to provide global coverage.[22] The Intergovernmental Panel on Climate Change (IPCC) has adopted the baseline reference period 1850–1900 as an approximation of pre-industrial global mean surface temperature.[21]
+The warming evident in the instrumental temperature record is consistent with a wide range of observations, documented by many independent scientific groups.[23] Although the most common measure of global warming is the increase in the near-surface atmospheric temperature, over 90% of the additional energy in the climate system over the last 50 years has been stored in the ocean, warming it.[24] The remainder of the additional energy has melted ice and warmed the continents and the atmosphere.[25] The ocean heat uptake drives thermal expansion which has contributed to observed sea level rise.[26] Further indicators of climate change include an increase in the frequency and intensity of heavy precipitation, melting of snow and land ice and increased atmospheric humidity.[27] Flora and fauna also portray behaviour consistent with warming, such as the earlier flowering of plants in spring.[28]
+Global warming refers to global averages, with the amount of warming varying by region. Since the pre-industrial period, global average land temperatures have increased almost twice as fast as global average temperatures.[29] This is due to the larger heat capacity of oceans and because oceans lose more heat by evaporation.[30] Patterns of warming are independent of the locations of greenhouse gas emissions because the gases persist long enough to diffuse across the planet; however, localized black carbon deposits on snow and ice do contribute to Arctic warming.[31]
+The Northern Hemisphere and North Pole have warmed much faster than the South Pole and Southern Hemisphere. The Northern Hemisphere not only has much more land, but also more snow area and sea ice, because of how the land masses are arranged around the Arctic Ocean. As these surfaces flip from being reflective to dark after the ice has melted, they start absorbing more heat. The Southern Hemisphere already had little sea ice in summer before it started warming.[32] Arctic temperatures have increased and are predicted to continue to increase during this century at over twice the rate of the rest of the world.[33] Melting of glaciers and ice sheets in the Arctic disrupts ocean circulation, uncluding a weakened Gulf Stream, causing increased warming in some areas.[34]
+Although record-breaking years attract considerable media attention, individual years are less significant than the overall global surface temperature, which is subject to short-term fluctuations that overlie long-term trends.[35] An example of such an episode is the slower rate of surface temperature increase from 1998 to 2012, which was described as the global warming hiatus.[36] Throughout this period, ocean heat storage continued to progress steadily upwards, and in subsequent years, surface temperatures have spiked upwards. The slower pace of warming can be attributed to a combination of natural fluctuations, reduced solar activity, and increased reflection sunlight of by particles from volcanic eruptions.[37]
+By itself, the climate system experiences various cycles which can last for years (such as the El Niño–Southern Oscillation) to decades or centuries.[38] Other changes are caused by an imbalance of energy at the top of the atmosphere: external forcings. These forcings are "external" to the climate system, but not always external to the Earth.[39] Examples of external forcings include changes in the composition of the atmosphere (e.g. increased concentrations of greenhouse gases), solar luminosity, volcanic eruptions, and variations in the Earth's orbit around the Sun.[40]
+Attribution of climate change is the effort to scientifically show which mechanisms are responsible for observed changes in Earth's climate. First, known internal climate variability and natural external forcings need to be ruled out. Therefore, a key approach is to use computer modelling of the climate system to determine unique "fingerprints" for all potential causes. By comparing these fingerprints with observed patterns and evolution of climate change, and the observed history of the forcings, the causes of the observed changes can be determined.[41] For example, solar forcing can be ruled out as major cause because its fingerprint is warming in the entire atmosphere, and only the lower atmosphere has warmed as expected for greenhouse gases.[42] The major causes of current climate change are primarily greenhouse gases, and secondarily land use changes, and aerosols and soot.[43]
+Greenhouse gases trap heat radiating from the Earth to space.[44] This heat, in the form of infrared radiation, gets absorbed and emitted by these gases in the atmosphere, thus warming the lower atmosphere and the surface. Before the Industrial Revolution, naturally occurring amounts of greenhouse gases caused the air near the surface to be warmer by about 33 °C (59 °F) than it would be in their absence.[45] Without the Earth's atmosphere, the Earth's average temperature would be well below the freezing temperature of water.[46] While water vapour (~50%) and clouds (~25%) are the biggest contributors to the greenhouse effect, they increase as a function of temperature and are therefore considered feedbacks. Increased concentrations of gases such as CO2 (~20%), ozone and N2O are external forcing on the other hand.[47] Ozone acts as a greenhouse gas in the lowest layer of the atmosphere, the troposphere. Furthermore, it is highly reactive and interacts with other greenhouse gases and aerosols.[48]
+Human activity since the Industrial Revolution, mainly extracting and burning fossil fuels,[49] has increased the amount of greenhouse gases in the atmosphere. This CO2, methane, tropospheric ozone, CFCs, and nitrous oxide has increased radiative forcing. In 2018, the concentrations of CO2 and methane had increased by about 45% and 160%, respectively, since pre-industrial times.[50] In 2013, CO2 readings taken at the world's primary benchmark site in Mauna Loa surpassing 400 ppm for the first time.[51] These levels are much higher than at any time during the last 800,000 years, the period for which reliable data have been collected from ice cores.[52] Less direct geological evidence indicates that CO2 values have not been this high for millions of years.[53]
+Global anthropogenic greenhouse gas emissions in 2018 excluding land use change were equivalent to 52 billion tonnes of carbon dioxide. Of these emissions, 72% was carbon dioxide from fossil fuel burning and industry, 19% was methane, largely from livestock,[54] 6% was nitrous oxide, mainly from agriculture, and 3% was fluorinated gases.[55] A further 4 billion tonnes of CO2 was released as a consequence of land use change, which is primarily due to deforestation.[56] From a production standpoint, the primary sources of global GHG emissions are estimated as: electricity and heat (25%), agriculture and forestry (24%), industry (21%), and transportation (14%).[57] Consumption based estimates of GHG emissions offer another useful way to understand sources of global warming, and may better capture the effects of trade.[58] From a consumption standpoint, the dominant sources of global 2010 emissions were found to be: food (30%), washing, heating, and lighting (26%); personal transport and freight (20%); and building construction (15%).[59]
+Despite the contribution of deforestation to GHG emissions, the Earth's land surface, particularly its forests, remain a significant carbon sink for CO2. Natural processes, such as carbon fixation in the soil and photosynthesis, more than offset the GHG contributions from deforestation. The land surface sink is estimated to remove about 11 billion tonnes of CO2 annually from the atmosphere, or about 29% of global CO2 emissions.[60] The ocean also serves as a significant carbon sink via a two-step process. First, CO2 dissolves in the surface water. Afterwards, the ocean's overturning circulation distributes it deep into the ocean's interior, where it accumulates over time as part of the carbon cycle. Over the last two decades, the world's oceans have removed between 20 and 30% of emitted CO2.[61] The strength of both the land and ocean sinks increase as CO2 levels in the atmosphere rise. In this respect they act as negative feedbacks in global warming.[62]
+Humans change the Earth's surface mainly to create more agricultural land. Today agriculture takes up 50% of the world's habitable land, while 37% is forests,[63] and that latter figure continues to decrease,[64] largely due to continued forest loss in the tropics.[65] This deforestation is the most significant aspect of land use change affecting global warming. The main causes are: deforestation through permanent land use change for agricultural products such as beef and palm oil (27%), forestry/forest products (26%), short term agricultural cultivation (24%), and wildfires (23%).[66]
+In addition to impacting greenhouse gas concentrations, land use changes affect global warming through a variety of other chemical and physical dynamics. Changing the type of vegetation in a region impacts the local temperature by changing how much sunlight gets reflected back into space, called albedo, and how much heat is lost by evaporation. For instance, the change from a dark forest to grassland makes the surface lighter, causing it to reflect more sunlight. Deforestation can also contribute to changing temperatures by affecting the release of aerosols and other chemical compounds that affect clouds; and by changing wind patterns when the land surface has different obstacles.[67] Globally, these effects are estimated to have led to a slight cooling, dominated by an increase in surface albedo.[68] But there is significant geographic variation in how this works. In the tropics the net effect is to produce a significant warming, while at latitudes closer to the poles a loss of albedo leads to an overall cooling effect.[67]
+Air pollution, in the form of aerosols, not only puts a large burden on human health, but also affects the climate on a large scale.[69] From 1961 to 1990, a gradual reduction in the amount of sunlight reaching the Earth's surface was observed, a phenomenon popularly known as global dimming,[70] typically attributed to aerosols from biofuel and fossil fuel burning.[71] Aerosol removal by precipitation gives tropospheric aerosols an atmospheric lifetime of only about a week, while stratospheric aerosols can remain in the atmosphere for a few years.[72] Globally, aerosols have been declining since 1990, removing some of the masking of global warming that they had been providing.[73]
+In addition to their direct effect by scattering and absorbing solar radiation, aerosols have indirect effects on the Earth's radiation budget. Sulfate aerosols act as cloud condensation nuclei and thus lead to clouds that have more and smaller cloud droplets. These clouds reflect solar radiation more efficiently than clouds with fewer and larger droplets.[74] This effect also causes droplets to be of more uniform size, which reduces the growth of raindrops and makes clouds more reflective to incoming sunlight.[75] Indirect effects of aerosols are the largest uncertainty in radiative forcing.[76]
+While aerosols typically limit global warming by reflecting sunlight, black carbon in soot that falls on snow or ice can contribute to global warming. Not only does this increase the absorption of sunlight, it also increases melting and sea level rise.[77] Limiting new black carbon deposits in the Arctic could reduce global warming by 0.2 °C by 2050.[78]
+As the Sun is the Earth's primary energy source, changes in incoming sunlight directly affect the climate system.[79] Solar irradiance has been measured directly by satellites,[80] and indirect measurements are available beginning in the early 1600s.[79] There has been no upward trend in the amount of the Sun's energy reaching the Earth, so it cannot be responsible for the current warming.[81] Physical climate models are also unable to reproduce the rapid warming observed in recent decades when taking into account only variations in solar output and volcanic activity.[82] Another line of evidence for the warming not being due to the Sun is how temperature changes differ at different levels in the Earth's atmosphere.[83] According to basic physical principles, the greenhouse effect produces warming of the lower atmosphere (the troposphere), but cooling of the upper atmosphere (the stratosphere).[84] If solar variations were responsible for the observed warming, warming of both the troposphere and the stratosphere would be expected, but that has not been the case.[42] Explosive volcanic eruptions represent the largest natural forcing over the industrial era. When the eruption is sufficiently strong with sulfur dioxide reaching the stratosphere, sunlight can be partially blocked for a couple of years, with a temperature signal lasting about twice as long.[85]
+The response of the climate system to an initial forcing is increased by self-reinforcing feedbacks and reduced by balancing feedbacks.[87] The main balancing feedback to global temperature change is radiative cooling to space as infrared radiation, which increases strongly with increasing temperature.[88] The main reinforcing feedbacks are the water vapour feedback, the ice–albedo feedback, and probably the net effect of clouds.[89] Uncertainty over feedbacks is the major reason why different climate models project different magnitudes of warming for a given amount of emissions.[90]
+As air gets warmer, it can hold more moisture. After an initial warming due to emissions of greenhouse gases, the atmosphere will hold more water. As water is a potent greenhouse gas, this further heats the climate: the water vapour feedback.[89] The reduction of snow cover and sea ice in the Arctic reduces the albedo of the Earth's surface.[91] More of the Sun's energy is now absorbed in these regions, contributing to Arctic amplification, which has caused Arctic temperatures to increase at more than twice the rate of the rest of the world.[92] Arctic amplification also causes methane to be released as permafrost melts, which is expected to surpass land use changes as the second strongest anthropogenic source of greenhouse gases by the end of the century.[93]
+Cloud cover may change in the future. If cloud cover increases, more sunlight will be reflected back into space, cooling the planet. Simultaneously, the clouds enhance the greenhouse effect, warming the planet. The opposite is true if cloud cover decreases. It depends on the cloud type and location which process is more important. Overall, the net feedback over the industrial era has probably been self-reinforcing.[94]
+Roughly half of each year's CO2 emissions have been absorbed by plants on land and in oceans.[95] Carbon dioxide and an extended growing season have stimulated plant growth making the land carbon cycle a balancing feedback. Climate change also increases droughts and heat waves that inhibit plant growth, which makes it uncertain whether this balancing feedback will persist in the future.[96] Soils contain large quantities of carbon and may release some when they heat up.[97] As more CO2 and heat are absorbed by the ocean, it is acidifying and ocean circulation can change, changing the rate at which the ocean can absorb atmospheric carbon.[98]
+Future warming depends on the strenght of climate feedbacks and on emissions of greenhouse gases.[99] The former is often estimated using climate models. A climate model is a representation of the physical, chemical, and biological processes that affect the climate system.[100] They also include changes in the Earth's orbit, historical changes in the Sun's activity, and volcanic forcing.[101] Computer models attempt to reproduce and predict the circulation of the oceans, the annual cycle of the seasons, and the flows of carbon between the land surface and the atmosphere.[102] There are more than two dozen scientific institutions that develop climate models.[103] Models not only project different future temperature with different emissions of greenhouse gases, but also do not fully agree on the strength of different feedbacks on climate sensitivity and the amount of inertia of the system.[104]
+The physical realism of models is tested by examining their ability to simulate contemporary or past climates.[105] Past models have underestimated the rate of Arctic shrinkage[106] and underestimated the rate of precipitation increase.[107] Sea level rise since 1990 was underestimated in older models, but now agrees well with observations.[108] The 2017 United States-published National Climate Assessment notes that "climate models may still be underestimating or missing relevant feedback processes".[109]
+Four Representative Concentration Pathways (RCPs) are used as input for climate models: "a stringent mitigation scenario (RCP2.6), two intermediate scenarios (RCP4.5 and RCP6.0) and one scenario with very high GHG [greenhouse gas] emissions (RCP8.5)".[110]  RCPs only look at concentrations of greenhouse gases, and so does not include the response of the carbon cycle.[111] Climate model projections summarized in the IPCC Fifth Assessment Report indicate that, during the 21st century, the global surface temperature is likely to rise a further 0.3 to 1.7 °C (0.5 to 3.1 °F) in a moderate scenario, or as much as 2.6 to 4.8 °C (4.7 to 8.6 °F) in an extreme scenario, depending on the rate of future greenhouse gas emissions and on climate feedback effects.[112]
+A subset of climate models add societal factors to a simple physical climate model. These models simulate how population, economic growth, and energy use affect – and interact with – the physical climate. With this information, these models can produce scenarios of how greenhouse gas emissions may vary in the future. This output is then used as input for physical climate models to generate climate change projections.[113] In some scenarios emissions continue to rise over the century, while others have reduced emissions.[114] Fossil fuel resources are abundant, and cannot be relied on to limit carbon emissions in the 21st century.[115] Emission scenarios can be combined with modelling of the carbon cycle to predict how atmospheric concentrations of greenhouse gases might change in the future.[116] According to these combined models, by 2100 the atmospheric concentration of CO2 could be as low as 380 or as high as 1400 ppm, depending on the Shared Socioeconomic Pathway (SSP) and the mitigation scenario.[117]
+The remaining carbon emissions budget is determined from modelling the carbon cycle and climate sensitivity to greenhouse gases.[118] According to the IPCC, global warming can be kept below 1.5 °C with a two-thirds chance if emissions after 2018 do not exceed 420 or 570 GtCO2 depending on the choice of the measure of global temperature. This amount corresponds to 10 to 13 years of current emissions. There are high uncertainties about the budget; for instance, it may be 100 GtCO2 smaller due to methane release from permafrost and wetlands.[119]
+The environmental effects of global warming are broad and far-reaching. They include effects on the oceans, ice, and weather and may occur gradually or rapidly. Evidence for these effects come from studying climate change in the past, modelling and modern observations.[121] Since the 1950s, droughts and heat waves have appeared simultaneously with increasing frequency.[122] Extremely wet or dry events within the monsoon period have increased in India and East Asia.[123] Various mechanisms have been identified that might explain extreme weather in mid-latitudes from the rapidly warming Arctic, such as the jet stream becoming more erratic.[124] The maximum rainfall and wind speed from hurricanes and typhoons is likely increasing.[125]
+Between 1993 and 2017, the global mean sea level rose on average by 3.1 ± 0.3 mm per year, with an acceleration detected as well.[126] Over the 21st century, the IPCC projects that in a very high emissions scenario the sea level could rise by 61–110 cm.[127] The rate of ice loss from glaciers and ice sheets in the Antarctic is a key area of uncertainty since this source could account for 90% of the potential sea level rise:[128] increased ocean warmth is undermining and threatening to unplug Antarctic glacier outlets, potentially resulting in more rapid sea level rise.[129] The retreat of non-polar glaciers also contributes to sea level rise.[130]
+Global warming has led to decades of shrinking and thinning of the Arctic sea ice, making it vulnerable to atmospheric anomalies.[131] Projections of declines in Arctic sea ice vary.[132] While ice-free summers are expected to be rare at 1.5 °C (2.7 °F) degrees of warming, they are set to occur once every three to ten years at a warming level of 2.0 °C (3.6 °F),[133] increasing the ice–albedo feedback.[134] Higher atmospheric CO2 concentrations have led to an increase in dissolved CO2, which causes ocean acidification.[135] Furthermore, oxygen levels decrease because oxygen is less soluble in warmer water, an effect known as ocean deoxygenation.[136]
+The long-term effects of global warming include further ice melt, ocean warming, sea level rise, and ocean acidification. On the timescale of centuries to millennia, the magnitude of global warming will be determined primarily by anthropogenic CO2 emissions.[137] This is due to carbon dioxide's very long lifetime in the atmosphere.[137] Carbon dioxide is slowly taking up by the ocean, such that ocean acidification will continue for hundreds to thousands of years.[138] The emissions are estimated to have prolonged the current interglacial period by at least 100,000 years.[139] Because the great mass of glaciers and ice caps depressed the Earth's crust, another long-term effect of ice melt and deglaciation is the gradual rising of landmasses, a process called post-glacial rebound.[140] Sea level rise will continue over many centuries, with an estimated rise of 2.3 metres per degree Celsius (4.2 ft/°F) after 2000 years.[141]
+If global warming exceeds 1.5 °C, there is a greater risk of passing through ‘tipping points’, thresholds beyond which certain impacts can no longer be avoided even if temperatures are reduced.[142] Some large-scale changes could occur abruptly, i.e. over a short time period. One potential source of abrupt tipping would be the rapid release of methane and carbon dioxide from permafrost, which would amplify global warming.[143] Another example is the possibility for the Atlantic Meridional Overturning Circulation to collapse,[144] which could trigger cooling in the North Atlantic, Europe, and North America.[145] If multiple temperature and carbon cycle tipping points re-inforce each other, or if there were to be strong threshold behaviour in cloud cover, there could be a global tipping into a hothouse Earth.[146]
+Recent warming has driven many terrestrial and freshwater species poleward and towards higher altitudes.[147] Higher atmospheric CO2 levels and an extended growing season have resulted in global greening, whereas heatwaves and drought have reduced ecosystem productivity in some regions. The future balance of these opposing effects is unclear.[148] Global warming has contributed to the expansion of drier climatic zones, such as, probably, the expansion of deserts in the subtropics.[149] Without substantial actions to reduce the rate of global warming, land-based ecosystems risk major shifts in their composition and structure.[150] Overall, it is expected that climate change will result in the extinction of many species and reduced diversity of ecosystems.[151]
+The ocean has heated more slowly than the land, but plants and animals in the ocean have migrated towards the colder poles as fast as or faster than species on land.[152] Just as on land, heat waves in the ocean occur more due to climate change, with harmful effects found on a wide range of organisms such as corals, kelp, and seabirds.[153] Ocean acidification threatens damage to coral reefs, fisheries, protected species, and other natural resources of value to society.[154] Coastal ecosystems are under stress, with almost half of wetlands having disappeared as a consquence of climate change and other human impacts. Harmful algae blooms have increased due to warming, ocean deoxygenation and eutrophication.[155]
+Ecological collapse possibilities. Bleaching has damaged the Great Barrier Reef and threatens reefs worldwide.[156]
+Extreme weather. Drought and high temperatures worsened the 2020 bushfires in Australia.[157]
+Arctic warming. Permafrost thaws undermine infrastructure and release methane in a positive feedback loop.[143]
+Habitat destruction. Many arctic animals rely on sea ice, which has been disappearing in a warming Arctic.[158]
+Pest propagation. Mild winters allow more pine beetles to survive to kill large swaths of forest.[159]
+The effects of climate change on human systems, mostly due to warming and shifts in precipitation, have been detected worldwide. The social impacts of climate change will be uneven across the world.[160] All regions are at risk of experiencing negative impacts,[161] with low-latitude, less developed areas facing the greatest risk.[162] Global warming has likely already increased global economic inequality, and is projected to do so in the future.[163] Regional impacts of climate change are now observable on all continents and across ocean regions.[164] The Arctic, Africa, small islands, and Asian megadeltas are regions that are likely to be especially affected by future climate change.[165] Many risks increase with higher magnitudes of global warming.[166]
+Crop production will probably be negatively affected in low-latitude countries, while effects at northern latitudes may be positive or negative.[167] Global warming of around 4 °C relative to late 20th century levels could pose a large risk to global and regional food security.[168] The impact of climate change on crop productivity for the four major crops was negative for wheat and maize, and neutral for soy and rice, in the years 1960–2013.[169] Up to an additional 183 million people worldwide, particularly those with lower incomes, are at risk of hunger as a consequence of warming.[170] While increased CO2 levels help crop growth at lower temperature increases, those crops do become less nutritious.[170] Based on local and indigenous knowledge, climate change is already affecting food security in mountain regions in South America and Asia, and in various drylands, particularly in Africa.[170] Regions dependent on glacier water, regions that are already dry, and small islands are also at increased risk of water stress due to climate change.[171]
+In small islands and mega deltas, inundation from sea level rise is expected to threaten vital infrastructure and human settlements.[172] This could lead to homelessness in countries with low-lying areas such as Bangladesh, as well as statelessness for populations in island nations, such as the Maldives and Tuvalu.[173] Climate change can be an important driver of migration, both within and between countries.[174]
+The majority of severe impacts of climate change are expected in sub-Saharan Africa and South-East Asia, where existing poverty is exacerbated.[175] The World Bank estimates that global warming could drive over 120 million people into poverty by 2030.[176] Current inequalities between men and women, between rich and poor and between people of different ethnicity have been observed to worsen as a consequence of climate variability and climate change.[177] Existing stresses include poverty, political conflicts, and ecosystem degradation. Regions may even become uninhabitable, with humidity and temperatures reaching levels too high for humans to survive.[178]
+Generally, impacts on public health will be more negative than positive.[179] Impacts include the direct effects of extreme weather, leading to injury and loss of life;[180] and indirect effects, such as undernutrition brought on by crop failures.[181] Various infectious diseases are more easily transmitted in a warming climate, such as dengue fever, which affects children most severely, and malaria.[182] Young children are further the most vulnerable to food shortages, and together with older people to extreme heat.[183] Climate change has been linked to an increase in violent conflict by amplifying poverty and economic shocks, which are well-documented drivers of these conflicts.[184] Links have been made between a wide range of violent behaviour including, violent crimes, civil unrest, and wars, but conclusive scientific evidence remains elusive.[185]
+Environmental migration. Sparser rainfall leads to desertification that harms agriculture and can displace populations.[186]
+Farming. Droughts, rising temperatures, and extreme weather negatively impact agriculture.[187]
+Tidal flooding. Sea level rise increases flooding in low lying coastal regions. Shown: Venice, Italy.[188]
+Storm intensification. Bangladesh after Cyclone Sidr is an example of catastrophic flooding from increased rainfall.[189]
+Heat wave intensification. Events like the June 2019 European heat wave are becoming more common.[190]
+Mitigation of and adaptation to climate change are two complementary responses to global warming. Successful adaptation is easier if there are substantial emission reductions. Many of the countries that have contributed least to global greenhouse gas emissions are among the most vulnerable to climate change, which raises questions about justice and fairness with regard to mitigation and adaptation.[192]
+Climate change impacts can be mitigated by reducing greenhouse gas emissions and by enhancing the capacity of Earth's surface to absorb greenhouse gases from the atmosphere.[193]
+In order to limit global warming to less than 1.5°C with a high likelihood of success, the IPCC estimates that global GHG emissions will need to be net zero by 2050,[194] or by 2070 with a 2°C target. This will require far-reaching, systemIc changes on an unprecedented scale in energy, land, cities, transport, buildings, and industry.[195] To make progress towards that goal, the United Nations Environment Programme estimates that, within the next decade, countries will need to triple the amount of reductions they have committed to in their current Paris agreements.[196]
+Long-term scenarios all point to rapid and significant investment in renewable energy and energy efficiency as key to reducing GHG emissions.[197] These technologies include solar and wind power, bioenergy, geothermal energy, and hydroelectricity. Combined, they are capable of supplying several times the world’s current energy needs.[198] Solar PV and wind, in particular, have seen substantial growth and progress over the last few years,[199] such that they are currently among the cheapest sources of new power generation.[200] Renewables represented 75% of all new electricity generation installed in 2019, with solar and wind constituting nearly all of that amount.[201] However, fossil fuels continue to dominate world energy supplies. In 2018 fossil fuels produced 80% of the world’s energy, with modern renewable sources, including solar and wind power, accounting for around 11%.[202]
+There are obstacles to the rapid development of renewable energy. Environmental and land use concerns are sometimes associated with large solar, wind and hydropower projects.[203] Solar and wind power also require energy storage systems and other modifications to the electricity grid to operate effectively,[204] although several storage technologies are now emerging to supplement the traditional use of  pumped-storage hydropower.[205] The use of rare metals and other hazardous materials has also been raised as a concern with solar power.[206] The use of bioenergy is often not carbon neutral, and may have negative consequences for food security,[207] largely due to the amount of land required compared to other renewable energy options.[208]
+For certain energy supply needs, as well as specific CO2-intensive heavy industries, carbon capture and storage may be a viable method of reducing CO2 emissions. Although high costs have been a concern with this technology,[209] it may be able to play a significant role in limiting atmospheric CO2 concentrations by mid-century.[210] Greenhouse gas emissions can be offset by enhancing Earth’s land carbon sink to sequester significantly larger amounts of CO2 beyond naturally occurring levels.[211] Forest preservation, reforestation and  tree planting on non-forest lands are considered the most effective, although they may present food security concerns. Soil management on croplands and grasslands is another effective mitigation technique. For all these approaches there remain large scientific uncertainties with implementing them on a global scale.[212]
+Individuals can also take actions to reduce their carbon footprint. These include: driving an EV or other energy efficient car and reducing vehicles miles by using mass transit or cycling; adopting a plant-based diet; reducing energy use in the home; limiting consumption of goods and services; and foregoing air travel.[213]
+Although there is no single pathway to limit global warming to 1.5 or 2°C,[214] most scenarios and strategies see a major increase in the use of renewable energy in combination with increased energy efficiency measures to generate the needed greenhouse gas reductions.[215] Forestry and agriculture components also include steps to reduce pressures on ecosystems and enhance their carbon sequestration capabilities.[216] Scenarios that limit global warming to 1.5°C generally project the large scale use of carbon dioxide removal methods to augment the greenhouse gas reduction approaches mentioned above.[217]
+Renewable energy would become the dominant form of electricity generation, rising to 85% or more by 2050 in some scenarios.  The use of electricity for other needs, such as heating, would rise to the point where electricity becomes the largest form of overall energy supply by 2050.[218] Investment in coal would be eliminated and coal use nearly phased out by 2050.[219]
+In transport, scenarios envision sharp increases in the market share of electric vehicles, low carbon fuel substitution for other transportation modes like shipping, and changes in transportation patterns to reduce overall demand, for example increased public transport.[220] Buildings will see additional electrification with the use of technologies like heat pumps, as well as continued energy efficiency improvements achieved via low energy building codes.[221] Industrial efforts will focus on increasing the energy efficiency of production processes, such as the use of cleaner technology for cement production,[222] designing and creating less energy intensive products, increasing product lifetimes, and developing incentives to reduce product demand.[223]
+The agriculture and forestry sector faces a triple challenge of limiting greenhouse gas emissions, preventing further conversion of forests to agricultural land, and meeting increases in world food demand.[224] A suite of actions could reduce agriculture/forestry based greenhouse gas emissions by 66% from 2010 levels by reducing growth in demand for food and other agricultural products, increasing land productivity, protecting and restoring forests, and reducing GHG emissions from agricultural production.[225]
+A wide range of policies, regulations and laws are being used to reduce greenhouse gases. Carbon pricing mechanisms include carbon taxes and emissions trading systems.[226] As of 2019, carbon pricing covers about 20% of global greenhouse gas emissions.[227] Renewable portfolio standards have been enacted in several countries to move utilities to increase the percentage of electricity they generate from renewable sources.[228] Phasing out of fossil fuel subsidies, currently estimated at $300 billion globally (about twice the level of renewable energy subsidies),[229] could reduce greenhouse gas emissions by 6%.[230] Subsidies could also be redirected to support the transition to clean energy.[231] More prescriptive methods that can reduce greenhouse gases include vehicle efficiency standards,[232] renewable fuel standards, and air pollution regulations on heavy industry.[233]
+As the use of fossil fuels is reduced, there are Just Transition considerations involving the social and economic challenges that arise. An example is the employment of workers in the affected industries, along with the well-being of the broader communities involved.[234] Climate justice considerations, such as those facing indigenous populations in the Arctic,[235] are another important aspect of mitigation policies.[236]
+Adaptation is “the process of adjustment to current or expected changes in climate and its effects”. As climate change varies across regions, adaptation does too.[237] While some adaptation responses call for trade-offs, others bring synergies and co-benefits.[238] Examples of adaptation are improved coastline protection, better disaster management, and the development of more resistant crops.[239] Increased use of air conditioning allows people to better cope with heat, but also increases energy demand.[240] Adaptation is especially important in developing countries since they are predicted to bear the brunt of the effects of global warming.[241] The capacity and potential for humans to adapt, called adaptive capacity, is unevenly distributed across different regions and populations, and developing countries generally have less capacity to adapt.[242] The public sector, private sector, and communities are all gaining experience with adaptation, and adaptation is becoming embedded within certain planning processes.[243] There are limits to adaptation and more severe climate change requires more transformative adaptation, which can be prohibitatively expensive.[237]
+Geoengineering or climate engineering is the deliberate large-scale modification of the climate to counteract climate change.[244] Techniques fall generally into the categories of solar radiation management and carbon dioxide removal, although various other schemes have been suggested. A 2018 review paper concluded that although geo-engineering is physically possible, all the techniques are in early stages of development, carry large risks and uncertainties and raise significant ethical and legal issues.[245]
+The geopolitics of climate change is complex and was often framed as a prisoners' dilemma, in which all countries benefit from mitigation done by other countries, but individual countries would lose from investing in a transition to a low-carbon economy themselves. Net importers of fossil fuels win economically from transitioning, and net exporters face stranded assets: fossil fuels they cannot sell.[246] Furthermore, the benefits to individual countries in terms of public health and local environmental improvements of coal phase out exceed the costs, potentially eliminating the free-rider problem.[247] The geopolitics may be further complicated by the supply chain of rare earth metals, which are necessary to produce clean technology.[248]
+As of 2020[update] nearly all countries in the world are parties to the United Nations Framework Convention on Climate Change (UNFCCC).[249] The objective of the Convention is to prevent dangerous human interference with the climate system.[250] As stated in the Convention, this requires that greenhouse gas concentrations are stabilized in the atmosphere at a level where ecosystems can adapt naturally to climate change, food production is not threatened, and economic development can be sustained.[251] The Framework Convention was agreed on in 1992, but global emissions have risen since then.[57] Its yearly conferences are the stage of global negotiations.[252]
+This mandate was sustained in the 1997 Kyoto Protocol to the Framework Convention.[253] In ratifying the Kyoto Protocol, most developed countries accepted legally binding commitments to limit their emissions. These first-round commitments expired in 2012.[254] United States President George W. Bush rejected the treaty on the basis that "it exempts 80% of the world, including major population centres such as China and India, from compliance, and would cause serious harm to the US economy".[255] During these negotiations, the G77 (a lobbying group in the United Nations representing developing countries)[256] pushed for a mandate requiring developed countries to "[take] the lead" in reducing their emissions.[257] This was justified on the basis that the developed countries' emissions had contributed most to the accumulation of greenhouse gases in the atmosphere, per-capita emissions were still relatively low in developing countries, and the emissions of developing countries would grow to meet their development needs.[258]
+In 2009 several UNFCCC Parties produced the Copenhagen Accord,[260] which has been widely portrayed as disappointing because of its low goals, leading poorer nations to reject it.[261] Nations associated with the Accord aimed to limit the future increase in global mean temperature to below 2 °C.[262] In 2015 all UN countries negotiated the Paris Agreement, which aims to keep climate change well below 2 °C and contains an aspirational goal of keeping warming under 1.5 °C. The agreement replaced the Kyoto Protocol. Unlike Kyoto, no binding emission targets are set in the Paris Agreement. Instead, the procedure of regularly setting ever more ambitious goals and reevaluating these goals every five years has been made binding.[263] The Paris Agreement reiterated that developing countries must be financially supported.[264] As of November 2019[update], 194 states and the European Union have signed the treaty and 186 states and the EU have ratified or acceded to the agreement.[265] In November 2019 the Trump administration notified the UN that it would withdraw the United States from the Paris Agreement in 2020.[266]
+In 2019, the British Parliament became the first national government in the world to officially declare a climate emergency.[267] Other countries and jurisdictions followed.[268] In November 2019 the European Parliament declared a "climate and environmental emergency",[269] and the European Commission presented its European Green Deal with which they hope to make the EU carbon-neutral in 2050.[270]
+While the ozone layer and climate change are considered separate problems, the solution to the former has significantly mitigated global warming. The estimated mitigation of the Montreal Protocol, an international agreement to stop emitting ozone-depleting gases, is estimated to have been more effective than the Kyoto Protocol, which was specifically designed to curb greenhouse gas emissions.[271] It has been argued that the Montreal Protocol may have done more than any other measure, as of 2017[update], to mitigate climate change as those substances were also powerful greenhouse gases.[272]
+In the scientific literature, there is an overwhelming consensus that global surface temperatures have increased in recent decades and that the trend is caused mainly by human-induced emissions of greenhouse gases.[274] No scientific body of national or international standing disagrees with this view.[275] Scientific discussion takes place in journal articles that are peer-reviewed, which scientists subject to assessment every couple of years in the Intergovernmental Panel on Climate Change reports.[276] In 2013, the IPCC Fifth Assessment Report stated that "is extremely likely that human influence has been the dominant cause of the observed warming since the mid-20th century".[277] Their 2018 report expressed the scientific consensus as: "human influence on climate has been the dominant cause of observed warming since the mid-20th century".[278]
+Consensus has further developed that some form of action should be taken to protect people against the impacts of climate change, and national science academies have called on world leaders to cut global emissions.[279] In 2017, in the second warning to humanity, 15,364 scientists from 184 countries stated that "the current trajectory of potentially catastrophic climate change due to rising greenhouse gases from burning fossil fuels, deforestation, and agricultural production – particularly from farming ruminants for meat consumption" is "especially troubling".[280] In 2019, a group of more than 11,000 scientists from 153 countries named climate change an "emergency" that would lead to "untold human suffering" if no big shifts in action takes place.[281] The emergency declaration emphasized that economic growth and population growth "are among the most important drivers of increases in CO2 emissions from fossil fuel combustion" and that "we need bold and drastic transformations regarding economic and population policies".[282]
+The global warming problem came to international public attention in the late 1980s.[283] Due to confusing media coverage in the early 1990s, issues such as ozone depletion and climate change were often mixed up, affecting public understanding of these issues.[284] Although there are a few areas of linkage, the relationship between the two is weak.[285]
+Significant regional differences exist in how concerned people are about climate change and how much they understand the issue.[286] In 2010, just a little over half the US population viewed it as a serious concern for either themselves or their families, while 73% of people in Latin America and 74% in developed Asia felt this way.[287] Similarly, in 2015 a median of 54% of respondents considered it "a very serious problem", but Americans and Chinese (whose economies are responsible for the greatest annual CO2 emissions) were among the least concerned.[286] Worldwide in 2011, people were more likely to attribute global warming to human activities than to natural causes, except in the US where nearly half of the population attributed global warming to natural causes.[288] Public reactions to global warming and concern about its effects have been increasing, with many perceiving it as the worst global threat.[289] In a 2019 CBS poll, 64% of the US population said that climate change is a "crisis" or a "serious problem", with 44% saying human activity was a significant contributor.[290]
+Public debate about climate change has been strongly affected by climate change denial and misinformation, which originated in the United States and has since spread to other countries, particularly Canada and Australia. The actors behind climate change denial form a well-funded and relatively coordinated coalition of fossil fuel companies, industry groups, conservative think tanks, and contrarian scientists.[292] Like the tobacco industry before, the main strategy of these groups has been to manufacture doubt about scientific data and results.[293] Many who deny, dismiss, or hold unwarranted doubt about the scientific consensus on anthropogenic global warming are labelled as "climate change skeptics", which several scientists have noted is a misnomer.[294]
+There are different variants of climate denial: some deny that warming takes place at all, some acknowledge warming but attribute it to natural influences, and some minimize the negative impacts of climate change.[295] Manufacturing uncertainty about the science later developed into a manufacturing of controversy: creating the belief that there remains significant uncertainty about climate change within the scientific community in order to delay policy changes.[296] Strategies to promote these ideas include a criticism of scientific institutions,[297] and questioning the motives of individual scientists.[298] An "echo chamber" of climate-denying blogs and media has further fomented misunderstanding of global warming.[299]
+Protests seeking more ambitious climate action increased in the 2010s in the form of fossil fuel divestment,[300] and worldwide demonstrations.[301] In particular, youth across the globe protested by skipping school, inspired by Swedish teenager Greta Thunberg in the school strike for climate.[302] Mass civil disobedience actions by Extinction Rebellion and Ende Gelände have ended in police intervention and large-scale arrests.[303] Litigation is increasingly used as a tool to strengthen climate action, with governments being the biggest target of lawsuits demanding that they become ambitious on climate action or enforce existing laws. Cases against fossil-fuel companies, from activists, shareholders and investors, generally seek compensation for loss and damage.[304]
+In 1681 Mariotte noted that glass, though transparent to sunlight, obstructs radiant heat.[305] Around 1774 de Saussure showed that non-luminous warm objects emit infrared heat, and used a glass-topped insulated box to trap and measure heat from sunlight.[306] In 1824 Joseph Fourier proposed by analogy a version of the greenhouse effect; transparent atmosphere lets through visible light, which warms the surface. The warmed surface emits infrared radiation, but the atmosphere is relatively opaque to infrared and slows the emission of energy, warming the planet.[307] Starting in 1859,[308] John Tyndall established that nitrogen and oxygen (99% of dry air) are transparent to infrared, but water vapour and traces of some gases (significantly methane and carbon dioxide) both absorb infrared and, when warmed, emit infrared radiation. Changing concentrations of these gases could have caused "all the mutations of climate which the researches of geologists reveal" including ice ages.[309]
+Svante Arrhenius noted that water vapour in air continuously varied, but carbon dioxide (CO2) was determined by long term geological processes. At the end of an ice age, warming from increased CO2 would increase the amount of water vapour, amplifying its effect in a feedback process. In 1896, he published the first climate model of its kind, showing that halving of CO2 could have produced the drop in temperature initiating the ice age. Arrhenius calculated the temperature increase expected from doubling CO2 to be around 5–6 °C (9.0–10.8 °F).[310] Other scientists were initially sceptical and believed the greenhouse effect to be saturated so that adding more CO2 would make no difference. Experts thought climate would be self-regulating.[311] From 1938 Guy Stewart Callendar published evidence that climate was warming and CO2 levels increasing,[312] but his calculations met the same objections.[311]
+Early calculations treated the atmosphere as a single layer: Gilbert Plass used digital computers to model the different layers and found added CO2 would cause warming. Hans Suess found evidence CO2 levels had been rising, Roger Revelle showed the oceans would not absorb the increase, and together they helped Charles Keeling to begin a record of continued increase, the Keeling Curve.[311] Scientists alerted the public,[313] and the dangers were highlighted at James Hansen's 1988 Congressional testimony.[314] The Intergovernmental Panel on Climate Change, set up in 1988 to provide formal advice to the world's governments, spurred interdisciplanary research.[315]
+Before the 1980s, when it was unclear whether warming by greenhouse gases would dominate aerosol-induced cooling, scientist often used the term ‘’inadvertent climate modification’’ to refer to humankinds’ impact on the climate. With increasing evidence of warming, the terms ‘’global warming’’ and ‘’climate change’’ were introduced, with the former referring only to increasing surface warming, and the latter to the full effect of greenhouse gases on climate.[316] Global warming became the dominant popular term after NASA climate scientist James Hansen used it in his 1988 testimony in the U.S. Senate.[314] In the 2000s, the term climate change increased in popularity.[317] Global warming is almost only used to refer to human-induced warming of the Earth system, whereas climate change is sometimes used to refer to natural as well as anthropogenic change.[318] The two terms are often used interchangeably.[319]
+Various scientists, politicians and news media have adopted the terms climate crisis or a climate emergency to talk about climate change, while using global heating instead of global warming.[320] The policy editor-in-chief of The Guardian explained why they included this language in their editorial guidelines: "We want to ensure that we are being scientifically precise, while also communicating clearly with readers on this very important issue".[321] Oxford Dictionary chose climate emergency as the word of the year 2019 and defines the term as "a situation in which urgent action is required to reduce or halt climate change and avoid potentially irreversible environmental damage resulting from it".[322]
+AR4 Working Group I Report
+AR4 Working Group II Report
+AR4 Working Group III Report
+AR4 Synthesis Report
+AR5 Working Group I Report
+AR5 Working Group II Report
+AR5 Working Group III Report
+AR5 Synthesis Report
+Special Report: SR15
+Special Report: Climate change and Land
+Special Report:  SROCC

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+The rising average temperature of Earth's climate system, called global warming, is driving changes in rainfall patterns, extreme weather, arrival of seasons, and more. Collectively, global warming and its effects are known as climate change.  While there have been prehistoric periods of global warming, observed changes since the mid-20th century have been unprecedented in rate and scale.[1]
+The Intergovernmental Panel on Climate Change (IPCC) concluded that "human influence on climate has been the dominant cause of observed warming since the mid-20th century". These findings have been recognized by the national science academies of major nations and are not disputed by any scientific body of national or international standing.[4] The largest human influence has been the emission of greenhouse gases, with over 90% of the impact from carbon dioxide and methane.[5] Fossil fuel burning is the principal source of these gases, with agricultural emissions and deforestation also playing significant roles. Temperature rise is enhanced by self-reinforcing climate feedbacks, such as loss of snow cover, increased water vapour, and melting permafrost.
+Land surfaces are heating faster than the ocean surface, leading to heat waves, wildfires, and the expansion of deserts.[6] Increasing atmospheric energy and rates of evaporation are causing more intense storms and weather extremes, damaging infrastructure and agriculture.[7] Surface temperature increases are greatest in the Arctic and have contributed to the retreat of glaciers, permafrost, and sea ice. Environmental impacts include the extinction or relocation of many species as their ecosystems change, most immediately in coral reefs, mountains, and the Arctic. Surface temperatures would stabilize and decline a little if emissions were cut off, but other impacts will continue for centuries, including rising sea levels from melting ice sheets, rising ocean temperatures, and ocean acidification from elevated levels of carbon dioxide.[8]
+Extreme weather. Drought and high temperatures worsened the 2020 bushfires in Australia.
+Habitat destruction. Many arctic animals rely on sea ice, which has been disappearing in a warming Arctic.
+Storm intensification. Bangladesh after Cyclone Sidr is an example of catastrophic flooding from increased rainfall.
+Heat wave intensification. Events like the June 2019 European heat wave are becoming more common.
+Farming. Droughts, rising temperatures, and extreme weather negatively impact agriculture.
+Tidal flooding. Sea level rise increases flooding in low lying coastal regions. Shown: Venice, Italy.
+Arctic warming. Permafrost thaws undermine infrastructure and release methane in a positive feedback loop.
+Ecological collapse possibilities. Bleaching has damaged the Great Barrier Reef and threatens reefs worldwide.
+Pest propagation. Mild winters allow more pine beetles to survive to kill large swaths of forest.
+Environmental migration. Sparser rainfall leads to desertification that harms agriculture and can displace populations.
+Mitigation efforts to address global warming include the development and deployment of low carbon energy technologies, policies to reduce fossil fuel emissions, reforestation, forest preservation, as well as the development of potential climate engineering technologies. Societies and governments are also working to adapt to current and future global warming impacts, including improved coastline protection, better disaster management, and the development of more resistant crops.
+Countries work together on climate change under the umbrella of the United Nations Framework Convention on Climate Change (UNFCCC), which has near-universal membership. The goal of the convention is to "prevent dangerous anthropogenic interference with the climate system". The IPCC has stressed the need to keep global warming below 1.5 °C (2.7 °F) compared to pre-industrial levels in order to avoid some irreversible impacts.[10] With current policies and pledges, global warming by the end of the century is expected to reach about 2.8 °C (5.0 °F).[11] At the current greenhouse gas (GHG) emission rate, the carbon budget for staying below 1.5 °C (2.7 °F) would be exhausted by 2028.[12]
+Multiple independently produced instrumental datasets show that the climate system is warming,[14] with the 2009–2018 decade being 0.93 ± 0.07 °C (1.67 ± 0.13 °F) warmer than the pre-industrial baseline (1850–1900).[15] Currently, surface temperatures are rising by about 0.2 °C (0.36 °F) per decade.[16] Since 1950, the number of cold days and nights has decreased, and the number of warm days and nights has increased.[17] Historical patterns of warming and cooling, like the Medieval Climate Anomaly and the Little Ice Age, were not as synchronous as current warming, but may have reached temperatures as high as those of the late-20th century in a limited set of regions.[18] There have been prehistorical episodes of global warming, such as the Paleocene–Eocene Thermal Maximum.[19] However, the observed rise in temperature and CO2 concentrations has been so rapid that even abrupt geophysical events that took place in Earth's history do not approach current rates.[20]
+Climate proxy records show that natural variations offset the early effects of the Industrial Revolution, so there was little net warming between the 18th century and the mid-19th century,[21] when thermometer records began to provide global coverage.[22] The Intergovernmental Panel on Climate Change (IPCC) has adopted the baseline reference period 1850–1900 as an approximation of pre-industrial global mean surface temperature.[21]
+The warming evident in the instrumental temperature record is consistent with a wide range of observations, documented by many independent scientific groups.[23] Although the most common measure of global warming is the increase in the near-surface atmospheric temperature, over 90% of the additional energy in the climate system over the last 50 years has been stored in the ocean, warming it.[24] The remainder of the additional energy has melted ice and warmed the continents and the atmosphere.[25] The ocean heat uptake drives thermal expansion which has contributed to observed sea level rise.[26] Further indicators of climate change include an increase in the frequency and intensity of heavy precipitation, melting of snow and land ice and increased atmospheric humidity.[27] Flora and fauna also portray behaviour consistent with warming, such as the earlier flowering of plants in spring.[28]
+Global warming refers to global averages, with the amount of warming varying by region. Since the pre-industrial period, global average land temperatures have increased almost twice as fast as global average temperatures.[29] This is due to the larger heat capacity of oceans and because oceans lose more heat by evaporation.[30] Patterns of warming are independent of the locations of greenhouse gas emissions because the gases persist long enough to diffuse across the planet; however, localized black carbon deposits on snow and ice do contribute to Arctic warming.[31]
+The Northern Hemisphere and North Pole have warmed much faster than the South Pole and Southern Hemisphere. The Northern Hemisphere not only has much more land, but also more snow area and sea ice, because of how the land masses are arranged around the Arctic Ocean. As these surfaces flip from being reflective to dark after the ice has melted, they start absorbing more heat. The Southern Hemisphere already had little sea ice in summer before it started warming.[32] Arctic temperatures have increased and are predicted to continue to increase during this century at over twice the rate of the rest of the world.[33] Melting of glaciers and ice sheets in the Arctic disrupts ocean circulation, uncluding a weakened Gulf Stream, causing increased warming in some areas.[34]
+Although record-breaking years attract considerable media attention, individual years are less significant than the overall global surface temperature, which is subject to short-term fluctuations that overlie long-term trends.[35] An example of such an episode is the slower rate of surface temperature increase from 1998 to 2012, which was described as the global warming hiatus.[36] Throughout this period, ocean heat storage continued to progress steadily upwards, and in subsequent years, surface temperatures have spiked upwards. The slower pace of warming can be attributed to a combination of natural fluctuations, reduced solar activity, and increased reflection sunlight of by particles from volcanic eruptions.[37]
+By itself, the climate system experiences various cycles which can last for years (such as the El Niño–Southern Oscillation) to decades or centuries.[38] Other changes are caused by an imbalance of energy at the top of the atmosphere: external forcings. These forcings are "external" to the climate system, but not always external to the Earth.[39] Examples of external forcings include changes in the composition of the atmosphere (e.g. increased concentrations of greenhouse gases), solar luminosity, volcanic eruptions, and variations in the Earth's orbit around the Sun.[40]
+Attribution of climate change is the effort to scientifically show which mechanisms are responsible for observed changes in Earth's climate. First, known internal climate variability and natural external forcings need to be ruled out. Therefore, a key approach is to use computer modelling of the climate system to determine unique "fingerprints" for all potential causes. By comparing these fingerprints with observed patterns and evolution of climate change, and the observed history of the forcings, the causes of the observed changes can be determined.[41] For example, solar forcing can be ruled out as major cause because its fingerprint is warming in the entire atmosphere, and only the lower atmosphere has warmed as expected for greenhouse gases.[42] The major causes of current climate change are primarily greenhouse gases, and secondarily land use changes, and aerosols and soot.[43]
+Greenhouse gases trap heat radiating from the Earth to space.[44] This heat, in the form of infrared radiation, gets absorbed and emitted by these gases in the atmosphere, thus warming the lower atmosphere and the surface. Before the Industrial Revolution, naturally occurring amounts of greenhouse gases caused the air near the surface to be warmer by about 33 °C (59 °F) than it would be in their absence.[45] Without the Earth's atmosphere, the Earth's average temperature would be well below the freezing temperature of water.[46] While water vapour (~50%) and clouds (~25%) are the biggest contributors to the greenhouse effect, they increase as a function of temperature and are therefore considered feedbacks. Increased concentrations of gases such as CO2 (~20%), ozone and N2O are external forcing on the other hand.[47] Ozone acts as a greenhouse gas in the lowest layer of the atmosphere, the troposphere. Furthermore, it is highly reactive and interacts with other greenhouse gases and aerosols.[48]
+Human activity since the Industrial Revolution, mainly extracting and burning fossil fuels,[49] has increased the amount of greenhouse gases in the atmosphere. This CO2, methane, tropospheric ozone, CFCs, and nitrous oxide has increased radiative forcing. In 2018, the concentrations of CO2 and methane had increased by about 45% and 160%, respectively, since pre-industrial times.[50] In 2013, CO2 readings taken at the world's primary benchmark site in Mauna Loa surpassing 400 ppm for the first time.[51] These levels are much higher than at any time during the last 800,000 years, the period for which reliable data have been collected from ice cores.[52] Less direct geological evidence indicates that CO2 values have not been this high for millions of years.[53]
+Global anthropogenic greenhouse gas emissions in 2018 excluding land use change were equivalent to 52 billion tonnes of carbon dioxide. Of these emissions, 72% was carbon dioxide from fossil fuel burning and industry, 19% was methane, largely from livestock,[54] 6% was nitrous oxide, mainly from agriculture, and 3% was fluorinated gases.[55] A further 4 billion tonnes of CO2 was released as a consequence of land use change, which is primarily due to deforestation.[56] From a production standpoint, the primary sources of global GHG emissions are estimated as: electricity and heat (25%), agriculture and forestry (24%), industry (21%), and transportation (14%).[57] Consumption based estimates of GHG emissions offer another useful way to understand sources of global warming, and may better capture the effects of trade.[58] From a consumption standpoint, the dominant sources of global 2010 emissions were found to be: food (30%), washing, heating, and lighting (26%); personal transport and freight (20%); and building construction (15%).[59]
+Despite the contribution of deforestation to GHG emissions, the Earth's land surface, particularly its forests, remain a significant carbon sink for CO2. Natural processes, such as carbon fixation in the soil and photosynthesis, more than offset the GHG contributions from deforestation. The land surface sink is estimated to remove about 11 billion tonnes of CO2 annually from the atmosphere, or about 29% of global CO2 emissions.[60] The ocean also serves as a significant carbon sink via a two-step process. First, CO2 dissolves in the surface water. Afterwards, the ocean's overturning circulation distributes it deep into the ocean's interior, where it accumulates over time as part of the carbon cycle. Over the last two decades, the world's oceans have removed between 20 and 30% of emitted CO2.[61] The strength of both the land and ocean sinks increase as CO2 levels in the atmosphere rise. In this respect they act as negative feedbacks in global warming.[62]
+Humans change the Earth's surface mainly to create more agricultural land. Today agriculture takes up 50% of the world's habitable land, while 37% is forests,[63] and that latter figure continues to decrease,[64] largely due to continued forest loss in the tropics.[65] This deforestation is the most significant aspect of land use change affecting global warming. The main causes are: deforestation through permanent land use change for agricultural products such as beef and palm oil (27%), forestry/forest products (26%), short term agricultural cultivation (24%), and wildfires (23%).[66]
+In addition to impacting greenhouse gas concentrations, land use changes affect global warming through a variety of other chemical and physical dynamics. Changing the type of vegetation in a region impacts the local temperature by changing how much sunlight gets reflected back into space, called albedo, and how much heat is lost by evaporation. For instance, the change from a dark forest to grassland makes the surface lighter, causing it to reflect more sunlight. Deforestation can also contribute to changing temperatures by affecting the release of aerosols and other chemical compounds that affect clouds; and by changing wind patterns when the land surface has different obstacles.[67] Globally, these effects are estimated to have led to a slight cooling, dominated by an increase in surface albedo.[68] But there is significant geographic variation in how this works. In the tropics the net effect is to produce a significant warming, while at latitudes closer to the poles a loss of albedo leads to an overall cooling effect.[67]
+Air pollution, in the form of aerosols, not only puts a large burden on human health, but also affects the climate on a large scale.[69] From 1961 to 1990, a gradual reduction in the amount of sunlight reaching the Earth's surface was observed, a phenomenon popularly known as global dimming,[70] typically attributed to aerosols from biofuel and fossil fuel burning.[71] Aerosol removal by precipitation gives tropospheric aerosols an atmospheric lifetime of only about a week, while stratospheric aerosols can remain in the atmosphere for a few years.[72] Globally, aerosols have been declining since 1990, removing some of the masking of global warming that they had been providing.[73]
+In addition to their direct effect by scattering and absorbing solar radiation, aerosols have indirect effects on the Earth's radiation budget. Sulfate aerosols act as cloud condensation nuclei and thus lead to clouds that have more and smaller cloud droplets. These clouds reflect solar radiation more efficiently than clouds with fewer and larger droplets.[74] This effect also causes droplets to be of more uniform size, which reduces the growth of raindrops and makes clouds more reflective to incoming sunlight.[75] Indirect effects of aerosols are the largest uncertainty in radiative forcing.[76]
+While aerosols typically limit global warming by reflecting sunlight, black carbon in soot that falls on snow or ice can contribute to global warming. Not only does this increase the absorption of sunlight, it also increases melting and sea level rise.[77] Limiting new black carbon deposits in the Arctic could reduce global warming by 0.2 °C by 2050.[78]
+As the Sun is the Earth's primary energy source, changes in incoming sunlight directly affect the climate system.[79] Solar irradiance has been measured directly by satellites,[80] and indirect measurements are available beginning in the early 1600s.[79] There has been no upward trend in the amount of the Sun's energy reaching the Earth, so it cannot be responsible for the current warming.[81] Physical climate models are also unable to reproduce the rapid warming observed in recent decades when taking into account only variations in solar output and volcanic activity.[82] Another line of evidence for the warming not being due to the Sun is how temperature changes differ at different levels in the Earth's atmosphere.[83] According to basic physical principles, the greenhouse effect produces warming of the lower atmosphere (the troposphere), but cooling of the upper atmosphere (the stratosphere).[84] If solar variations were responsible for the observed warming, warming of both the troposphere and the stratosphere would be expected, but that has not been the case.[42] Explosive volcanic eruptions represent the largest natural forcing over the industrial era. When the eruption is sufficiently strong with sulfur dioxide reaching the stratosphere, sunlight can be partially blocked for a couple of years, with a temperature signal lasting about twice as long.[85]
+The response of the climate system to an initial forcing is increased by self-reinforcing feedbacks and reduced by balancing feedbacks.[87] The main balancing feedback to global temperature change is radiative cooling to space as infrared radiation, which increases strongly with increasing temperature.[88] The main reinforcing feedbacks are the water vapour feedback, the ice–albedo feedback, and probably the net effect of clouds.[89] Uncertainty over feedbacks is the major reason why different climate models project different magnitudes of warming for a given amount of emissions.[90]
+As air gets warmer, it can hold more moisture. After an initial warming due to emissions of greenhouse gases, the atmosphere will hold more water. As water is a potent greenhouse gas, this further heats the climate: the water vapour feedback.[89] The reduction of snow cover and sea ice in the Arctic reduces the albedo of the Earth's surface.[91] More of the Sun's energy is now absorbed in these regions, contributing to Arctic amplification, which has caused Arctic temperatures to increase at more than twice the rate of the rest of the world.[92] Arctic amplification also causes methane to be released as permafrost melts, which is expected to surpass land use changes as the second strongest anthropogenic source of greenhouse gases by the end of the century.[93]
+Cloud cover may change in the future. If cloud cover increases, more sunlight will be reflected back into space, cooling the planet. Simultaneously, the clouds enhance the greenhouse effect, warming the planet. The opposite is true if cloud cover decreases. It depends on the cloud type and location which process is more important. Overall, the net feedback over the industrial era has probably been self-reinforcing.[94]
+Roughly half of each year's CO2 emissions have been absorbed by plants on land and in oceans.[95] Carbon dioxide and an extended growing season have stimulated plant growth making the land carbon cycle a balancing feedback. Climate change also increases droughts and heat waves that inhibit plant growth, which makes it uncertain whether this balancing feedback will persist in the future.[96] Soils contain large quantities of carbon and may release some when they heat up.[97] As more CO2 and heat are absorbed by the ocean, it is acidifying and ocean circulation can change, changing the rate at which the ocean can absorb atmospheric carbon.[98]
+Future warming depends on the strenght of climate feedbacks and on emissions of greenhouse gases.[99] The former is often estimated using climate models. A climate model is a representation of the physical, chemical, and biological processes that affect the climate system.[100] They also include changes in the Earth's orbit, historical changes in the Sun's activity, and volcanic forcing.[101] Computer models attempt to reproduce and predict the circulation of the oceans, the annual cycle of the seasons, and the flows of carbon between the land surface and the atmosphere.[102] There are more than two dozen scientific institutions that develop climate models.[103] Models not only project different future temperature with different emissions of greenhouse gases, but also do not fully agree on the strength of different feedbacks on climate sensitivity and the amount of inertia of the system.[104]
+The physical realism of models is tested by examining their ability to simulate contemporary or past climates.[105] Past models have underestimated the rate of Arctic shrinkage[106] and underestimated the rate of precipitation increase.[107] Sea level rise since 1990 was underestimated in older models, but now agrees well with observations.[108] The 2017 United States-published National Climate Assessment notes that "climate models may still be underestimating or missing relevant feedback processes".[109]
+Four Representative Concentration Pathways (RCPs) are used as input for climate models: "a stringent mitigation scenario (RCP2.6), two intermediate scenarios (RCP4.5 and RCP6.0) and one scenario with very high GHG [greenhouse gas] emissions (RCP8.5)".[110]  RCPs only look at concentrations of greenhouse gases, and so does not include the response of the carbon cycle.[111] Climate model projections summarized in the IPCC Fifth Assessment Report indicate that, during the 21st century, the global surface temperature is likely to rise a further 0.3 to 1.7 °C (0.5 to 3.1 °F) in a moderate scenario, or as much as 2.6 to 4.8 °C (4.7 to 8.6 °F) in an extreme scenario, depending on the rate of future greenhouse gas emissions and on climate feedback effects.[112]
+A subset of climate models add societal factors to a simple physical climate model. These models simulate how population, economic growth, and energy use affect – and interact with – the physical climate. With this information, these models can produce scenarios of how greenhouse gas emissions may vary in the future. This output is then used as input for physical climate models to generate climate change projections.[113] In some scenarios emissions continue to rise over the century, while others have reduced emissions.[114] Fossil fuel resources are abundant, and cannot be relied on to limit carbon emissions in the 21st century.[115] Emission scenarios can be combined with modelling of the carbon cycle to predict how atmospheric concentrations of greenhouse gases might change in the future.[116] According to these combined models, by 2100 the atmospheric concentration of CO2 could be as low as 380 or as high as 1400 ppm, depending on the Shared Socioeconomic Pathway (SSP) and the mitigation scenario.[117]
+The remaining carbon emissions budget is determined from modelling the carbon cycle and climate sensitivity to greenhouse gases.[118] According to the IPCC, global warming can be kept below 1.5 °C with a two-thirds chance if emissions after 2018 do not exceed 420 or 570 GtCO2 depending on the choice of the measure of global temperature. This amount corresponds to 10 to 13 years of current emissions. There are high uncertainties about the budget; for instance, it may be 100 GtCO2 smaller due to methane release from permafrost and wetlands.[119]
+The environmental effects of global warming are broad and far-reaching. They include effects on the oceans, ice, and weather and may occur gradually or rapidly. Evidence for these effects come from studying climate change in the past, modelling and modern observations.[121] Since the 1950s, droughts and heat waves have appeared simultaneously with increasing frequency.[122] Extremely wet or dry events within the monsoon period have increased in India and East Asia.[123] Various mechanisms have been identified that might explain extreme weather in mid-latitudes from the rapidly warming Arctic, such as the jet stream becoming more erratic.[124] The maximum rainfall and wind speed from hurricanes and typhoons is likely increasing.[125]
+Between 1993 and 2017, the global mean sea level rose on average by 3.1 ± 0.3 mm per year, with an acceleration detected as well.[126] Over the 21st century, the IPCC projects that in a very high emissions scenario the sea level could rise by 61–110 cm.[127] The rate of ice loss from glaciers and ice sheets in the Antarctic is a key area of uncertainty since this source could account for 90% of the potential sea level rise:[128] increased ocean warmth is undermining and threatening to unplug Antarctic glacier outlets, potentially resulting in more rapid sea level rise.[129] The retreat of non-polar glaciers also contributes to sea level rise.[130]
+Global warming has led to decades of shrinking and thinning of the Arctic sea ice, making it vulnerable to atmospheric anomalies.[131] Projections of declines in Arctic sea ice vary.[132] While ice-free summers are expected to be rare at 1.5 °C (2.7 °F) degrees of warming, they are set to occur once every three to ten years at a warming level of 2.0 °C (3.6 °F),[133] increasing the ice–albedo feedback.[134] Higher atmospheric CO2 concentrations have led to an increase in dissolved CO2, which causes ocean acidification.[135] Furthermore, oxygen levels decrease because oxygen is less soluble in warmer water, an effect known as ocean deoxygenation.[136]
+The long-term effects of global warming include further ice melt, ocean warming, sea level rise, and ocean acidification. On the timescale of centuries to millennia, the magnitude of global warming will be determined primarily by anthropogenic CO2 emissions.[137] This is due to carbon dioxide's very long lifetime in the atmosphere.[137] Carbon dioxide is slowly taking up by the ocean, such that ocean acidification will continue for hundreds to thousands of years.[138] The emissions are estimated to have prolonged the current interglacial period by at least 100,000 years.[139] Because the great mass of glaciers and ice caps depressed the Earth's crust, another long-term effect of ice melt and deglaciation is the gradual rising of landmasses, a process called post-glacial rebound.[140] Sea level rise will continue over many centuries, with an estimated rise of 2.3 metres per degree Celsius (4.2 ft/°F) after 2000 years.[141]
+If global warming exceeds 1.5 °C, there is a greater risk of passing through ‘tipping points’, thresholds beyond which certain impacts can no longer be avoided even if temperatures are reduced.[142] Some large-scale changes could occur abruptly, i.e. over a short time period. One potential source of abrupt tipping would be the rapid release of methane and carbon dioxide from permafrost, which would amplify global warming.[143] Another example is the possibility for the Atlantic Meridional Overturning Circulation to collapse,[144] which could trigger cooling in the North Atlantic, Europe, and North America.[145] If multiple temperature and carbon cycle tipping points re-inforce each other, or if there were to be strong threshold behaviour in cloud cover, there could be a global tipping into a hothouse Earth.[146]
+Recent warming has driven many terrestrial and freshwater species poleward and towards higher altitudes.[147] Higher atmospheric CO2 levels and an extended growing season have resulted in global greening, whereas heatwaves and drought have reduced ecosystem productivity in some regions. The future balance of these opposing effects is unclear.[148] Global warming has contributed to the expansion of drier climatic zones, such as, probably, the expansion of deserts in the subtropics.[149] Without substantial actions to reduce the rate of global warming, land-based ecosystems risk major shifts in their composition and structure.[150] Overall, it is expected that climate change will result in the extinction of many species and reduced diversity of ecosystems.[151]
+The ocean has heated more slowly than the land, but plants and animals in the ocean have migrated towards the colder poles as fast as or faster than species on land.[152] Just as on land, heat waves in the ocean occur more due to climate change, with harmful effects found on a wide range of organisms such as corals, kelp, and seabirds.[153] Ocean acidification threatens damage to coral reefs, fisheries, protected species, and other natural resources of value to society.[154] Coastal ecosystems are under stress, with almost half of wetlands having disappeared as a consquence of climate change and other human impacts. Harmful algae blooms have increased due to warming, ocean deoxygenation and eutrophication.[155]
+Ecological collapse possibilities. Bleaching has damaged the Great Barrier Reef and threatens reefs worldwide.[156]
+Extreme weather. Drought and high temperatures worsened the 2020 bushfires in Australia.[157]
+Arctic warming. Permafrost thaws undermine infrastructure and release methane in a positive feedback loop.[143]
+Habitat destruction. Many arctic animals rely on sea ice, which has been disappearing in a warming Arctic.[158]
+Pest propagation. Mild winters allow more pine beetles to survive to kill large swaths of forest.[159]
+The effects of climate change on human systems, mostly due to warming and shifts in precipitation, have been detected worldwide. The social impacts of climate change will be uneven across the world.[160] All regions are at risk of experiencing negative impacts,[161] with low-latitude, less developed areas facing the greatest risk.[162] Global warming has likely already increased global economic inequality, and is projected to do so in the future.[163] Regional impacts of climate change are now observable on all continents and across ocean regions.[164] The Arctic, Africa, small islands, and Asian megadeltas are regions that are likely to be especially affected by future climate change.[165] Many risks increase with higher magnitudes of global warming.[166]
+Crop production will probably be negatively affected in low-latitude countries, while effects at northern latitudes may be positive or negative.[167] Global warming of around 4 °C relative to late 20th century levels could pose a large risk to global and regional food security.[168] The impact of climate change on crop productivity for the four major crops was negative for wheat and maize, and neutral for soy and rice, in the years 1960–2013.[169] Up to an additional 183 million people worldwide, particularly those with lower incomes, are at risk of hunger as a consequence of warming.[170] While increased CO2 levels help crop growth at lower temperature increases, those crops do become less nutritious.[170] Based on local and indigenous knowledge, climate change is already affecting food security in mountain regions in South America and Asia, and in various drylands, particularly in Africa.[170] Regions dependent on glacier water, regions that are already dry, and small islands are also at increased risk of water stress due to climate change.[171]
+In small islands and mega deltas, inundation from sea level rise is expected to threaten vital infrastructure and human settlements.[172] This could lead to homelessness in countries with low-lying areas such as Bangladesh, as well as statelessness for populations in island nations, such as the Maldives and Tuvalu.[173] Climate change can be an important driver of migration, both within and between countries.[174]
+The majority of severe impacts of climate change are expected in sub-Saharan Africa and South-East Asia, where existing poverty is exacerbated.[175] The World Bank estimates that global warming could drive over 120 million people into poverty by 2030.[176] Current inequalities between men and women, between rich and poor and between people of different ethnicity have been observed to worsen as a consequence of climate variability and climate change.[177] Existing stresses include poverty, political conflicts, and ecosystem degradation. Regions may even become uninhabitable, with humidity and temperatures reaching levels too high for humans to survive.[178]
+Generally, impacts on public health will be more negative than positive.[179] Impacts include the direct effects of extreme weather, leading to injury and loss of life;[180] and indirect effects, such as undernutrition brought on by crop failures.[181] Various infectious diseases are more easily transmitted in a warming climate, such as dengue fever, which affects children most severely, and malaria.[182] Young children are further the most vulnerable to food shortages, and together with older people to extreme heat.[183] Climate change has been linked to an increase in violent conflict by amplifying poverty and economic shocks, which are well-documented drivers of these conflicts.[184] Links have been made between a wide range of violent behaviour including, violent crimes, civil unrest, and wars, but conclusive scientific evidence remains elusive.[185]
+Environmental migration. Sparser rainfall leads to desertification that harms agriculture and can displace populations.[186]
+Farming. Droughts, rising temperatures, and extreme weather negatively impact agriculture.[187]
+Tidal flooding. Sea level rise increases flooding in low lying coastal regions. Shown: Venice, Italy.[188]
+Storm intensification. Bangladesh after Cyclone Sidr is an example of catastrophic flooding from increased rainfall.[189]
+Heat wave intensification. Events like the June 2019 European heat wave are becoming more common.[190]
+Mitigation of and adaptation to climate change are two complementary responses to global warming. Successful adaptation is easier if there are substantial emission reductions. Many of the countries that have contributed least to global greenhouse gas emissions are among the most vulnerable to climate change, which raises questions about justice and fairness with regard to mitigation and adaptation.[192]
+Climate change impacts can be mitigated by reducing greenhouse gas emissions and by enhancing the capacity of Earth's surface to absorb greenhouse gases from the atmosphere.[193]
+In order to limit global warming to less than 1.5°C with a high likelihood of success, the IPCC estimates that global GHG emissions will need to be net zero by 2050,[194] or by 2070 with a 2°C target. This will require far-reaching, systemIc changes on an unprecedented scale in energy, land, cities, transport, buildings, and industry.[195] To make progress towards that goal, the United Nations Environment Programme estimates that, within the next decade, countries will need to triple the amount of reductions they have committed to in their current Paris agreements.[196]
+Long-term scenarios all point to rapid and significant investment in renewable energy and energy efficiency as key to reducing GHG emissions.[197] These technologies include solar and wind power, bioenergy, geothermal energy, and hydroelectricity. Combined, they are capable of supplying several times the world’s current energy needs.[198] Solar PV and wind, in particular, have seen substantial growth and progress over the last few years,[199] such that they are currently among the cheapest sources of new power generation.[200] Renewables represented 75% of all new electricity generation installed in 2019, with solar and wind constituting nearly all of that amount.[201] However, fossil fuels continue to dominate world energy supplies. In 2018 fossil fuels produced 80% of the world’s energy, with modern renewable sources, including solar and wind power, accounting for around 11%.[202]
+There are obstacles to the rapid development of renewable energy. Environmental and land use concerns are sometimes associated with large solar, wind and hydropower projects.[203] Solar and wind power also require energy storage systems and other modifications to the electricity grid to operate effectively,[204] although several storage technologies are now emerging to supplement the traditional use of  pumped-storage hydropower.[205] The use of rare metals and other hazardous materials has also been raised as a concern with solar power.[206] The use of bioenergy is often not carbon neutral, and may have negative consequences for food security,[207] largely due to the amount of land required compared to other renewable energy options.[208]
+For certain energy supply needs, as well as specific CO2-intensive heavy industries, carbon capture and storage may be a viable method of reducing CO2 emissions. Although high costs have been a concern with this technology,[209] it may be able to play a significant role in limiting atmospheric CO2 concentrations by mid-century.[210] Greenhouse gas emissions can be offset by enhancing Earth’s land carbon sink to sequester significantly larger amounts of CO2 beyond naturally occurring levels.[211] Forest preservation, reforestation and  tree planting on non-forest lands are considered the most effective, although they may present food security concerns. Soil management on croplands and grasslands is another effective mitigation technique. For all these approaches there remain large scientific uncertainties with implementing them on a global scale.[212]
+Individuals can also take actions to reduce their carbon footprint. These include: driving an EV or other energy efficient car and reducing vehicles miles by using mass transit or cycling; adopting a plant-based diet; reducing energy use in the home; limiting consumption of goods and services; and foregoing air travel.[213]
+Although there is no single pathway to limit global warming to 1.5 or 2°C,[214] most scenarios and strategies see a major increase in the use of renewable energy in combination with increased energy efficiency measures to generate the needed greenhouse gas reductions.[215] Forestry and agriculture components also include steps to reduce pressures on ecosystems and enhance their carbon sequestration capabilities.[216] Scenarios that limit global warming to 1.5°C generally project the large scale use of carbon dioxide removal methods to augment the greenhouse gas reduction approaches mentioned above.[217]
+Renewable energy would become the dominant form of electricity generation, rising to 85% or more by 2050 in some scenarios.  The use of electricity for other needs, such as heating, would rise to the point where electricity becomes the largest form of overall energy supply by 2050.[218] Investment in coal would be eliminated and coal use nearly phased out by 2050.[219]
+In transport, scenarios envision sharp increases in the market share of electric vehicles, low carbon fuel substitution for other transportation modes like shipping, and changes in transportation patterns to reduce overall demand, for example increased public transport.[220] Buildings will see additional electrification with the use of technologies like heat pumps, as well as continued energy efficiency improvements achieved via low energy building codes.[221] Industrial efforts will focus on increasing the energy efficiency of production processes, such as the use of cleaner technology for cement production,[222] designing and creating less energy intensive products, increasing product lifetimes, and developing incentives to reduce product demand.[223]
+The agriculture and forestry sector faces a triple challenge of limiting greenhouse gas emissions, preventing further conversion of forests to agricultural land, and meeting increases in world food demand.[224] A suite of actions could reduce agriculture/forestry based greenhouse gas emissions by 66% from 2010 levels by reducing growth in demand for food and other agricultural products, increasing land productivity, protecting and restoring forests, and reducing GHG emissions from agricultural production.[225]
+A wide range of policies, regulations and laws are being used to reduce greenhouse gases. Carbon pricing mechanisms include carbon taxes and emissions trading systems.[226] As of 2019, carbon pricing covers about 20% of global greenhouse gas emissions.[227] Renewable portfolio standards have been enacted in several countries to move utilities to increase the percentage of electricity they generate from renewable sources.[228] Phasing out of fossil fuel subsidies, currently estimated at $300 billion globally (about twice the level of renewable energy subsidies),[229] could reduce greenhouse gas emissions by 6%.[230] Subsidies could also be redirected to support the transition to clean energy.[231] More prescriptive methods that can reduce greenhouse gases include vehicle efficiency standards,[232] renewable fuel standards, and air pollution regulations on heavy industry.[233]
+As the use of fossil fuels is reduced, there are Just Transition considerations involving the social and economic challenges that arise. An example is the employment of workers in the affected industries, along with the well-being of the broader communities involved.[234] Climate justice considerations, such as those facing indigenous populations in the Arctic,[235] are another important aspect of mitigation policies.[236]
+Adaptation is “the process of adjustment to current or expected changes in climate and its effects”. As climate change varies across regions, adaptation does too.[237] While some adaptation responses call for trade-offs, others bring synergies and co-benefits.[238] Examples of adaptation are improved coastline protection, better disaster management, and the development of more resistant crops.[239] Increased use of air conditioning allows people to better cope with heat, but also increases energy demand.[240] Adaptation is especially important in developing countries since they are predicted to bear the brunt of the effects of global warming.[241] The capacity and potential for humans to adapt, called adaptive capacity, is unevenly distributed across different regions and populations, and developing countries generally have less capacity to adapt.[242] The public sector, private sector, and communities are all gaining experience with adaptation, and adaptation is becoming embedded within certain planning processes.[243] There are limits to adaptation and more severe climate change requires more transformative adaptation, which can be prohibitatively expensive.[237]
+Geoengineering or climate engineering is the deliberate large-scale modification of the climate to counteract climate change.[244] Techniques fall generally into the categories of solar radiation management and carbon dioxide removal, although various other schemes have been suggested. A 2018 review paper concluded that although geo-engineering is physically possible, all the techniques are in early stages of development, carry large risks and uncertainties and raise significant ethical and legal issues.[245]
+The geopolitics of climate change is complex and was often framed as a prisoners' dilemma, in which all countries benefit from mitigation done by other countries, but individual countries would lose from investing in a transition to a low-carbon economy themselves. Net importers of fossil fuels win economically from transitioning, and net exporters face stranded assets: fossil fuels they cannot sell.[246] Furthermore, the benefits to individual countries in terms of public health and local environmental improvements of coal phase out exceed the costs, potentially eliminating the free-rider problem.[247] The geopolitics may be further complicated by the supply chain of rare earth metals, which are necessary to produce clean technology.[248]
+As of 2020[update] nearly all countries in the world are parties to the United Nations Framework Convention on Climate Change (UNFCCC).[249] The objective of the Convention is to prevent dangerous human interference with the climate system.[250] As stated in the Convention, this requires that greenhouse gas concentrations are stabilized in the atmosphere at a level where ecosystems can adapt naturally to climate change, food production is not threatened, and economic development can be sustained.[251] The Framework Convention was agreed on in 1992, but global emissions have risen since then.[57] Its yearly conferences are the stage of global negotiations.[252]
+This mandate was sustained in the 1997 Kyoto Protocol to the Framework Convention.[253] In ratifying the Kyoto Protocol, most developed countries accepted legally binding commitments to limit their emissions. These first-round commitments expired in 2012.[254] United States President George W. Bush rejected the treaty on the basis that "it exempts 80% of the world, including major population centres such as China and India, from compliance, and would cause serious harm to the US economy".[255] During these negotiations, the G77 (a lobbying group in the United Nations representing developing countries)[256] pushed for a mandate requiring developed countries to "[take] the lead" in reducing their emissions.[257] This was justified on the basis that the developed countries' emissions had contributed most to the accumulation of greenhouse gases in the atmosphere, per-capita emissions were still relatively low in developing countries, and the emissions of developing countries would grow to meet their development needs.[258]
+In 2009 several UNFCCC Parties produced the Copenhagen Accord,[260] which has been widely portrayed as disappointing because of its low goals, leading poorer nations to reject it.[261] Nations associated with the Accord aimed to limit the future increase in global mean temperature to below 2 °C.[262] In 2015 all UN countries negotiated the Paris Agreement, which aims to keep climate change well below 2 °C and contains an aspirational goal of keeping warming under 1.5 °C. The agreement replaced the Kyoto Protocol. Unlike Kyoto, no binding emission targets are set in the Paris Agreement. Instead, the procedure of regularly setting ever more ambitious goals and reevaluating these goals every five years has been made binding.[263] The Paris Agreement reiterated that developing countries must be financially supported.[264] As of November 2019[update], 194 states and the European Union have signed the treaty and 186 states and the EU have ratified or acceded to the agreement.[265] In November 2019 the Trump administration notified the UN that it would withdraw the United States from the Paris Agreement in 2020.[266]
+In 2019, the British Parliament became the first national government in the world to officially declare a climate emergency.[267] Other countries and jurisdictions followed.[268] In November 2019 the European Parliament declared a "climate and environmental emergency",[269] and the European Commission presented its European Green Deal with which they hope to make the EU carbon-neutral in 2050.[270]
+While the ozone layer and climate change are considered separate problems, the solution to the former has significantly mitigated global warming. The estimated mitigation of the Montreal Protocol, an international agreement to stop emitting ozone-depleting gases, is estimated to have been more effective than the Kyoto Protocol, which was specifically designed to curb greenhouse gas emissions.[271] It has been argued that the Montreal Protocol may have done more than any other measure, as of 2017[update], to mitigate climate change as those substances were also powerful greenhouse gases.[272]
+In the scientific literature, there is an overwhelming consensus that global surface temperatures have increased in recent decades and that the trend is caused mainly by human-induced emissions of greenhouse gases.[274] No scientific body of national or international standing disagrees with this view.[275] Scientific discussion takes place in journal articles that are peer-reviewed, which scientists subject to assessment every couple of years in the Intergovernmental Panel on Climate Change reports.[276] In 2013, the IPCC Fifth Assessment Report stated that "is extremely likely that human influence has been the dominant cause of the observed warming since the mid-20th century".[277] Their 2018 report expressed the scientific consensus as: "human influence on climate has been the dominant cause of observed warming since the mid-20th century".[278]
+Consensus has further developed that some form of action should be taken to protect people against the impacts of climate change, and national science academies have called on world leaders to cut global emissions.[279] In 2017, in the second warning to humanity, 15,364 scientists from 184 countries stated that "the current trajectory of potentially catastrophic climate change due to rising greenhouse gases from burning fossil fuels, deforestation, and agricultural production – particularly from farming ruminants for meat consumption" is "especially troubling".[280] In 2019, a group of more than 11,000 scientists from 153 countries named climate change an "emergency" that would lead to "untold human suffering" if no big shifts in action takes place.[281] The emergency declaration emphasized that economic growth and population growth "are among the most important drivers of increases in CO2 emissions from fossil fuel combustion" and that "we need bold and drastic transformations regarding economic and population policies".[282]
+The global warming problem came to international public attention in the late 1980s.[283] Due to confusing media coverage in the early 1990s, issues such as ozone depletion and climate change were often mixed up, affecting public understanding of these issues.[284] Although there are a few areas of linkage, the relationship between the two is weak.[285]
+Significant regional differences exist in how concerned people are about climate change and how much they understand the issue.[286] In 2010, just a little over half the US population viewed it as a serious concern for either themselves or their families, while 73% of people in Latin America and 74% in developed Asia felt this way.[287] Similarly, in 2015 a median of 54% of respondents considered it "a very serious problem", but Americans and Chinese (whose economies are responsible for the greatest annual CO2 emissions) were among the least concerned.[286] Worldwide in 2011, people were more likely to attribute global warming to human activities than to natural causes, except in the US where nearly half of the population attributed global warming to natural causes.[288] Public reactions to global warming and concern about its effects have been increasing, with many perceiving it as the worst global threat.[289] In a 2019 CBS poll, 64% of the US population said that climate change is a "crisis" or a "serious problem", with 44% saying human activity was a significant contributor.[290]
+Public debate about climate change has been strongly affected by climate change denial and misinformation, which originated in the United States and has since spread to other countries, particularly Canada and Australia. The actors behind climate change denial form a well-funded and relatively coordinated coalition of fossil fuel companies, industry groups, conservative think tanks, and contrarian scientists.[292] Like the tobacco industry before, the main strategy of these groups has been to manufacture doubt about scientific data and results.[293] Many who deny, dismiss, or hold unwarranted doubt about the scientific consensus on anthropogenic global warming are labelled as "climate change skeptics", which several scientists have noted is a misnomer.[294]
+There are different variants of climate denial: some deny that warming takes place at all, some acknowledge warming but attribute it to natural influences, and some minimize the negative impacts of climate change.[295] Manufacturing uncertainty about the science later developed into a manufacturing of controversy: creating the belief that there remains significant uncertainty about climate change within the scientific community in order to delay policy changes.[296] Strategies to promote these ideas include a criticism of scientific institutions,[297] and questioning the motives of individual scientists.[298] An "echo chamber" of climate-denying blogs and media has further fomented misunderstanding of global warming.[299]
+Protests seeking more ambitious climate action increased in the 2010s in the form of fossil fuel divestment,[300] and worldwide demonstrations.[301] In particular, youth across the globe protested by skipping school, inspired by Swedish teenager Greta Thunberg in the school strike for climate.[302] Mass civil disobedience actions by Extinction Rebellion and Ende Gelände have ended in police intervention and large-scale arrests.[303] Litigation is increasingly used as a tool to strengthen climate action, with governments being the biggest target of lawsuits demanding that they become ambitious on climate action or enforce existing laws. Cases against fossil-fuel companies, from activists, shareholders and investors, generally seek compensation for loss and damage.[304]
+In 1681 Mariotte noted that glass, though transparent to sunlight, obstructs radiant heat.[305] Around 1774 de Saussure showed that non-luminous warm objects emit infrared heat, and used a glass-topped insulated box to trap and measure heat from sunlight.[306] In 1824 Joseph Fourier proposed by analogy a version of the greenhouse effect; transparent atmosphere lets through visible light, which warms the surface. The warmed surface emits infrared radiation, but the atmosphere is relatively opaque to infrared and slows the emission of energy, warming the planet.[307] Starting in 1859,[308] John Tyndall established that nitrogen and oxygen (99% of dry air) are transparent to infrared, but water vapour and traces of some gases (significantly methane and carbon dioxide) both absorb infrared and, when warmed, emit infrared radiation. Changing concentrations of these gases could have caused "all the mutations of climate which the researches of geologists reveal" including ice ages.[309]
+Svante Arrhenius noted that water vapour in air continuously varied, but carbon dioxide (CO2) was determined by long term geological processes. At the end of an ice age, warming from increased CO2 would increase the amount of water vapour, amplifying its effect in a feedback process. In 1896, he published the first climate model of its kind, showing that halving of CO2 could have produced the drop in temperature initiating the ice age. Arrhenius calculated the temperature increase expected from doubling CO2 to be around 5–6 °C (9.0–10.8 °F).[310] Other scientists were initially sceptical and believed the greenhouse effect to be saturated so that adding more CO2 would make no difference. Experts thought climate would be self-regulating.[311] From 1938 Guy Stewart Callendar published evidence that climate was warming and CO2 levels increasing,[312] but his calculations met the same objections.[311]
+Early calculations treated the atmosphere as a single layer: Gilbert Plass used digital computers to model the different layers and found added CO2 would cause warming. Hans Suess found evidence CO2 levels had been rising, Roger Revelle showed the oceans would not absorb the increase, and together they helped Charles Keeling to begin a record of continued increase, the Keeling Curve.[311] Scientists alerted the public,[313] and the dangers were highlighted at James Hansen's 1988 Congressional testimony.[314] The Intergovernmental Panel on Climate Change, set up in 1988 to provide formal advice to the world's governments, spurred interdisciplanary research.[315]
+Before the 1980s, when it was unclear whether warming by greenhouse gases would dominate aerosol-induced cooling, scientist often used the term ‘’inadvertent climate modification’’ to refer to humankinds’ impact on the climate. With increasing evidence of warming, the terms ‘’global warming’’ and ‘’climate change’’ were introduced, with the former referring only to increasing surface warming, and the latter to the full effect of greenhouse gases on climate.[316] Global warming became the dominant popular term after NASA climate scientist James Hansen used it in his 1988 testimony in the U.S. Senate.[314] In the 2000s, the term climate change increased in popularity.[317] Global warming is almost only used to refer to human-induced warming of the Earth system, whereas climate change is sometimes used to refer to natural as well as anthropogenic change.[318] The two terms are often used interchangeably.[319]
+Various scientists, politicians and news media have adopted the terms climate crisis or a climate emergency to talk about climate change, while using global heating instead of global warming.[320] The policy editor-in-chief of The Guardian explained why they included this language in their editorial guidelines: "We want to ensure that we are being scientifically precise, while also communicating clearly with readers on this very important issue".[321] Oxford Dictionary chose climate emergency as the word of the year 2019 and defines the term as "a situation in which urgent action is required to reduce or halt climate change and avoid potentially irreversible environmental damage resulting from it".[322]
+AR4 Working Group I Report
+AR4 Working Group II Report
+AR4 Working Group III Report
+AR4 Synthesis Report
+AR5 Working Group I Report
+AR5 Working Group II Report
+AR5 Working Group III Report
+AR5 Synthesis Report
+Special Report: SR15
+Special Report: Climate change and Land
+Special Report:  SROCC

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+Mesothelae
+Opisthothelae
+ See Spider taxonomy.
+Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom,[2] and spinnerets that extrude silk.[3] They are the largest order of arachnids and rank seventh in total species diversity among all orders of organisms.[4] Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of July 2019[update], at least 48,200 spider species, and 120 families have been recorded by taxonomists.[1] However, there has been dissension within the scientific community as to how all these families should be classified, as evidenced by the over 20 different classifications that have been proposed since 1900.[5]
+Anatomically, spiders (as with all arachnids) differ from other arthropods in that the usual body segments are fused into two tagmata, the prosoma, or cephalothorax, and opisthosoma, or abdomen, and joined by a small, cylindrical pedicel (however, as there is currently neither paleontological nor embryological evidence that spiders ever had a separate thorax-like division, there exists an argument against the validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly, arguments can be formed against use of the term abdomen, as the opisthosoma of all spiders contains a heart and respiratory organs, organs atypical of an abdomen[6]). Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.
+Their abdomens bear appendages that have been modified into spinnerets that extrude silk from up to six types of glands. Spider webs vary widely in size, shape and the amount of sticky thread used. It now appears that the spiral orb web may be one of the earliest forms, and spiders that produce tangled cobwebs are more abundant and diverse than orb-web spiders. Spider-like arachnids with silk-producing spigots appeared in the Devonian period about 386 million years ago, but these animals apparently lacked spinnerets. True spiders have been found in Carboniferous rocks from 318 to 299 million years ago, and are very similar to the most primitive surviving suborder, the Mesothelae. The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, before 200 million years ago.
+The species Bagheera kiplingi was described as herbivorous in 2008,[7] but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. It is estimated that the world's 25 million tons of spiders kill 400–800 million tons of prey per year.[8] Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, so they liquefy their food by flooding it with digestive enzymes. They also grind food with the bases of their pedipalps, as arachnids do not have the mandibles that crustaceans and insects have.
+To avoid being eaten by the females, which are typically much larger, male spiders identify themselves to potential mates by a variety of complex courtship rituals. Males of most species survive a few matings, limited mainly by their short life spans. Females weave silk egg-cases, each of which may contain hundreds of eggs. Females of many species care for their young, for example by carrying them around or by sharing food with them. A minority of species are social, building communal webs that may house anywhere from a few to 50,000 individuals. Social behavior ranges from precarious toleration, as in the widow spiders, to co-operative hunting and food-sharing. Although most spiders live for at most two years, tarantulas and other mygalomorph spiders can live up to 25 years in captivity.
+While the venom of a few species is dangerous to humans, scientists are now researching the use of spider venom in medicine and as non-polluting pesticides. Spider silk provides a combination of lightness, strength and elasticity that is superior to that of synthetic materials, and spider silk genes have been inserted into mammals and plants to see if these can be used as silk factories. As a result of their wide range of behaviors, spiders have become common symbols in art and mythology symbolizing various combinations of patience, cruelty and creative powers. An abnormal fear of spiders is called arachnophobia.
+1: pedipalp
+ 2: trichobothria
+ 3: carapace of prosoma (cephalothorax)
+ 4: opisthosoma (abdomen)
+ 5: eyes – AL (anterior lateral)
+    AM (anterior median)
+    PL (posterior lateral)
+    PM (posterior median)
+Leg segments:
+ 6: costa
+ 7: trochanter
+ 8: patella
+ 9: tibia
+10: metatarsus
+11: tarsus
+13: claw
+14: chelicera
+15: sternum of prosoma
+16: pedicel (also called pedicle)
+17: book lung sac
+18: book lung stigma
+19: epigastric fold
+20: epigyne
+21: anterior spinneret
+22: posterior spinneret
+Spiders are chelicerates and therefore arthropods.[9] As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo.[10] Being chelicerates, their bodies consist of two tagmata, sets of segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete fusion of the segments that in an insect would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma.[9] In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel.[11] The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws".[10][12] The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.[9]
+Spiders and scorpions are members of one chelicerate group, the arachnids.[12] Scorpions' chelicerae have three sections and are used in feeding.[13] Spiders' chelicerae have two sections and terminate in fangs that are generally venomous, and fold away behind the upper sections while not in use. The upper sections generally have thick "beards" that filter solid lumps out of their food, as spiders can take only liquid food.[11] Scorpions' pedipalps generally form large claws for capturing prey,[13] while those of spiders are fairly small appendages whose bases also act as an extension of the mouth; in addition, those of male spiders have enlarged last sections used for sperm transfer.[11]
+In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex carapace, while the underside is covered by two rather flat plates. The abdomen is soft and egg-shaped. It shows no sign of segmentation, except that the primitive Mesothelae, whose living members are the Liphistiidae, have segmented plates on the upper surface.[11]
+Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end.[14] However, in spiders, it occupies only the upper part of the abdomen, and blood is discharged into the hemocoel by one artery that opens at the rear end of the abdomen and by branching arteries that pass through the pedicle and open into several parts of the cephalothorax. Hence spiders have open circulatory systems.[11] The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.[12]
+Spiders have developed several different respiratory anatomies, based on book lungs, a tracheal system, or both. Mygalomorph and Mesothelae spiders have two pairs of book lungs filled with haemolymph, where openings on the ventral surface of the abdomen allow air to enter and diffuse oxygen. This is also the case for some basal araneomorph spiders, like the family Hypochilidae, but the remaining members of this group have just the anterior pair of book lungs intact while the posterior pair of breathing organs are partly or fully modified into tracheae, through which oxygen is diffused into the haemolymph or directly to the tissue and organs.[11] The tracheal system has most likely evolved in small ancestors to help resist desiccation.[12] The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets.[11] Spiders that have tracheae generally have higher metabolic rates and better water conservation.[15] Spiders are ectotherms, so environmental temperatures affect their activity.[16]
+Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae.[11] The families Uloboridae and Holarchaeidae, and some Liphistiidae spiders, have lost their venom glands, and kill their prey with silk instead.[17] Like most arachnids, including scorpions,[12] spiders have a narrow gut that can only cope with liquid food and two sets of filters to keep solids out.[11] They use one of two different systems of external digestion. Some pump digestive enzymes from the midgut into the prey and then suck the liquified tissues of the prey into the gut, eventually leaving behind the empty husk of the prey. Others grind the prey to pulp using the chelicerae and the bases of the pedipalps, while flooding it with enzymes; in these species, the chelicerae and the bases of the pedipalps form a preoral cavity that holds the food they are processing.[11]
+The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The midgut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.[11]
+Most spiders convert nitrogenous waste products into uric acid, which can be excreted as a dry material. Malphigian tubules ("little tubes") extract these wastes from the blood in the hemocoel and dump them into the cloacal chamber, from which they are expelled through the anus.[11] Production of uric acid and its removal via Malphigian tubules are a water-conserving feature that has evolved independently in several arthropod lineages that can live far away from water,[18] for example the tubules of insects and arachnids develop from completely different parts of the embryo.[12] However, a few primitive spiders, the suborder Mesothelae and infraorder Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"),[11] which use large amounts of water to excrete nitrogenous waste products as ammonia.[18]
+The basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia.[19] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen;[11][12][19] in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.[15]
+Despite the relatively small central nervous system, some spiders (like Portia) exhibit complex behaviour, including the ability to use a trial-and-error approach.[20][21]
+Spiders have primarily four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another.[11] The principal pair at the front are of the type called pigment-cup ocelli ("little eyes"), which in most arthropods are only capable of detecting the direction from which light is coming, using the shadow cast by the walls of the cup. However, in spiders these eyes are capable of forming images.[22][23] The other pairs, called secondary eyes, are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the principal eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torchlight reflected from the tapeta. On the other hand, jumping spiders' secondary eyes have no tapeta.[11]
+Other differences between the principal and secondary eyes are that the latter have rhabdomeres that point away from incoming light, just like in vertebrates, while the arrangement is the opposite in the former. The principal eyes are also the only ones with eye muscles, allowing them to move the retina. Having no muscles, the secondary eyes are immobile.[24]
+Some jumping spiders' visual acuity exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects; in fact the human eye is only about five times sharper than a jumping spider's. They achieve this by a telephotographic series of lenses, a four-layer retina and the ability to swivel their eyes and integrate images from different stages in the scan. The downside is that the scanning and integrating processes are relatively slow.[20]
+There are spiders with a reduced number of eyes. Of these, those with six eyes (such as Periegops suterii) are the most numerous and are missing a pair of eyes on the anterior median line;[25] other species have four eyes and some just two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.
+As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae.[22] An adult Araneus may have up to 1,000 such chemosensitive setae, most on the tarsi of the first pair of legs. Males have more chemosensitive bristles on their pedipalps than females. They have been shown to be responsive to sex pheromones produced by females, both contact and air-borne.[26] The jumping spider Evarcha culicivora uses the scent of blood from mammals and other vertebrates, which is obtained by capturing blood-filled mosquitoes, to attract the opposite sex. Because they are able to tell the sexes apart, it is assumed the blood scent is mixed with pheromones.[27] Spiders also have in the joints of their limbs slit sensillae that detect force and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.[11]
+Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' proprioceptors, sensors that report the force exerted by muscles and the degree of bending in the body and joints, are well-understood. On the other hand, little is known about what other internal sensors spiders or other arthropods may have.[22]
+Each of the eight legs of a spider consists of seven distinct parts. The part closest to and attaching the leg to the cephalothorax is the coxa; the next segment is the short trochanter that works as a hinge for the following long segment, the femur; next is the spider's knee, the patella, which acts as the hinge for the tibia; the metatarsus is next, and it connects the tibia to the tarsus (which may be thought of as a foot of sorts); the tarsus ends in a claw made up of either two or three points, depending on the family to which the spider belongs. Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors.[28] The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter).[29] As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up.[11] Spiders can generate pressures up to eight times their resting level to extend their legs,[30] and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs.[11] Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.[29]
+Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine bristles between the paired claws at the tips of their legs. These tufts, known as scopulae, consist of bristles whose ends are split into as many as 1,000 branches, and enable spiders with scopulae to walk up vertical glass and upside down on ceilings. It appears that scopulae get their grip from contact with extremely thin layers of water on surfaces.[11] Spiders, like most other arachnids, keep at least four legs on the surface while walking or running.[31]
+The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk.[11]
+Silk is mainly composed of a protein very similar to that used in insect silk. It is initially a liquid, and hardens not by exposure to air but as a result of being drawn out, which changes the internal structure of the protein.[32] It is similar in tensile strength to nylon and biological materials such as chitin, collagen and cellulose, but is much more elastic. In other words, it can stretch much further before breaking or losing shape.[11]
+Some spiders have a cribellum, a modified spinneret with up to 40,000 spigots, each of which produces a single very fine fiber. The fibers are pulled out by the calamistrum, a comblike set of bristles on the jointed tip of the cribellum, and combined into a composite woolly thread that is very effective in snagging the bristles of insects. The earliest spiders had cribella, which produced the first silk capable of capturing insects, before spiders developed silk coated with sticky droplets. However, most modern groups of spiders have lost the cribellum.[11]
+Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a "safety rope"; for nest-building; and as "parachutes" by the young of some species.[11]
+Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods,[33] male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs onto which they ejaculate and then transfer the sperm to special syringe-styled structures, palpal bulbs or palpal organs, borne on the tips of the pedipalps of mature males. When a male detects signs of a female nearby he checks whether she is of the same species and whether she is ready to mate; for example in species that produce webs or "safety ropes", the male can identify the species and sex of these objects by "smell".[11]
+Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female via one or two openings on the underside of her abdomen.[11]
+Female spiders' reproductive tracts are arranged in one of two ways. The ancestral arrangement ("haplogyne" or "non-entelegyne") consists of a single genital opening, leading to two seminal receptacles (spermathecae) in which females store sperm. In the more advanced arrangement ("entelegyne" ), there are two further openings leading directly to the spermathecae, creating a "flow through" system rather than a "first-in first-out" one. Eggs are as a general rule only fertilized during oviposition when the stored sperm is released from its chamber, rather than in the ovarian cavity.[34] A few exceptions exist, such as Parasteatoda tepidariorum. In these species the female appears to be able to activate the dormant sperm before oviposition, allowing them to migrate to the ovarian cavity where fertilization occurs.[35][36][37] The only known example of direct fertilization between male and female is an Israeli spider, Harpactea sadistica, which has evolved traumatic insemination. In this species the male will penetrate its pedipalps through the female's body wall and inject his sperm directly into her ovaries, where the embryos inside the fertilized eggs will start to develop before being laid.[38]
+Males of the genus Tidarren amputate one of their palps before maturation and enter adult life with one palp only. The palps are 20% of the male's body mass in this species, and detaching one of the two improves mobility. In the Yemeni species Tidarren argo, the remaining palp is then torn off by the female. The separated palp remains attached to the female's epigynum for about four hours and apparently continues to function independently. In the meantime, the female feeds on the palpless male.[39] In over 60% of cases, the female of the Australian redback spider kills and eats the male after it inserts its second palp into the female's genital opening; in fact, the males co-operate by trying to impale themselves on the females' fangs. Observation shows that most male redbacks never get an opportunity to mate, and the "lucky" ones increase the likely number of offspring by ensuring that the females are well-fed.[40] However, males of most species survive a few matings, limited mainly by their short life spans. Some even live for a while in their mates' webs.[41]
+The tiny male of the Golden orb weaver (Trichonephila clavipes) (near the top of the leaf) is protected from the female by producing the right vibrations in the web, and may be too small to be worth eating.
+Orange spider egg sac hanging from ceiling
+Gasteracantha mammosa spiderlings next to their eggs capsule
+Wolf spider carrying its young on its abdomen
+Females lay up to 3,000 eggs in one or more silk egg sacs,[11] which maintain a fairly constant humidity level.[41] In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.[11]
+Baby spiders pass all their larval stages inside the egg and hatch as spiderlings, very small and sexually immature but similar in shape to adults. Some spiders care for their young, for example a wolf spider's brood clings to rough bristles on the mother's back,[11] and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.[41]
+Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch.[42] In some species males mate with newly-molted females, which are too weak to be dangerous to the males.[41] Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years,[11][43] and an Australian female trapdoor spider was documented to have lived in the wild for 43 years, dying of a parasitic wasp attack.[44]
+Spiders occur in a large range of sizes. The smallest, Patu digua from Colombia, are less than 0.37 mm (0.015 in) in body length. The largest and heaviest spiders occur among tarantulas, which can have body lengths up to 90 mm (3.5 in) and leg spans up to 250 mm (9.8 in).[45]
+Only three classes of pigment (ommochromes, bilins and guanine) have been identified in spiders, although other pigments have been detected but not yet characterized. Melanins, carotenoids and pterins, very common in other animals, are apparently absent. In some species, the exocuticle of the legs and prosoma is modified by a tanning process, resulting in a brown coloration.[46]
+Bilins are found, for example, in Micrommata virescens, resulting in its green color. Guanine is responsible for the white markings of the European garden spider Araneus diadematus. It is in many species accumulated in specialized cells called guanocytes. In genera such as Tetragnatha, Leucauge, Argyrodes or Theridiosoma, guanine creates their silvery appearance. While guanine is originally an end-product of protein metabolism, its excretion can be blocked in spiders, leading to an increase in its storage.[46] Structural colors occur in some species, which are the result of the diffraction, scattering or interference of light, for example by modified setae or scales. The white prosoma of Argiope results from bristles reflecting the light, Lycosa and Josa both have areas of modified cuticle that act as light reflectors.[46]
+While in many spiders color is fixed throughout their lifespan, in some groups, color may be variable in response to environmental and internal conditions.[46] Choice of prey may be able to alter the color of spiders. For example, the abdomen of Theridion grallator will become orange if the spider ingests certain species of Diptera and adult Lepidoptera, but if it consumes Homoptera or larval Lepidoptera, then the abdomen becomes green.[47] Environmentally induced color changes may be morphological (occurring over several days) or physiological (occurring near instantly). Morphological changes require pigment synthesis and degradation. In contrast to this, physiological changes occur by changing the position of pigment-containing cells.[46] An example of morphological color changes is background matching. Misumena vatia for instance can change its body color to match the substrate it lives on which makes it more difficult to be detected by prey.[48] An example of physiological color change is observed in Cyrtophora cicatrosa, which can change its body color from white to brown near instantly.[46]
+Although spiders are generally regarded as predatory, the jumping spider Bagheera kiplingi gets over 90% of its food from fairly solid plant material produced by acacias as part of a mutually beneficial relationship with a species of ant.[49]
+Juveniles of some spiders in the families Anyphaenidae, Corinnidae, Clubionidae, Thomisidae and Salticidae feed on plant nectar. Laboratory studies show that they do so deliberately and over extended periods, and periodically clean themselves while feeding. These spiders also prefer sugar solutions to plain water, which indicates that they are seeking nutrients. Since many spiders are nocturnal, the extent of nectar consumption by spiders may have been underestimated. Nectar contains amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.[50]
+Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.[50]
+The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.[11]
+Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it.[51] A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.[11]
+Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus Hyptiotes and the family Theridiosomatidae stretch their webs and then release them when prey strike them, but do not actively move their webs. Those of the family Deinopidae weave even smaller webs, hold them outstretched between their first two pairs of legs, and lunge and push the webs as much as twice their own body length to trap prey, and this move may increase the webs' area by a factor of up to ten. Experiments have shown that Deinopis spinosus has two different techniques for trapping prey: backwards strikes to catch flying insects, whose vibrations it detects; and forward strikes to catch ground-walking prey that it sees. These two techniques have also been observed in other deinopids. Walking insects form most of the prey of most deinopids, but one population of Deinopis subrufa appears to live mainly on tipulid flies that they catch with the backwards strike.[52]
+Mature female bolas spiders of the genus Mastophora build "webs" that consist of only a single "trapeze line", which they patrol. They also construct a bolas made of a single thread, tipped with a large ball of very wet sticky silk. They emit chemicals that resemble the pheromones of moths, and then swing the bolas at the moths. Although they miss on about 50% of strikes, they catch about the same weight of insects per night as web-weaving spiders of similar size. The spiders eat the bolas if they have not made a kill in about 30 minutes, rest for a while, and then make new bolas.[53][54] Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.[55]
+The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush predators that lurk in burrows, often closed by trapdoors and often surrounded by networks of silk threads that alert these spiders to the presence of prey.[15] Other ambush predators do without such aids, including many crab spiders,[11] and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking.[46] Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.[11]
+Some jumping spiders of the genus Portia hunt other spiders in ways that seem intelligent,[20] outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species.[56] However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.[20]
+Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective bristles to resemble the shiny bodies of ants. In some spider species, males and females mimic different ant species, as female spiders are usually much larger than males. Ant-mimicking spiders also modify their behavior to resemble that of the target species of ant; for example, many adopt a zig-zag pattern of movement, ant-mimicking jumping spiders avoid jumping, and spiders of the genus Synemosyna walk on the outer edges of leaves in the same way as Pseudomyrmex. Ant mimicry in many spiders and other arthropods may be for protection from predators that hunt by sight, including birds, lizards and spiders. However, several ant-mimicking spiders prey either on ants or on the ants' "livestock", such as aphids. When at rest, the ant-mimicking crab spider Amyciaea does not closely resemble Oecophylla, but while hunting it imitates the behavior of a dying ant to attract worker ants. After a kill, some ant-mimicking spiders hold their victims between themselves and large groups of ants to avoid being attacked.[57]
+There is strong evidence that spiders' coloration is camouflage that helps them to evade their major predators, birds and parasitic wasps, both of which have good color vision. Many spider species are colored so as to merge with their most common backgrounds, and some have disruptive coloration, stripes and blotches that break up their outlines. In a few species, such as the Hawaiian happy-face spider, Theridion grallator, several coloration schemes are present in a ratio that appears to remain constant, and this may make it more difficult for predators to recognize the species. Most spiders are insufficiently dangerous or unpleasant-tasting for warning coloration to offer much benefit. However, a few species with powerful venom, large jaws or irritant bristles have patches of warning colors, and some actively display these colors when threatened.[46][58]
+Many of the family Theraphosidae, which includes tarantulas and baboon spiders, have urticating hairs on their abdomens and use their legs to flick them at attackers. These bristles are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom.[59] A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles.[60] The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.[61]
+A few spider species that build webs live together in large colonies and show social behavior, although not as complex as in social insects. Anelosimus eximius (in the family Theridiidae) can form colonies of up to 50,000 individuals.[62] The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social.[63] Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food.[64] Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae).[65] Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this.[66] The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings.[49] Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.[67]
+There is no consistent relationship between the classification of spiders and the types of web they build: species in the same genus may build very similar or significantly different webs. Nor is there much correspondence between spiders' classification and the chemical composition of their silks. Convergent evolution in web construction, in other words use of similar techniques by remotely related species, is rampant. Orb web designs and the spinning behaviors that produce them are the best understood. The basic radial-then-spiral sequence visible in orb webs and the sense of direction required to build them may have been inherited from the common ancestors of most spider groups.[68] However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a subgroup that evolved from orb-web spiders, and non-orb spiders have over 40% more species and are four times as abundant as orb-web spiders. Their greater success may be because sphecid wasps, which are often the dominant predators of spiders, much prefer to attack spiders that have flat webs.[69]
+About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey.[68]
+The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.[68]
+Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the backlighting from the sky; enabling oscillations to catch insects in slow horizontal flight. However, there is no single explanation for the common use of horizontal orb webs.[68]
+Spiders often attach highly visible silk bands, called decorations or stabilimenta, to their webs. Field research suggests that webs with more decorative bands captured more prey per hour.[70] However, a laboratory study showed that spiders reduce the building of these decorations if they sense the presence of predators.[71]
+There are several unusual variants of orb web, many of them convergently evolved, including: attachment of lines to the surface of water, possibly to trap insects in or on the surface; webs with twigs through their centers, possibly to hide the spiders from predators; "ladderlike" webs that appear most effective in catching moths. However, the significance of many variations is unclear.[68]
+In 1973, Skylab 3 took two orb-web spiders into space to test their web-spinning capabilities in zero gravity. At first, both produced rather sloppy webs, but they adapted quickly.[72]
+Members of the family Theridiidae weave irregular, tangled, three-dimensional webs, popularly known as cobwebs. There seems to be an evolutionary trend towards a reduction in the amount of sticky silk used, leading to its total absence in some species. The construction of cobwebs is less stereotyped than that of orb-webs, and may take several days.[69]
+The Linyphiidae generally make horizontal but uneven sheets, with tangles of stopping threads above. Insects that hit the stopping threads fall onto the sheet or are shaken onto it by the spider, and are held by sticky threads on the sheet until the spider can attack from below.[73]
+Although the fossil record of spiders is considered poor,[74] almost 1000 species have been described from fossils.[75] Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber.[75] The oldest known amber that contains fossil arthropods dates from 130 million years ago in the Early Cretaceous period. In addition to preserving spiders' anatomy in very fine detail, pieces of amber show spiders mating, killing prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached;[76] the oldest fossil web found so far is 100 million years old.[77] Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.[76]
+The oldest known exclusively terrestrial arachnid is the trigonotarbid Palaeotarbus jerami, from about 420 million years ago in the Silurian period, and had a triangular cephalothorax and segmented abdomen, as well as eight legs and a pair of pedipalps.[78] Attercopus fimbriunguis, from 386 million years ago in the Devonian period, bears the earliest known silk-producing spigots, and was therefore hailed as a spider at the time of its discovery.[79] However, these spigots may have been mounted on the underside of the abdomen rather than on spinnerets, which are modified appendages and whose mobility is important in the building of webs. Hence Attercopus and the similar Permian arachnid Permarachne may not have been true spiders, and probably used silk for lining nests or producing egg cases rather than for building webs.[3] The largest known fossil spider as of 2011 is the araneid Nephila jurassica, from about 165 million years ago, recorded from Daohuogo, Inner Mongolia in China.[80] Its body length is almost 25 mm, (i.e., almost one inch).
+Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae.[79]
+The mesothelid Paleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets.[81] Although the Permian period 299 to 251 million years ago saw rapid diversification of flying insects, there are very few fossil spiders from this period.[79]
+The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appear in the Triassic well before 200 million years ago. Some Triassic mygalomorphs appear to be members of the family Hexathelidae, whose modern members include the notorious Sydney funnel-web spider, and their spinnerets appear adapted for building funnel-shaped webs to catch jumping insects. Araneomorphae account for the great majority of modern spiders, including those that weave the familiar orb-shaped webs. The Jurassic and Cretaceous periods provide a large number of fossil spiders, including representatives of many modern families.[79]
+Xiphosura (horseshoe crabs)
+Eurypterida†
+Chasmataspidida†
+Scorpiones
+Opiliones (harvestmen)
+Pseudoscorpiones
+Solifugae (sun spiders)
+Palpigradi (microwhip scorpions)
+Trigonotarbida†
+Araneae (spiders)
+Haptopoda†
+Amblypygi (whip spiders)
+Thelyphonida (whip scorpions)
+Schizomida
+Ricinulei (hooded tickspiders)
+Anactinotrichida
+Acariformes (mites)
+It is now agreed that spiders (Araneae) are monophyletic (i.e., members of a group of organisms that form a clade, consisting of a last common ancestor and all of its descendants).[83] There has been debate about what their closest evolutionary relatives are, and how all of these evolved from the ancestral chelicerates, which were marine animals. The cladogram on the right is based on J. W. Shultz' analysis (2007). Other views include proposals that: scorpions are more closely related to the extinct marine scorpioid eurypterids than to spiders; spiders and Amblypygi are a monophyletic group. The appearance of several multi-way branchings in the tree on the right shows that there are still uncertainties about relationships between the groups involved.[83]
+Arachnids lack some features of other chelicerates, including backward-pointing mouths and gnathobases ("jaw bases") at the bases of their legs;[83] both of these features are part of the ancestral arthropod feeding system.[84] Instead, they have mouths that point forwards and downwards, and all have some means of breathing air.[83] Spiders (Araneae) are distinguished from other arachnid groups by several characteristics, including spinnerets and, in males, pedipalps that are specially adapted for sperm transfer.[85]
+Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two infraorders, Mygalomorphae and Araneomorphae. Over 48,000 living species of spiders (order Araneae) have been identified and as of 2019 grouped into 120 families and about 4,100 genera by arachnologists.[1]
+The only living members of the primitive Mesothelae are the family Liphistiidae, found only in Southeast Asia, China, and Japan.[85] Most of the Liphistiidae construct silk-lined burrows with thin trapdoors, although some species of the genus Liphistius build camouflaged silk tubes with a second trapdoor as an emergency exit. Members of the genus Liphistius run silk "tripwires" outwards from their tunnels to help them detect approaching prey, while those of the genus Heptathela do not and instead rely on their built-in vibration sensors.[88] Spiders of the genus Heptathela have no venom glands, although they do have venom gland outlets on the fang tip.[89]
+The extinct families Arthrolycosidae, found in Carboniferous and Permian rocks, and Arthromygalidae, so far found only in Carboniferous rocks, have been classified as members of the Mesothelae.[90]
+The Mygalomorphae, which first appeared in the Triassic period,[79] are generally heavily built and ″hairy″, with large, robust chelicerae and fangs (technically, spiders do not have true hairs, but rather setae).[91][85] Well-known examples include tarantulas, ctenizid trapdoor spiders and the Australasian funnel-web spiders.[11] Most spend the majority of their time in burrows, and some run silk tripwires out from these, but a few build webs to capture prey. However, mygalomorphs cannot produce the pirifom silk that the Araneomorphae use as an instant adhesive to glue silk to surfaces or to other strands of silk, and this makes web construction more difficult for mygalomorphs. Since mygalomorphs rarely "balloon" by using air currents for transport, their populations often form clumps.[85] In addition to arthropods, some mygalomorphs are known to prey on frogs, small mammals, lizards, snakes, snails, and small birds.[92][93]
+In addition to accounting for over 90% of spider species, the Araneomorphae, also known as the "true spiders", include orb-web spiders, the cursorial wolf spiders, and jumping spiders,[85] as well as the only known herbivorous spider, Bagheera kiplingi.[49] They are distinguished by having fangs that oppose each other and cross in a pinching action, in contrast to the Mygalomorphae, which have fangs that are nearly parallel in alignment.[94]
+Although spiders are widely feared, only a few species are dangerous to people.[96] Spiders will only bite humans in self-defense, and few produce worse effects than a mosquito bite or bee sting.[97] Most of those with medically serious bites, such as recluse spiders (genus Loxosceles) and widow spiders (genus Latrodectus), would rather flee and bite only when trapped, although this can easily arise by accident.[98][99] The defensive tactics of Australian funnel-web spiders (family Atracidae) include fang display. Their venom, although they rarely inject much, has resulted in 13 attributed human deaths over 50 years.[100] They have been deemed to be the world's most dangerous spiders on clinical and venom toxicity grounds,[96] though this claim has also been attributed to the Brazilian wandering spider (genus Phoneutria).[101]
+There were about 100 reliably reported deaths from spider bites in the 20th century,[102] compared to about 1,500 from jellyfish stings.[103] Many alleged cases of spider bites may represent incorrect diagnoses,[104] which would make it more difficult to check the effectiveness of treatments for genuine bites.[105] A review published in 2016 agreed with this conclusion, showing that 78% of 134 published medical case studies of supposed spider bites did not meet the necessary criteria for a spider bite to be verified. In the case of the two genera with the highest reported number of bites, Loxosceles and Latrodectus, spider bites were not verified in over 90% of the reports. Even when verification had occurred, details of the treatment and its effects were often lacking.[106]
+Spider venoms may be a less polluting alternative to conventional pesticides, as they are deadly to insects but the great majority are harmless to vertebrates. Australian funnel web spiders are a promising source, as most of the world's insect pests have had no opportunity to develop any immunity to their venom, and funnel web spiders thrive in captivity and are easy to "milk". It may be possible to target specific pests by engineering genes for the production of spider toxins into viruses that infect species such as cotton bollworms.[107]
+The Ch'ol Maya use a beverage created from the tarantula species Brachypelma vagans for the treatment of a condition they term 'tarantula wind', the symptoms of which include chest pain, asthma and coughing.[108]
+Possible medical uses for spider venoms are being investigated, for the treatment of cardiac arrhythmia,[109] Alzheimer's disease,[110] strokes,[111] and erectile dysfunction.[112]
+The peptide GsMtx-4, found in the venom of Brachypelma vagans, is being researched to determine whether or not it could effectively be used for the treatment of cardiac arrhythmia, muscular dystrophy or glioma.[108]
+Because spider silk is both light and very strong, attempts are being made to produce it in goats' milk and in the leaves of plants, by means of genetic engineering.[113][114]
+Spiders can also be used as food. Cooked tarantulas are considered a delicacy in Cambodia,[115] and by the Piaroa Indians of southern Venezuela – provided the highly irritant bristles, the spiders' main defense system, are removed first.[116]
+Arachnophobia is a specific phobia—it is the abnormal fear of spiders or anything reminiscent of spiders, such as webs or spiderlike shapes. It is one of the most common specific phobias,[117][118] and some statistics show that 50% of women and 10% of men show symptoms.[119] It may be an exaggerated form of an instinctive response that helped early humans to survive,[120] or a cultural phenomenon that is most common in predominantly European societies.[121]
+Spiders have been the focus of stories and mythologies of various cultures for centuries.[122] Uttu, the ancient Sumerian goddess of weaving, was envisioned as a spider spinning her web.[123][124] According to her main myth, she resisted her father Enki's sexual advances by ensconcing herself in her web,[124] but let him in after he promised her fresh produce as a marriage gift,[124] thereby allowing him to intoxicate her with beer and rape her.[124] Enki's wife Ninhursag heard Uttu's screams and rescued her,[124] removing Enki's semen from her vagina and planting it in the ground to produce eight previously-nonexistent plants.[124] In a story told by the Roman poet Ovid in his Metamorphoses, Arachne was a Lydian girl who challenged the goddess Athena to a weaving contest.[125][126] Arachne won, but Athena destroyed her tapestry out of jealousy,[126][127] causing Arachne to hang herself.[126][127] In an act of mercy, Athena brought Arachne back to life as the first spider.[126][127] Stories about the trickster-spider Anansi are prominent in the folktales of West Africa and the Caribbean.[128]
+In some cultures, spiders have symbolized patience due to their hunting technique of setting webs and waiting for prey, as well as mischief and malice due to their venomous bites.[129] The Italian tarantella is a dance to rid the young woman of the lustful effects of a spider bite. Web-spinning also caused the association of the spider with creation myths, as they seem to have the ability to produce their own worlds.[130] Dreamcatchers are depictions of spiderwebs. The Moche people of ancient Peru worshipped nature.[131] They placed emphasis on animals and often depicted spiders in their art.[132]

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+Recycling is the process of converting waste materials into new materials and objects. The recyclability of a material depends on its ability to reacquire the properties it had in its virgin or original state.[1] It is an alternative to "conventional" waste disposal that can save material and help lower greenhouse gas emissions. Recycling can prevent the waste of potentially useful materials and reduce the consumption of fresh raw materials, thereby reducing: energy usage, air pollution (from incineration), and water pollution (from landfilling).
+Recycling is a key component of modern waste reduction and is the third component of the "Reduce, Reuse, and Recycle" waste hierarchy.[2][3] Thus, recycling aims at environmental sustainability by substituting raw material inputs into and redirecting waste outputs out of the economic system.[4] There are some ISO standards related to recycling such as ISO 15270:2008 for plastics waste and ISO 14001:2015 for environmental management control of recycling practice.
+Recyclable materials include many kinds of glass, paper, cardboard, metal, plastic, tires, textiles, batteries, and electronics. The composting or other reuse of biodegradable waste—such as food or garden waste—is also a form of recycling.[5] Materials to be recycled are either delivered to a household recycling center or picked up from curbside bins, then sorted, cleaned, and reprocessed into new materials destined for manufacturing new products.
+In the strictest sense, recycling of a material would produce a fresh supply of the same material—for example, used office paper would be converted into new office paper or used polystyrene foam into new polystyrene. This is accomplished when recycling certain types of materials, such as metal cans, which can become a can again and again, indefinitely, without losing purity in the product.[6] However, this is often difficult or too expensive (compared with producing the same product from raw materials or other sources), so "recycling" of many products or materials involves their reuse in producing different materials (for example, paperboard) instead. Another form of recycling is the salvage of certain materials from complex products, either due to their intrinsic value (such as lead from car batteries, or gold from printed circuit boards), or due to their hazardous nature (e.g., removal and reuse of mercury from thermometers and thermostats).
+Recycling has been a common practice for most of human history, with recorded advocates as far back as Plato in the fourth century BC.[citation needed] During periods when resources were scarce and hard to come by, archaeological studies of ancient waste dumps show less household waste (such as ash, broken tools, and pottery)—implying more waste was being recycled in the absence of new material.[7]
+In pre-industrial times, there is evidence of scrap bronze and other metals being collected in Europe and melted down for continuous reuse.[8] Paper recycling was first recorded in 1031 when Japanese shops sold repulped paper.[9][10] In Britain dust and ash from wood and coal fires was collected by "dustmen" and downcycled as a base material used in brick making. The main driver for these types of recycling was the economic advantage of obtaining recycled feedstock instead of acquiring virgin material, as well as a lack of public waste removal in ever more densely populated areas.[7] In 1813, Benjamin Law developed the process of turning rags into "shoddy" and "mungo" wool in Batley, Yorkshire. This material combined recycled fibers with virgin wool.[11] The West Yorkshire shoddy industry in towns such as Batley and Dewsbury lasted from the early 19th century to at least 1914.
+Industrialization spurred demand for affordable materials; aside from rags, ferrous scrap metals were coveted as they were cheaper to acquire than virgin ore. Railroads both purchased and sold scrap metal in the 19th century, and the growing steel and automobile industries purchased scrap in the early 20th century. Many secondary goods were collected, processed and sold by peddlers who scoured dumps and city streets for discarded machinery, pots, pans, and other sources of metal. By World War I, thousands of such peddlers roamed the streets of American cities, taking advantage of market forces to recycle post-consumer materials back into industrial production.[12]
+Beverage bottles were recycled with a refundable deposit at some drink manufacturers in Great Britain and Ireland around 1800, notably Schweppes.[13] An official recycling system with refundable deposits was established in Sweden for bottles in 1884 and aluminum beverage cans in 1982; the law led to a recycling rate for beverage containers of 84–99 percent depending on type, and a glass bottle can be refilled over 20 times on average.[14]
+New chemical industries created in the late 19th century both invented new materials (e.g. Bakelite [1907]) and promised to transform valueless into valuable materials. Proverbially, you could not make a silk purse of a sow's ear—until the US firm Arthur D. Little published in 1921 "On the Making of Silk Purses from Sows' Ears", its research proving that when "chemistry puts on overalls and gets down to business ... new values appear. New and better paths are opened to reach the goals desired."[15]
+Recycling (or "salvage", as it was then usually known) was a major issue for governments throughout World War II. Financial constraints and significant material shortages due to war efforts made it necessary for countries to reuse goods and recycle materials.[16] These resource shortages caused by the world wars, and other such world-changing occurrences, greatly encouraged recycling.[17] The struggles of war claimed much of the material resources available, leaving little for the civilian population.[16] It became necessary for most homes to recycle their waste, as recycling offered an extra source of materials allowing people to make the most of what was available to them. Recycling household materials meant more resources for war efforts and a better chance of victory.[16] Massive government promotion campaigns, such as the National Salvage Campaign in Britain and the Salvage for Victory campaign in the United States, were carried out on the home front in every combative nation, urging citizens to donate metal, paper, rags, and rubber as a matter of patriotism.
+A considerable investment in recycling occurred in the 1970s, due to rising energy costs.[18] Recycling aluminium uses only 5% of the energy required by virgin production; glass, paper and other metals have less dramatic but very significant energy savings when recycled feedstock is used.[19]
+Although consumer electronics such as the television have been popular since the 1920s, recycling of them was almost unheard of until early 1991.[20] The first electronic waste recycling scheme was implemented in Switzerland, beginning with collection of old refrigerators but gradually expanding to cover all devices.[21] After these schemes were set up, many countries did not have the capacity to deal with the sheer quantity of e-waste they generated or its hazardous nature. They began to export the problem to developing countries without enforced environmental legislation. This is cheaper, as recycling computer monitors in the United States costs 10 times more than in China. Demand in Asia for electronic waste began to grow when scrap yards found that they could extract valuable substances such as copper, silver, iron, silicon, nickel, and gold, during the recycling process.[22] The 2000s saw a large increase in both the sale of electronic devices and their growth as a waste stream: in 2002, e-waste grew faster than any other type of waste in the EU.[23] This caused investment in modern, automated facilities to cope with the influx of redundant appliances, especially after strict laws were implemented in 2003.[24][25][26][27]
+As of 2014, the European Union had about 50% of world share of the waste and recycling industries, with over 60,000 companies employing 500,000 persons, with a turnover of €24 billion.[28] Countries have to reach recycling rates of at least 50%, while the lead countries were around 65% and the EU average was 39% as of 2013.[29]
+The EU average has been rising steadily, to 45% in 2015.[30][31]
+In 2018, changes in the recycling market have sparked a global "crisis" in the industry. On 31 December 2017, China announced its "National Sword" policy, setting new standards for imports of recyclable material and banning materials that were deemed too "dirty" or "hazardous". The new policy caused drastic disruptions in the global market in recycling and reduced the prices of scrap plastic and low-grade paper. Exports of recyclable materials from G7 countries to China dropped dramatically, with many exports shifting to countries in southeast Asia. The crisis generated significant concern about the practices and environmental sustainability of the recycling industry. The abrupt shift caused countries to accept more recyclable materials than they could process, raising fundamental questions about shipping recycling waste from economically developed countries to countries with few environmental regulations—a practice that predated the crisis.[32]
+For a recycling program to work, having a large, stable supply of recyclable material is crucial. Three legislative options have been used to create such a supply: mandatory recycling collection, container deposit legislation, and refuse bans. Mandatory collection laws set recycling targets for cities to aim for, usually in the form that a certain percentage of a material must be diverted from the city's waste stream by a target date. The city is then responsible for working to meet this target.[5]
+Container deposit legislation involves offering a refund for the return of certain containers, typically glass, plastic, and metal. When a product in such a container is purchased, a small surcharge is added to the price. This surcharge can be reclaimed by the consumer if the container is returned to a collection point. These programs have been very successful, often resulting in an 80 percent recycling rate.[33] Despite such good results, the shift in collection costs from local government to industry and consumers has created strong opposition to the creation of such programs in some areas.[5] A variation on this is where the manufacturer bears responsibility for the recycling of their goods. In the European Union, the WEEE Directive requires producers of consumer electronics to reimburse the recyclers' costs.[34]
+An alternative way to increase the supply of recyclates is to ban the disposal of certain materials as waste, often including used oil, old batteries, tires, and garden waste. One aim of this method is to create a viable economy for proper disposal of banned products. Care must be taken that enough of these recycling services exist, or such bans simply lead to increased illegal dumping.[5]
+Legislation has also been used to increase and maintain a demand for recycled materials. Four methods of such legislation exist: minimum recycled content mandates, utilization rates, procurement policies, and recycled product labeling.[5]
+Both minimum recycled content mandates and utilization rates increase demand directly by forcing manufacturers to include recycling in their operations. Content mandates specify that a certain percentage of a new product must consist of recycled material. Utilization rates are a more flexible option: industries are permitted to meet the recycling targets at any point of their operation or even contract recycling out in exchange for tradeable credits. Opponents to both of these methods point to the large increase in reporting requirements they impose, and claim that they rob the industry of necessary flexibility.[5][35]
+Governments have used their own purchasing power to increase recycling demand through what are called "procurement policies". These policies are either "set-asides", which reserve a certain amount of spending solely towards recycled products, or "price preference" programs which provide a larger budget when recycled items are purchased. Additional regulations can target specific cases: in the United States, for example, the Environmental Protection Agency mandates the purchase of oil, paper, tires and building insulation from recycled or re-refined sources whenever possible.[5]
+The final government regulation towards increased demand is recycled product labeling. When producers are required to label their packaging with amount of recycled material in the product (including the packaging), consumers are better able to make educated choices. Consumers with sufficient buying power can then choose more environmentally conscious options, prompt producers to increase the amount of recycled material in their products, and indirectly increase demand. Standardized recycling labeling can also have a positive effect on supply of recyclates if the labeling includes information on how and where the product can be recycled.[5]
+"Recyclate" is a raw material that is sent to, and processed in a waste recycling plant or materials recovery facility which will be used to form new products.[36] The material is collected in various methods and delivered to a facility where it undergoes re-manufacturing so that it can be used in the production of new materials or products. For example, plastic bottles that are collected can be re-used and made into plastic pellets, a new product.[37]
+The quality of recyclates is recognized as one of the principal challenges that needs to be addressed for the success of a long-term vision of a green economy and achieving zero waste. Recyclate quality is generally referring to how much of the raw material is made up of target material compared to the amount of non-target material and other non-recyclable material.[38] For example, steel and metal are materials with a higher recyclate quality. It's estimated that two-thirds of all new steel manufactured comes from recycled steel.[39] Only target material is likely to be recycled, so a higher amount of non-target and non-recyclable material will reduce the quantity of recycling product.[38] A high proportion of non-target and non-recyclable material can make it more difficult for re-processors to achieve "high-quality" recycling. If the recyclate is of poor quality, it is more likely to end up being down-cycled or, in more extreme cases, sent to other recovery options or landfilled.[38] For example, to facilitate the re-manufacturing of clear glass products there are tight restrictions for colored glass going into the re-melt process. Another example is the downcycling of plastic, in which products such as plastic food packaging are often downcycled into lower quality products, and do not get recycled into the same plastic food packaging.
+The quality of recyclate not only supports high-quality recycling, but it can also deliver significant environmental benefits by reducing, reusing and keeping products out of landfills.[38] High-quality recycling can help support growth in the economy by maximizing the economic value of the waste material collected.[38] Higher income levels from the sale of quality recyclates can return value which can be significant to local governments, households, and businesses.[38] Pursuing high-quality recycling can also provide consumer and business confidence in the waste and resource management sector and may encourage investment in that sector.
+There are many actions along the recycling supply chain that can influence and affect the material quality of recyclate.[40] It begins with the waste producers who place non-target and non-recyclable wastes in recycling collection. This can affect the quality of final recyclate streams or require further efforts to discard those materials at later stages in the recycling process.[40] The different collection systems can result in different levels of contamination. Depending on which materials are collected together, extra effort is required to sort this material back into separate streams and can significantly reduce the quality of the final product.[40] Transportation and the compaction of materials can make it more difficult to separate material back into separate waste streams. Sorting facilities are not one hundred per cent effective in separating materials, despite improvements in technology and quality recyclate which can see a loss in recyclate quality.[40] The storage of materials outside, where the product can become wet, can cause problems for re-processors. Reprocessing facilities may require further sorting steps to further reduce the amount of non-target and non-recyclable material.[40] Each action along the recycling path plays a part in the quality of recyclate.
+The Recyclate Quality Action Plan of Scotland sets out a number of proposed actions that the Scottish Government would like to take forward in order to drive up the quality of the materials being collected for recycling and sorted at materials recovery facilities before being exported or sold on to the reprocessing market.[40]
+The plan's objectives are to:[41]
+The plan focuses on three key areas, with fourteen actions which were identified to increase the quality of materials collected, sorted and presented to the processing market in Scotland.[41]
+The three areas of focus are:[40]
+A number of different systems have been implemented to collect recyclates from the general waste stream. These systems lie along the spectrum of trade-off between public convenience and government ease and expense. The three main categories of collection are "drop-off centers", "buy-back centers", and "curbside collection".[5]
+Curbside collection encompasses many subtly different systems, which differ mostly on where in the process the recyclates are sorted and cleaned. The main categories are mixed waste collection, commingled recyclables, and source separation.[5] A waste collection vehicle generally picks up the waste.
+At one end of the spectrum is mixed waste collection, in which all recyclates are collected mixed in with the rest of the waste, and the desired material is then sorted out and cleaned at a central sorting facility. This results in a large amount of recyclable waste, paper especially, being too soiled to reprocess, but has advantages as well: the city need not pay for a separate collection of recyclates and no public education is needed. Any changes to which materials are recyclable is easy to accommodate as all sorting happens in a central location.[5]
+In a commingled or single-stream system, all recyclables for collection are mixed but kept separate from other waste. This greatly reduces the need for post-collection cleaning but does require public education on what materials are recyclable.[5][8]
+Source separation is the other extreme, where each material is cleaned and sorted prior to collection. This method requires the least post-collection sorting and produces the purest recyclates, but incurs additional operating costs for collection of each separate material. An extensive public education program is also required, which must be successful if recyclate contamination is to be avoided.[5] In Oregon, USA, its environmental authority Oregon DEQ surveyed multi-family property managers and about half of them reported problems including contamination of recyclables due to trespassers such as transients gaining access to the collection areas.[42]
+Source separation used to be the preferred method due to the high sorting costs incurred by commingled (mixed waste) collection. However, advances in sorting technology have lowered this overhead substantially. Many areas which had developed source separation programs have since switched to what is called co-mingled collection.[8]
+Buy-back centers differ in that the cleaned recyclates are purchased, thus providing a clear incentive for use and creating a stable supply. The post-processed material can then be sold. If this is profitable, this conserves the emission of greenhouse gases; if unprofitable, it increases the emission of greenhouse gasses. Government subsidies are necessary to make buy-back centres a viable enterprise. In 1993, according to the U.S. National Waste & Recycling Association, it costs on average $50 to process a ton of material, which can be resold for $30.[5]
+In the US, the value per ton of mixed recyclables was $180 in 2011, $80 in 2015, and $100 in 2017.[43]
+In 2017, glass is essentially valueless, because of the low cost of sand, its major component; low oil costs thwarts plastic recycling.[43]
+In 2017, Napa, California was reimbursed about 20% of its costs in recycling.[43]
+Drop-off centers require the waste producer to carry the recyclates to a central location, either an installed or mobile collection station or the reprocessing plant itself. They are the easiest type of collection to establish but suffer from low and unpredictable throughput.
+For some waste materials such as plastic, recent technical devices called recyclebots[44] enable a form of distributed recycling. Preliminary life-cycle analysis (LCA) indicates that such distributed recycling of HDPE to make filament of 3D printers in rural regions is energetically favorable to either using virgin resin or conventional recycling processes because of reductions in transportation energy.[45][46]
+Once commingled recyclates are collected and delivered to a materials recovery facility, the different types of materials must be sorted. This is done in a series of stages, many of which involve automated processes such that a truckload of material can be fully sorted in less than an hour.[8] Some plants can now sort the materials automatically, known as single-stream recycling. Automatic sorting may be aided by robotics and machine-learning.[47][48] In plants, a variety of materials is sorted such as paper, different types of plastics, glass, metals, food scraps, and most types of batteries.[49] A 30 percent increase in recycling rates has been seen in the areas where these plants exist.[50] In the United States, there are over 300 materials recovery facilities.[51]
+Initially, the commingled recyclates are removed from the collection vehicle and placed on a conveyor belt spread out in a single layer. Large pieces of corrugated fiberboard and plastic bags are removed by hand at this stage, as they can cause later machinery to jam.[8]
+Next, automated machinery such as disk screens and air classifiers separate the recyclates by weight, splitting lighter paper and plastic from heavier glass and metal. Cardboard is removed from the mixed paper and the most common types of plastic, PET (#1) and HDPE (#2), are collected. This separation is usually done by hand but has become automated in some sorting centers: a spectroscopic scanner is used to differentiate between different types of paper and plastic based on the absorbed wavelengths, and subsequently divert each material into the proper collection channel.[8]
+Strong magnets are used to separate out ferrous metals, such as iron, steel, and tin cans. Non-ferrous metals are ejected by magnetic eddy currents in which a rotating magnetic field induces an electric current around the aluminum cans, which in turn creates a magnetic eddy current inside the cans. This magnetic eddy current is repulsed by a large magnetic field, and the cans are ejected from the rest of the recyclate stream.[8]
+Finally, glass is sorted according to its color: brown, amber, green, or clear. It may either be sorted by hand,[8] or via an automated machine that uses colored filters to detect different colors. Glass fragments smaller than 10 millimetres (0.39 in) across cannot be sorted automatically, and are mixed together as "glass fines".[52]
+This process of recycling as well as reusing the recycled material has proven advantageous because it reduces amount of waste sent to landfills, conserves natural resources, saves energy, reduces greenhouse gas emissions, and helps create new jobs. Recycled materials can also be converted into new products that can be consumed again, such as paper, plastic, and glass.[53]
+The City and County of San Francisco's Department of the Environment is attempting to achieve a citywide goal of generating zero waste by 2020.[54] San Francisco's refuse hauler, Recology, operates an effective recyclables sorting facility which helped the city reach a record-breaking diversion rate of 80%.[55]
+Although many government programs are concentrated on recycling at home, 64% of waste in the United Kingdom is generated by industry.[56] The focus of many recycling programs done by industry is the cost–effectiveness of recycling. The ubiquitous nature of cardboard packaging makes cardboard a commonly recycled waste product by companies that deal heavily in packaged goods, like retail stores, warehouses, and distributors of goods. Other industries deal in niche or specialized products, depending on the nature of the waste materials that are present.
+The glass, lumber, wood pulp and paper manufacturers all deal directly in commonly recycled materials; however, old rubber tires may be collected and recycled by independent tire dealers for a profit.
+Levels of metals recycling are generally low. In 2010, the International Resource Panel, hosted by the United Nations Environment Programme (UNEP) published reports on metal stocks that exist within society[57] and their recycling rates.[57] The Panel reported that the increase in the use of metals during the 20th and into the 21st century has led to a substantial shift in metal stocks from below ground to use in applications within society above ground. For example, the in-use stock of copper in the USA grew from 73 to 238 kg per capita between 1932 and 1999.
+The report authors observed that, as metals are inherently recyclable, the metal stocks in society can serve as huge mines above ground (the term "urban mining" has been coined with this idea in mind[58]). However, they found that the recycling rates of many metals are very low. The report warned that the recycling rates of some rare metals used in applications such as mobile phones, battery packs for hybrid cars and fuel cells, are so low that unless future end-of-life recycling rates are dramatically stepped up these critical metals will become unavailable for use in modern technology.
+The military recycles some metals. The U.S. Navy's Ship Disposal Program uses ship breaking to reclaim the steel of old vessels. Ships may also be sunk to create an artificial reef. Uranium is a very dense metal that has qualities superior to lead and titanium for many military and industrial uses. The uranium left over from processing it into nuclear weapons and fuel for nuclear reactors is called depleted uranium, and is used by all branches of the U.S. military for the development of such things as armour-piercing shells and shielding.
+The construction industry may recycle concrete and old road surface pavement, selling their waste materials for profit.
+Some industries, like the renewable energy industry and solar photovoltaic technology, in particular, are being proactive in setting up recycling policies even before there is considerable volume to their waste streams, anticipating future demand during their rapid growth.[59]
+Recycling of plastics is more difficult, as most programs are not able to reach the necessary level of quality. Recycling of PVC often results in downcycling of the material, which means only products of lower quality standard can be made with the recycled material. A new approach which allows an equal level of quality is the Vinyloop process. It was used after the London Olympics 2012 to fulfill the PVC Policy.[60]
+E-waste is a growing problem, accounting for 20–50 million metric tons of global waste per year according to the EPA. It is also the fastest growing waste stream in the EU.[23] Many recyclers do not recycle e-waste responsibly. After the cargo barge Khian Sea dumped 14,000 metric tons of toxic ash in Haiti, the Basel Convention was formed to stem the flow of hazardous substances into poorer countries. They created the e-Stewards certification to ensure that recyclers are held to the highest standards for environmental responsibility and to help consumers identify responsible recyclers. This works alongside other prominent legislation, such as the Waste Electrical and Electronic Equipment Directive of the EU the United States National Computer Recycling Act, to prevent poisonous chemicals from entering waterways and the atmosphere.
+In the recycling process, television sets, monitors, cell phones, and computers are typically tested for reuse and repaired. If broken, they may be disassembled for parts still having high value if labor is cheap enough. Other e-waste is shredded to pieces roughly 10 centimetres (3.9 in) in size, and manually checked to separate out toxic batteries and capacitors which contain poisonous metals. The remaining pieces are further shredded to 10 millimetres (0.39 in) particles and passed under a magnet to remove ferrous metals. An eddy current ejects non-ferrous metals, which are sorted by density either by a centrifuge or vibrating plates. Precious metals can be dissolved in acid, sorted, and smelted into ingots. The remaining glass and plastic fractions are separated by density and sold to re-processors. Television sets and monitors must be manually disassembled to remove lead from CRTs or the mercury backlight from LCDs.[61][62][63]
+Plastic recycling is the process of recovering scrap or waste plastic and reprocessing the material into useful products, sometimes completely different in form from their original state. For instance, this could mean melting down soft drink bottles and then casting them as plastic chairs and tables.[64] For some types of plastic, the same piece of plastic can only be recycled about 2–3 times before its quality decreases to the point where it can no longer be used.[6]
+Some plastics are remelted to form new plastic objects; for example, PET water bottles can be converted into polyester destined for clothing. A disadvantage of this type of recycling is that the molecular weight of the polymer can change further and the levels of unwanted substances in the plastic can increase with each remelt.[citation needed]
+A commercial-built recycling facility was sent to the International Space Station in late 2019. The facility will take in plastic waste and unneeded plastic parts and physically convert them into spools of feedstock for the space station additive manufacturing facility used for in-space 3D printing.[65]
+For some polymers, it is possible to convert them back into monomers, for example, PET can be treated with an alcohol and a catalyst to form a dialkyl terephthalate. The terephthalate diester can be used with ethylene glycol to form a new polyester polymer, thus making it possible to use the pure polymer again. In 2019, Eastman Chemical Company announced initiatives of methanolysis and syngas designed to handle a greater variety of used material.[66]
+Another process involves the conversion of assorted polymers into petroleum by a much less precise thermal depolymerization process. Such a process would be able to accept almost any polymer or mix of polymers, including thermoset materials such as vulcanized rubber tires and the biopolymers in feathers and other agricultural waste. Like natural petroleum, the chemicals produced can be used as fuels or as feedstock. A RESEM Technology[67] plant of this type in Carthage, Missouri, US, uses turkey waste as input material. Gasification is a similar process but is not technically recycling since polymers are not likely to become the result.
+Plastic Pyrolysis can convert petroleum based waste streams such as plastics into quality fuels, carbons. Given below is the list of suitable plastic raw materials for pyrolysis:
+The (ideal) recycling process can be differentiated into three loops, one for manufacture (production-waste recycling) and two for disposal of the product (product and material recycling).[68]
+The product's manufacturing phase, which consists of material processing and fabrication, forms the production-waste recycling loop. Industrial waste materials are fed back into, and reused in, the same production process.
+The product's disposal process requires two recycling loops: product recycling and material recycling.[68]
+The product or product parts are reused in the product recycling phase. This happens in one of two ways: the product is used retaining the product functionality ("reuse") or the product continues to be used but with altered functionality ("further use").[68] The product design is unmodified, or only slightly modified, in both scenarios.
+Product disassembly requires material recycling where product materials are recovered and recycled. Ideally, the materials are processed so they can flow back into the production process.[68]
+In order to meet recyclers' needs while providing manufacturers a consistent, uniform system, a coding system was developed. The recycling code for plastics was introduced in 1988 by the plastics industry through the Society of the Plastics Industry.[69] Because municipal recycling programs traditionally have targeted packaging—primarily bottles and containers—the resin coding system offered a means of identifying the resin content of bottles and containers commonly found in the residential waste stream.[70]
+Plastic products are printed with numbers 1–7 depending on the type of resin. Type 1 (polyethylene terephthalate) is commonly found in soft drink and water bottles. Type 2 (high-density polyethylene) is found in most hard plastics such as milk jugs, laundry detergent bottles, and some dishware. Type 3 (polyvinyl chloride) includes items such as shampoo bottles, shower curtains, hula hoops, credit cards, wire jacketing, medical equipment, siding, and piping. Type 4 (low-density polyethylene) is found in shopping bags, squeezable bottles, tote bags, clothing, furniture, and carpet. Type 5 is polypropylene and makes up syrup bottles, straws, Tupperware, and some automotive parts. Type 6 is polystyrene and makes up meat trays, egg cartons, clamshell containers, and compact disc cases. Type 7 includes all other plastics such as bulletproof materials, 3- and 5-gallon water bottles, cell phone and tablet frames, safety goggles and sunglasses.[71] Having a recycling code or the chasing arrows logo on a material is not an automatic indicator that a material is recyclable but rather an explanation of what the material is. Types 1 and 2 are the most commonly recycled.
+There is some debate over whether recycling is economically efficient. According to a Natural Resources Defense Council study, waste collection and landfill disposal creates less than one job per 1,000 tons of waste material managed; in contrast, the collection, processing, and manufacturing of recycled materials creates 6–13 or more jobs per 1,000 tons.[75] However, the cost effectiveness of creating the additional jobs remains unproven. According to the U.S. Recycling Economic Informational Study, there are over 50,000 recycling establishments that have created over a million jobs in the US.[76]
+Two years after New York City declared that implementing recycling programs would be "a drain on the city", New York City leaders realized that an efficient recycling system could save the city over $20 million.[77] Municipalities often see fiscal benefits from implementing recycling programs, largely due to the reduced landfill costs.[78] A study conducted by the Technical University of Denmark according to the Economist found that in 83 percent of cases, recycling is the most efficient method to dispose of household waste.[8][19] However, a 2004 assessment by the Danish Environmental Assessment Institute concluded that incineration was the most effective method for disposing of drink containers, even aluminium ones.[79]
+Fiscal efficiency is separate from economic efficiency. Economic analysis of recycling does not include what economists call externalities, which are unpriced costs and benefits that accrue to individuals outside of private transactions. Examples include: decreased air pollution and greenhouse gases from incineration, reduced hazardous waste leaching from landfills, reduced energy consumption, and reduced waste and resource consumption, which leads to a reduction in environmentally damaging mining and timber activity. About 4,000 minerals are known, of which only a few hundred are relatively common.[80] Known reserves of phosphorus will be exhausted within the next 100 years at current rates of usage.[81][82] Without mechanisms such as taxes or subsidies to internalize externalities, businesses may ignore them despite the costs imposed on society.[citation needed] To make such nonfiscal benefits economically relevant, advocates have pushed for legislative action to increase the demand for recycled materials.[5] The United States Environmental Protection Agency (EPA) has concluded in favor of recycling, saying that recycling efforts reduced the country's carbon emissions by a net 49 million metric tonnes in 2005.[8] In the United Kingdom, the Waste and Resources Action Programme stated that Great Britain's recycling efforts reduce CO2 emissions by 10–15 million tonnes a year.[8] Recycling is more efficient in densely populated areas, as there are economies of scale involved.[5]
+Certain requirements must be met for recycling to be economically feasible and environmentally effective. These include an adequate source of recyclates, a system to extract those recyclates from the waste stream, a nearby factory capable of reprocessing the recyclates, and a potential demand for the recycled products. These last two requirements are often overlooked—without both an industrial market for production using the collected materials and a consumer market for the manufactured goods, recycling is incomplete and in fact only "collection".[5]
+Free-market economist Julian Simon remarked "There are three ways society can organize waste disposal: (a) commanding, (b) guiding by tax and subsidy, and (c) leaving it to the individual and the market". These principles appear to divide economic thinkers today.[83]
+Frank Ackerman favours a high level of government intervention to provide recycling services. He believes that recycling's benefit cannot be effectively quantified by traditional laissez-faire economics. Allen Hershkowitz supports intervention, saying that it is a public service equal to education and policing. He argues that manufacturers should shoulder more of the burden of waste disposal.[83]
+Paul Calcott and Margaret Walls advocate the second option. A deposit refund scheme and a small refuse charge would encourage recycling but not at the expense of fly-tipping. Thomas C. Kinnaman concludes that a landfill tax would force consumers, companies and councils to recycle more.[83]
+Most free-market thinkers detest subsidy and intervention because they waste resources. Terry Anderson and Donald Leal think that all recycling programmes should be privately operated, and therefore would only operate if the money saved by recycling exceeds its costs. Daniel K. Benjamin argues that it wastes people's resources and lowers the wealth of a population.[83]
+The National Waste & Recycling Association (NWRA) then again reported in May 2015, that recycling and waste made a $6.7 billion economic impact in Ohio, U.S., and employed 14,000 people.[84]
+Certain countries trade in unprocessed recyclates. Some have complained that the ultimate fate of recyclates sold to another country is unknown and they may end up in landfills instead of being reprocessed. According to one report, in America, 50–80 percent of computers destined for recycling are actually not recycled.[85][86] There are reports of illegal-waste imports to China being dismantled and recycled solely for monetary gain, without consideration for workers' health or environmental damage. Although the Chinese government has banned these practices, it has not been able to eradicate them.[87] In 2008, the prices of recyclable waste plummeted before rebounding in 2009. Cardboard averaged about £53/tonne from 2004 to 2008, dropped to £19/tonne, and then went up to £59/tonne in May 2009. PET plastic averaged about £156/tonne, dropped to £75/tonne and then moved up to £195/tonne in May 2009.[88]
+Certain regions have difficulty using or exporting as much of a material as they recycle. This problem is most prevalent with glass: both Britain and the U.S. import large quantities of wine bottled in green glass. Though much of this glass is sent to be recycled, outside the American Midwest there is not enough wine production to use all of the reprocessed material. The extra must be downcycled into building materials or re-inserted into the regular waste stream.[5][8]
+Similarly, the northwestern United States has difficulty finding markets for recycled newspaper, given the large number of pulp mills in the region as well as the proximity to Asian markets. In other areas of the U.S., however, demand for used newsprint has seen wide fluctuation.[5]
+In some U.S. states, a program called RecycleBank pays people to recycle, receiving money from local municipalities for the reduction in landfill space which must be purchased. It uses a single stream process in which all material is automatically sorted.[89]
+Critics[who?] dispute the net economic and environmental benefits of recycling over its costs, and suggest that proponents of recycling often make matters worse and suffer from confirmation bias. Specifically, critics argue that the costs and energy used in collection and transportation detract from (and outweigh) the costs and energy saved in the production process; also that the jobs produced by the recycling industry can be a poor trade for the jobs lost in logging, mining, and other industries associated with production; and that materials such as paper pulp can only be recycled a few times before material degradation prevents further recycling.[90]
+Much of the difficulty inherent in recycling comes from the fact that most products are not designed with recycling in mind. The concept of sustainable design aims to solve this problem, and was laid out in the book Cradle to Cradle: Remaking the Way We Make Things by architect William McDonough and chemist Michael Braungart.[91] They suggest that every product (and all packaging it requires) should have a complete "closed-loop" cycle mapped out for each component—a way in which every component will either return to the natural ecosystem through biodegradation or be recycled indefinitely.[8][92]
+Complete recycling is impossible from a practical standpoint. In summary, substitution and recycling strategies only delay the depletion of non-renewable stocks and therefore may buy time in the transition to true or strong sustainability, which ultimately is only guaranteed in an economy based on renewable resources.[93]:21
+While recycling diverts waste from entering directly into landfill sites, current recycling misses the dispersive components. These critics believe that complete recycling is impracticable as highly dispersed wastes become so diluted that the energy needed for their recovery becomes increasingly excessive.
+As with environmental economics, care must be taken to ensure a complete view of the costs and benefits involved. For example, paperboard packaging for food products is more easily recycled than most plastic, but is heavier to ship and may result in more waste from spoilage.[94]
+The amount of energy saved through recycling depends upon the material being recycled and the type of energy accounting that is used. Correct accounting for this saved energy can be accomplished with life-cycle analysis using real energy values, and in addition, exergy, which is a measure of how much useful energy can be used. In general, it takes far less energy to produce a unit mass of recycled materials than it does to make the same mass of virgin materials.[95][96][97]
+Some scholars use emergy (spelled with an m) analysis, for example, budgets for the amount of energy of one kind (exergy) that is required to make or transform things into another kind of product or service. Emergy calculations take into account economics which can alter pure physics-based results. Using emergy life-cycle analysis researchers have concluded that materials with large refining costs have the greatest potential for high recycle benefits. Moreover, the highest emergy efficiency accrues from systems geared toward material recycling, where materials are engineered to recycle back into their original form and purpose, followed by adaptive reuse systems where the materials are recycled into a different kind of product, and then by-product reuse systems where parts of the products are used to make an entirely different product.[98]
+The Energy Information Administration (EIA) states on its website that "a paper mill uses 40 percent less energy to make paper from recycled paper than it does to make paper from fresh lumber."[99] Some critics argue that it takes more energy to produce recycled products than it does to dispose of them in traditional landfill methods, since the curbside collection of recyclables often requires a second waste truck. However, recycling proponents point out that a second timber or logging truck is eliminated when paper is collected for recycling, so the net energy consumption is the same. An emergy life-cycle analysis on recycling revealed that fly ash, aluminum, recycled concrete aggregate, recycled plastic, and steel yield higher efficiency ratios, whereas the recycling of lumber generates the lowest recycle benefit ratio. Hence, the specific nature of the recycling process, the methods used to analyse the process, and the products involved affect the energy savings budgets.[98]
+It is difficult to determine the amount of energy consumed or produced in waste disposal processes in broader ecological terms, where causal relations dissipate into complex networks of material and energy flow. For example, "cities do not follow all the strategies of ecosystem development. Biogeochemical paths become fairly straight relative to wild ecosystems, with very reduced recycling, resulting in large flows of waste and low total energy efficiencies. By contrast, in wild ecosystems, one population's wastes are another population's resources, and succession results in efficient exploitation of available resources. However, even modernized cities may still be in the earliest stages of a succession that may take centuries or millennia to complete."[100]:720 How much energy is used in recycling also depends on the type of material being recycled and the process used to do so. Aluminium is generally agreed to use far less energy when recycled rather than being produced from scratch. The EPA states that "recycling aluminum cans, for example, saves 95 percent of the energy required to make the same amount of aluminum from its virgin source, bauxite."[101][102] In 2009, more than half of all aluminium cans produced came from recycled aluminium.[103] Similarly, it has been estimated that new steel produced with recycled cans reduces greenhouse gas emissions by 75%.[104]
+Every year, millions of tons of materials are being exploited from the earth's crust, and processed into consumer and capital goods. After decades to centuries, most of these materials are "lost". With the exception of some pieces of art or religious relics, they are no longer engaged in the consumption process. Where are they? Recycling is only an intermediate solution for such materials, although it does prolong the residence time in the anthroposphere. For thermodynamic reasons, however, recycling cannot prevent the final need for an ultimate sink.[105]:1
+Economist Steven Landsburg has suggested that the sole benefit of reducing landfill space is trumped by the energy needed and resulting pollution from the recycling process.[106] Others, however, have calculated through life-cycle assessment that producing recycled paper uses less energy and water than harvesting, pulping, processing, and transporting virgin trees.[107] When less recycled paper is used, additional energy is needed to create and maintain farmed forests until these forests are as self-sustainable as virgin forests.
+Other studies have shown that recycling in itself is inefficient to perform the "decoupling" of economic development from the depletion of non-renewable raw materials that is necessary for sustainable development.[108] The international transportation or recycle material flows through "... different trade networks of the three countries result in different flows, decay rates, and potential recycling returns".[109]:1 As global consumption of a natural resources grows, their depletion is inevitable. The best recycling can do is to delay; complete closure of material loops to achieve 100 percent recycling of nonrenewables is impossible as micro-trace materials dissipate into the environment causing severe damage to the planet's ecosystems.[110][111][112] Historically, this was identified as the metabolic rift by Karl Marx, who identified the unequal exchange rate between energy and nutrients flowing from rural areas to feed urban cities that create effluent wastes degrading the planet's ecological capital, such as loss in soil nutrient production.[113][114] Energy conservation also leads to what is known as Jevon's paradox, where improvements in energy efficiency lowers the cost of production and leads to a rebound effect where rates of consumption and economic growth increases.[112][115]
+The amount of money actually saved through recycling depends on the efficiency of the recycling program used to do it. The Institute for Local Self-Reliance argues that the cost of recycling depends on various factors, such as landfill fees and the amount of disposal that the community recycles. It states that communities begin to save money when they treat recycling as a replacement for their traditional waste system rather than an add-on to it and by "redesigning their collection schedules and/or trucks".[116]
+In some cases, the cost of recyclable materials also exceeds the cost of raw materials. Virgin plastic resin costs 40 percent less than recycled resin.[117] Additionally, a United States Environmental Protection Agency (EPA) study that tracked the price of clear glass from 15 July to 2 August 1991, found that the average cost per ton ranged from $40 to $60[118] while a USGS report shows that the cost per ton of raw silica sand from years 1993 to 1997 fell between $17.33 and $18.10.[119]
+Comparing the market cost of recyclable material with the cost of new raw materials ignores economic externalities—the costs that are currently not counted by the market. Creating a new piece of plastic, for instance, may cause more pollution and be less sustainable than recycling a similar piece of plastic, but these factors will not be counted in market cost. A life cycle assessment can be used to determine the levels of externalities and decide whether the recycling may be worthwhile despite unfavorable market costs. Alternatively, legal means (such as a carbon tax) can be used to bring externalities into the market, so that the market cost of the material becomes close to the true cost.
+The recycling of waste electrical and electronic equipment can create a significant amount of pollution. This problem is specifically occurrent in India and China. Informal recycling in an underground economy of these countries has generated an environmental and health disaster. High levels of lead (Pb), polybrominated diphenylethers (PBDEs), polychlorinated dioxins and furans, as well as polybrominated dioxins and furans (PCDD/Fs and PBDD/Fs), concentrated in the air, bottom ash, dust, soil, water, and sediments in areas surrounding recycling sites.[120] These materials can make work sites harmful to the workers themselves and the surrounding environment.
+Economist Steven Landsburg, author of a paper entitled "Why I Am Not an Environmentalist",[121] claimed that paper recycling actually reduces tree populations. He argues that because paper companies have incentives to replenish their forests, large demands for paper lead to large forests while reduced demand for paper leads to fewer "farmed" forests.[122]
+When foresting companies cut down trees, more are planted in their place; however, such "farmed" forests are inferior to natural forests in several ways. Farmed forests are not able to fix the soil as quickly as natural forests. This can cause widespread soil erosion and often requiring large amounts of fertilizer to maintain the soil, while containing little tree and wild-life biodiversity compared to virgin forests.[123] Also, the new trees planted are not as big as the trees that were cut down, and the argument that there will be "more trees" is not compelling to forestry advocates when they are counting saplings.
+In particular, wood from tropical rainforests is rarely harvested for paper because of their heterogeneity.[124] According to the United Nations Framework Convention on Climate Change secretariat, the overwhelming direct cause of deforestation is subsistence farming (48% of deforestation) and commercial agriculture (32%), which is linked to food, not paper production.[125]
+The reduction of greenhouse gas emission reduction also benefits from the development of the recycling industry. In Kitakyushu, the only green growth model city in Asia selected by OECD, recycling industries are strongly promoted and financially supported as part of the Eco-town program in Japan. Given the industrial sector in Kitakyushu accounts for more than 60% energy consumption of the city, the development of recycling industry results in substantial energy reduction due to the economies of scale effects; the concentration of CO is, thus, found to decline accordingly.[126]
+Other non-conventional methods of material recycling, like Waste-to-Energy (WTE) systems, have garnered increased attention in the recent past due to the polarizing nature of their emissions. While viewed as a sustainable method of capturing energy from material waste feedstocks by many, others have cited numerous explanations for why the technology has not been scaled globally.[127]
+In some countries, recycling is performed by the entrepreneurial poor such as the karung guni, zabbaleen, the rag-and-bone man, waste picker, and junk man. With the creation of large recycling organizations that may be profitable, either by law or economies of scale,[128][129] the poor are more likely to be driven out of the recycling and the remanufacturing job market. To compensate for this loss of income, a society may need to create additional forms of societal programs to help support the poor.[130] Like the parable of the broken window, there is a net loss to the poor and possibly the whole of a society to make recycling artificially profitable, e.g. through the law. However, in Brazil and Argentina, waste pickers/informal recyclers work alongside the authorities, in fully or semi-funded cooperatives, allowing informal recycling to be legitimized as a paid public sector job.[131]
+Because the social support of a country is likely to be less than the loss of income to the poor undertaking recycling, there is a greater chance the poor will come in conflict with the large recycling organizations.[132][133] This means fewer people can decide if certain waste is more economically reusable in its current form rather than being reprocessed. Contrasted to the recycling poor, the efficiency of their recycling may actually be higher for some materials because individuals have greater control over what is considered "waste".[130]
+One labor-intensive underused waste is electronic and computer waste. Because this waste may still be functional and wanted mostly by those on lower incomes, who may sell or use it at a greater efficiency than large recyclers.
+Some recycling advocates believe that laissez-faire individual-based recycling does not cover all of society's recycling needs. Thus, it does not negate the need for an organized recycling program.[130] Local government can consider the activities of the recycling poor as contributing to the ruining of property.
+Changes that have been demonstrated to increase recycling rates include:
+"Between 1960 and 2000, the world production of plastic resins increased 25 times its original amount, while recovery of the material remained below 5 percent."[134]:131 Many studies have addressed recycling behaviour and strategies to encourage community involvement in recycling programs. It has been argued[135] that recycling behavior is not natural because it requires a focus and appreciation for long-term planning, whereas humans have evolved to be sensitive to short-term survival goals; and that to overcome this innate predisposition, the best solution would be to use social pressure to compel participation in recycling programs. However, recent studies have concluded that social pressure will not work in this context.[136] One reason for this is that social pressure functions well in small group sizes of 50 to 150 individuals (common to nomadic hunter–gatherer peoples) but not in communities numbering in the millions, as we see today. Another reason is that individual recycling does not take place in the public view.
+Following the increasing popularity of recycling collection being sent to the same landfills as trash, some people kept on putting recyclables on the recyclables bin.[137] In Baltimore, the government kept collecting glass separately for seven years even though it did not recycle it.[138]
+Art objects are more and more often made from recycled material.
+In a study done by social psychologist Shawn Burn,[139] it was found that personal contact with individuals within a neighborhood is the most effective way to increase recycling within a community. In his study, he had 10 block leaders talk to their neighbors and persuade them to recycle. A comparison group was sent fliers promoting recycling. It was found that the neighbors that were personally contacted by their block leaders recycled much more than the group without personal contact. As a result of this study, Shawn Burn believes that personal contact within a small group of people is an important factor in encouraging recycling. Another study done by Stuart Oskamp[140] examines the effect of neighbors and friends on recycling. It was found in his studies that people who had friends and neighbors that recycled were much more likely to also recycle than those who didn't have friends and neighbors that recycled.
+Many schools have created recycling awareness clubs in order to give young students an insight on recycling. These schools believe that the clubs actually encourage students to not only recycle at school but at home as well.

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+All other species in Canini
+Foxes are small to medium-sized, omnivorous mammals belonging to several genera of the family Canidae. Foxes have a flattened skull, upright triangular ears, a pointed, slightly upturned snout, and a long bushy tail (or brush).
+Twelve species belong to the monophyletic "true foxes" group of genus Vulpes. Approximately another 25 current or extinct species are always or sometimes called foxes; these foxes are either part of the paraphyletic group of the South American foxes, or of the outlying group, which consists of the bat-eared fox, gray fox, and island fox.[1] Foxes live on every continent except Antarctica. By far the most common and widespread species of fox is the red fox (Vulpes vulpes) with about 47 recognized subspecies.[2] The global distribution of foxes, together with their widespread reputation for cunning, has contributed to their prominence in popular culture and folklore in many societies around the world. The hunting of foxes with packs of hounds, long an established pursuit in Europe, especially in the British Isles, was exported by European settlers to various parts of the New World.
+The word fox comes from Old English, which derived from Proto-Germanic *fuhsaz.[nb 1] This in turn derives from Proto-Indo-European *puḱ-, meaning ’thick-haired; tail’.[nb 2] Male foxes are known as dogs, tods or reynards, females as vixens, and young as cubs, pups, or kits, though the latter name is not to be confused with a distinct species called kit foxes. Vixen is one of very few words in modern English that retains the Middle English southern dialect "v" pronunciation instead of "f" (i.e. northern English "fox" versus southern English "vox").[3] A group of foxes is referred to as a skulk, leash, or earth.[4][5]
+Within the Canidae, the results of DNA analysis shows  several phylogenetic divisions:
+Foxes are generally smaller than some other members of the family Canidae such as wolves and jackals, while they may be larger than some within the family, such as Raccoon dogs. In the largest species, the red fox, males weigh on average between 4.1 and 8.7 kilograms (9 and 19 1⁄4 pounds),[7] while the smallest species, the fennec fox, weighs just 0.7 to 1.6 kg (1 1⁄2 to 3 1⁄2 lb).[8] Foxy features typically include a triangular face, pointed ears, an elongated rostrum, and a bushy tail. Foxes are digitigrade; they walk on their toes. Unlike most members of the family Canidae, foxes have partially retractable claws.[9] Fox vibrissae, or whiskers, are black. The whiskers on the muzzle, mystaciae vibrissae, average 100–110 millimetres (3 7⁄8–4 3⁄8 inches) long, while the whiskers everywhere else on the head average to be shorter in length. Whiskers (carpal vibrissae) are also on the forelimbs and average 40 mm (1 5⁄8 in) long, pointing downward and backward.[2] Other physical characteristics vary according to habitat and adaptive significance.
+Fox species differ in fur color, length, and density. Coat colors range from pearly white to black-and-white to black flecked with white or grey on the underside. Fennec foxes (and other species of fox adapted to life in the desert, such as kit foxes), for example, have large ears and short fur to aid in keeping the body cool.[2][9] Arctic foxes, on the other hand, have tiny ears and short limbs as well as thick, insulating fur, which aid in keeping the body warm.[10] Red foxes, by contrast, have a typical auburn pelt, the tail normally ending with a white marking.[11] A fox's coat color and texture may vary due to the change in seasons; fox pelts are richer and denser in the colder months and lighter in the warmer months. To get rid of the dense winter coat, foxes moult once a year around April; the process begins from the feet, up the legs, and then along the back.[9] Coat color may also change as the individual ages.[2]
+A fox's dentition, like all other canids, is I 3/3, C 1/1, PM 4/4, M 3/2 = 42. (Bat-eared foxes have six extra molars, totalling in 48 teeth.) Foxes have pronounced carnassial pairs, which is characteristic of a carnivore. These pairs consist of the upper premolar and the lower first molar, and work together to shear tough material like flesh. Foxes' canines are pronounced, also characteristic of a carnivore, and are excellent in gripping prey.[12]
+In the wild, the typical lifespan of a fox is one to three years, although individuals may live up to ten years. Unlike many canids, foxes are not always pack animals. Typically, they live in small family groups, but some (such as Arctic foxes) are known to be solitary.[2][9]
+Foxes are omnivores.[13][14] Their diet is made up primarily of invertebrates such as insects and small vertebrates such as reptiles and birds. They may also eat eggs and vegetation. Many species are generalist predators, but some (such as the crab-eating fox) have more specialized diets. Most species of fox consume around 1 kg (2.2 lb) of food every day. Foxes cache excess food, burying it for later consumption, usually under leaves, snow, or soil.[9][15] While hunting, foxes tend to use a particular pouncing technique, such that they crouch down to camouflage themselves in the terrain and then use their hind legs to leap up with great force and land on top of their chosen prey.[2] Using their pronounced canine teeth, they can then grip the prey's neck and shake it until it is dead or can be readily disemboweled.[2]
+The gray fox is one of only two canine species known to regularly climb trees; the other is the raccoon dog.[16]
+The male fox's scrotum is held up close to the body with the testes inside even after they descend. Like other canines, the male fox has a baculum, or penile bone.[2][17][18] The testes of red foxes are smaller than those of Arctic foxes.[19] Sperm formation in red foxes begins in August–September, with the testicles attaining their greatest weight in December–February.[20]
+Vixens are in heat for one to six days, making their reproductive cycle twelve months long. As with other canines, the ova are shed during estrus without the need for the stimulation of copulating. Once the egg is fertilized, the vixen enters a period of gestation that can last from 52 to 53 days. Foxes tend to have an average litter size of four to five with an 80 percent success rate in becoming pregnant.[2][21] Litter sizes can vary greatly according to species and environment – the Arctic fox, for example, can have up to eleven kits.[22]
+The vixen usually has six or eight mammae.[23] Each teat has 8 to 20 lactiferous ducts, which connect the mammary gland to the nipple, allowing for milk to be carried to the nipple.[citation needed]
+The fox's vocal repertoire is vast:
+In the case of domesticated foxes, the whining seems to remain in adult individuals as a sign of excitement and submission in the presence of their owners.[2]
+Canids commonly known as foxes include the following genera and species:[2]
+Several fox species are endangered in their native environments. Pressures placed on foxes include habitat loss and being hunted for pelts, other trade, or control.[25] Due in part to their opportunistic hunting style and industriousness, foxes are commonly resented as nuisance animals.[26] On the other hand, foxes, while often considered pests themselves, have been successfully employed to control pests on fruit farms while leaving the fruit intact.[27]
+The island fox, though considered a near-threatened species throughout the world, is becoming increasingly endangered in its endemic environment of the California Channel Islands.[28] A population on an island is smaller than those on the mainland because of limited resources like space, food and shelter.[29] Island populations, therefore, are highly susceptible to external threats ranging from introduced predatory species and humans to extreme weather.[29] On the California Channel Islands, it was found that the population of the island fox was so low due to an outbreak of canine distemper virus from 1999 to 2000[30] as well as predation by non-native golden eagles.[31] Since 1993, the eagles have caused the population to decline by as much as 95%.[30] Because of the low number of foxes, the population went through an Allee effect; this is where at low enough densities, an individual's fitness decreases.[28] Conservationists, therefore, had to take healthy breeding pairs out of the wild population to breed them in captivity until they had enough foxes to release back into the wild.[30] Nonnative grazers were also removed so that native plants would be able to grow back to their natural height, thereby providing adequate cover and protection for the foxes against golden eagles.[31]
+Darwin's fox is considered critically endangered because of their small known population of 250 mature individuals as well as their restricted distribution.[32] On the Chilean mainland, the population is limited to Nahuelbuta National Park and the surrounding Valdivian rainforest.[32] Similarly on Chiloé Island, their population is limited to the forests that extend from the southernmost to the northwestern most part of the island.[32] Though the Nahuelbuta National Park is protected, 90% of the species live on Chiloé Island.[33] A major problem the species faces, therefore, is their dwindling, limited habitat due to the cutting and burning of the unprotected forests.[32] Because of deforestation, the Darwin's fox habitat is shrinking, allowing for their competitor's (chilla fox) preferred habitat of open space, to increase; the Darwin's fox, subsequently, is being outcompeted.[34] Another problem they face is their inability to fight off diseases transmitted by the increasing number of pet dogs.[32] To conserve these animals, researchers suggest the need for the forests that link the Nahuelbuta National Park to the coast of Chile and in turn Chiloé Island and its forests, to be protected.[34] They also suggest that other forests around Chile be examined to determine whether Darwin's foxes have previously existed there or can live there in the future, should the need to reintroduce the species to those areas arise.[34] And finally, the researchers advise for the creation of a captive breeding program, in Chile, because of the limited number of mature individuals in the wild.[34]
+Foxes are often considered pests or nuisance creatures for their opportunistic attacks on poultry and other small livestock. Fox attacks on humans are not common.[35]
+Many foxes adapt well to human environments, with several species classified as "resident urban carnivores" for their ability to sustain populations entirely within urban boundaries.[36] Foxes in urban areas can live longer and can have smaller litter sizes than foxes in non-urban areas.[36] Urban foxes are ubiquitous in Europe, where they show altered behaviors compared to non-urban foxes, including increased population density, smaller territory, and pack foraging.[37] Foxes have been introduced in numerous locations, with varying effects on indigenous flora and fauna.[38]
+In some countries, foxes are major predators of rabbits and hens. Population oscillations of these two species were the first nonlinear oscillation studied, and led to the derivation of the Lotka–Volterra equation.[39][40]
+Fox hunting originated in the United Kingdom in the 16th century. Hunting with dogs is now banned in the United Kingdom,[41][42][43][44] though hunting without dogs is still permitted. Red foxes were introduced into Australia in the early 19th century for sport, and have since become widespread through much of the country. They have caused population decline among many native species and prey on livestock, especially new lambs.[45] Fox hunting is practiced as recreation in several other countries including Canada, France, Ireland, Italy, Russia, United States and Australia.
+There are many records of domesticated red foxes and others, but rarely of sustained domestication. A recent and notable exception is the Russian silver fox,[46] which resulted in visible and behavioral changes, and is a case study of an animal population modeling according to human domestication needs. The current group of domesticated silver foxes are the result of nearly fifty years of experiments in the Soviet Union and Russia to domesticate the silver morph of the red fox. This selective breeding resulted in physical and behavioral traits appearing that are frequently seen in domestic cats, dogs, and other animals, such as pigmentation changes, floppy ears, and curly tails.[47] Notably, the new foxes became more tame, allowing themselves to be petted, whimpering to get attention and sniffing and licking their caretakers.[48]
+In the United Kingdom, a number of cases of non-fatal attacks on humans were reported. They often involved children, or if there were gaps in homes through which foxes could pass.[49]
+Foxes are among the comparatively few mammals which have been able to adapt themselves to a certain degree to living in urban (mostly suburban) human environments. Their omnivorous diet allows them to survive on discarded food waste, and their skittish and often nocturnal nature means that they are often able to avoid detection, despite their larger size. Urban foxes, however, have been identified as threats to cats and small dogs, and for this reason there is often pressure to exclude them from these environments.[50]
+The San Joaquin kit fox is a highly endangered species that has, ironically, become adapted to urban living in the San Joaquin Valley and Salinas Valley of southern California. Its diet includes mice, ground squirrels, rabbits, hares, bird eggs, and insects, and it has claimed habitats in open areas, golf courses, drainage basins, and school grounds.[50]
+The fox appears in many cultures, usually in folklore. However, there are slight variations in their depictions in folklore. In Western folklore and also in Persian folklore, foxes are depicted as a symbol of cunning and trickery – a reputation derived especially from their reputed ability to evade hunters. This is usually represented as a character possessing these traits. These traits are used on a wide variety of characters, either making them a nuisance to the story, a misunderstood hero, or a devious villain.
+In Asian folklore, foxes are depicted as a familiar spirit possessed of magic powers. Similar to Western folklore, foxes are depicted as mischievous, usually tricking other people, with the ability to disguise as an attractive female human. However, there are other depictions of foxes as a mystical, sacred creature, that can either bring wonder or ruin.[51] Nine-tailed foxes appear in Chinese folklore, literature, and mythology, in which, depending on the tale can be a good or a bad omen.[52] The motif was eventually introduced from Chinese to Japanese and Korean cultures.[53]
+The constellation Vulpecula represents a fox.[54]

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+The great white shark (Carcharodon carcharias), also known as the great white, white shark or "white pointer", is a species of large mackerel shark which can be found in the coastal surface waters of all the major oceans. It is notable for its size, with larger female individuals growing to 6.1 m (20 ft) in length and 1,905–2,268 kg (4,200–5,000 lb) in weight at maturity.[3][4][5] However, most are smaller; males measure 3.4 to 4.0 m (11 to 13 ft), and females measure 4.6 to 4.9 m (15 to 16 ft) on average.[4][6] According to a 2014 study, the lifespan of great white sharks is estimated to be as long as 70 years or more, well above previous estimates,[7] making it one of the longest lived cartilaginous fishes currently known.[8] According to the same study, male great white sharks take 26 years to reach sexual maturity, while the females take 33 years to be ready to produce offspring.[9] Great white sharks can swim at speeds of 25 km/hr (16 mph)[10] for short bursts and to depths of 1,200 m (3,900 ft).[11]
+The great white shark has no known natural predators other than, on very rare occasions, the killer whale.[12] It is arguably the world's largest-known extant macropredatory fish, and is one of the primary predators of marine mammals, up to the size of large baleen whales. This shark is also known to prey upon a variety of other marine animals, including fish, and seabirds. It is the only known surviving species of its genus Carcharodon, and is responsible for more recorded human bite incidents than any other shark.[13][14]
+The species faces numerous ecological challenges which has resulted in international protection. The IUCN lists the great white shark as a vulnerable species,[2] and it is included in Appendix II of CITES.[15] It is also protected by several national governments such as Australia (as of 2018).[16]
+The novel Jaws by Peter Benchley and its subsequent film adaptation by Steven Spielberg depicted the great white shark as a "ferocious man-eater". Humans are not the preferred prey of the great white shark,[17] but the great white is nevertheless responsible for the largest number of reported and identified fatal unprovoked shark attacks on humans.[18]
+Due to their need to travel long distances for seasonal migration and extremely demanding diet, it is not logistically feasible to keep great white sharks in captivity. No aquarium in the world is currently believed to own one.[19]
+The great white shark was one of the many amphibia originally described by Carl Linnaeus in his landmark 1758 10th edition of Systema Naturae,[20] with its first scientific name, Squalus carcharias. Later, Sir Andrew Smith gave it Carcharodon as its generic name in 1833, and also in 1873. The generic name was identified with Linnaeus' specific name and the current scientific name, Carcharodon carcharias, was finalized. Carcharodon comes from the Ancient Greek words κάρχαρος (kárkharos, 'sharp' or 'jagged'), and ὀδούς (odoús), ὀδών (odṓn, 'tooth').[21]
+The earliest-known fossils of the great white shark are about 16 million years old, during the mid-Miocene epoch.[1] However, the phylogeny of the great white is still in dispute. The original hypothesis for the great white's origins is that it shares a common ancestor with a prehistoric shark, such as the C. megalodon. C. megalodon had teeth that were superficially not too dissimilar with those of great white sharks, but its teeth were far larger. Although cartilaginous skeletons do not fossilize, C. megalodon is estimated to have been considerably larger than the great white shark, estimated at up to 17 m (56 ft) and 59,413 kg (130,983 lb).[22] Similarities among the physical remains and the extreme size of both the great white and C. megalodon led many scientists to believe these sharks were closely related, and the name Carcharodon megalodon was applied to the latter.
+However, a new hypothesis proposes that the great white is also more closely related to an ancient mako shark, Isurus hastalis, than to the C. megalodon. The theory seems to be supported with the earlier discovery of a complete set of jaws with 222 teeth and 45 vertebrae of the extinct transitional species Carcharodon hubbelli in 1988.[23] In addition, the new hypothesis assigns C. megalodon to the genus Carcharocles, which also comprises the other megatoothed sharks.[24]
+Great white sharks live in almost all coastal and offshore waters which have water temperature between 12 and 24 °C (54 and 75 °F), with greater concentrations in the United States (Northeast and California), South Africa, Japan, Oceania, Chile, and the Mediterranean including Sea of Marmara and Bosphorus.[25][26] One of the densest-known populations is found around Dyer Island, South Africa.[27]
+The great white is an epipelagic fish, observed mostly in the presence of rich game, such as fur seals (Arctocephalus ssp.), sea lions, cetaceans, other sharks, and large bony fish species. In the open ocean, it has been recorded at depths as great as 1,200��m (3,900 ft).[11] These findings challenge the traditional notion that the great white is a coastal species.[11]
+According to a recent study, California great whites have migrated to an area between Baja California Peninsula and Hawaii known as the White Shark Café to spend at least 100 days before migrating back to Baja. On the journey out, they swim slowly and dive down to around 900 m (3,000 ft). After they arrive, they change behaviour and do short dives to about 300 m (980 ft) for up to ten minutes. Another white shark that was tagged off the South African coast swam to the southern coast of Australia and back within the year. A similar study tracked a different great white shark from South Africa swimming to Australia's northwestern coast and back, a journey of 20,000 km (12,000 mi; 11,000 nmi) in under nine months.[28]
+These observations argue against traditional theories that white sharks are coastal territorial predators, and open up the possibility of interaction between shark populations that were previously thought to have been discrete. The reasons for their migration and what they do at their destination is still unknown. Possibilities include seasonal feeding or mating.[29]
+In the Northwest Atlantic the white shark populations off the New England coast were nearly eradicated due to over-fishing.[30] However, in recent years the populations have begun to grow greatly,[31] largely due to the increase in seal populations on Cape Cod, Massachusetts since the enactment of the Marine Mammal Protection Act in 1972.[32] Currently very little is known about the hunting and movement patterns of great whites off Cape Cod, but ongoing studies hope to offer insight into this growing shark population.[33]
+A 2018 study indicated that white sharks prefer to congregate deep in anticyclonic eddies in the North Atlantic Ocean. The sharks studied tended to favour the warm water eddies, spending the daytime hours at 450 meters and coming to the surface at night.[34]
+The great white shark has a robust, large, conical snout. The upper and lower lobes on the tail fin are approximately the same size which is similar to some mackerel sharks. A great white displays countershading, by having a white underside and a grey dorsal area (sometimes in a brown or blue shade) that gives an overall mottled appearance. The coloration makes it difficult for prey to spot the shark because it breaks up the shark's outline when seen from the side. From above, the darker shade blends with the sea and from below it exposes a minimal silhouette against the sunlight. Leucism is extremely rare in this species, but has been documented in one great white shark (a pup that washed ashore in Australia and died).[35] Great white sharks, like many other sharks, have rows of serrated teeth behind the main ones, ready to replace any that break off. When the shark bites, it shakes its head side-to-side, helping the teeth saw off large chunks of flesh.[36] Great white sharks, like other mackerel sharks, have larger eyes than other shark species in proportion to their body size. The iris of the eye is a deep blue instead of black.[37]
+In great white sharks, sexual dimorphism is present, and females are generally larger than males. Male great whites on average measure 3.4 to 4.0 m (11 to 13 ft) long, while females at 4.6 to 4.9 m (15 to 16 ft).[6] Adults of this species weigh 522–771 kg (1,151–1,700 lb) on average;[40] however, mature females can have an average mass of 680–1,110 kg (1,500–2,450 lb).[4] The largest females have been verified up to 6.1 m (20 ft) in length and an estimated 1,905 kg (4,200 lb) in weight,[3][4] perhaps up to 2,268 kg (5,000 lb).[5] The maximum size is subject to debate because some reports are rough estimations or speculations performed under questionable circumstances.[41] Among living cartilaginous fish, only the whale shark (Rhincodon typus), the basking shark (Cetorhinus maximus) and the giant manta ray (Manta birostris), in that order, are on average larger and heavier. These three species are generally quite docile in disposition and given to passively filter-feeding on very small organisms.[40] This makes the great white shark the largest extant macropredatory fish. Great white sharks are at around 1.2 m (3.9 ft) when born, and grow about 25 cm (9.8 in) each year.[42]
+According to J. E. Randall, the largest white shark reliably measured was a 6 m (20 ft) individual reported from Ledge Point, Western Australia in 1987.[43] Another great white specimen of similar size has been verified by the Canadian Shark Research Center: A female caught by David McKendrick of Alberton, Prince Edward Island, in August 1988 in the Gulf of St. Lawrence off Prince Edward Island. This female great white was 6.1 m (20 ft) long.[4] However, there was a report considered reliable by some experts in the past, of a larger great white shark specimen from Cuba in 1945.[39][44][45][46] This specimen was reportedly 6.4 m (21 ft) long and had a body mass estimated at 3,324 kg (7,328 lb).[39][45] However, later studies also revealed that this particular specimen was actually around 4.9 m (16 ft) in length, a specimen in the average maximum size range.[4]
+The largest great white recognized by the International Game Fish Association (IGFA) is one caught by Alf Dean in south Australian waters in 1959, weighing 1,208 kg (2,663 lb).[41] Several larger great whites caught by anglers have since been verified, but were later disallowed from formal recognition by IGFA monitors for rules violations.
+A number of very large unconfirmed great white shark specimens have been recorded.[47] For decades, many ichthyological works, as well as the Guinness Book of World Records, listed two great white sharks as the largest individuals: In the 1870s, a 10.9 m (36 ft) great white captured in southern Australian waters, near Port Fairy, and an 11.3 m (37 ft) shark trapped in a herring weir in New Brunswick, Canada, in the 1930s. However, these measurements were not obtained in a rigorous, scientifically valid manner, and researchers have questioned the reliability of these measurements for a long time, noting they were much larger than any other accurately reported sighting. Later studies proved these doubts to be well founded. This New Brunswick shark may have been a misidentified basking shark, as the two have similar body shapes. The question of the Port Fairy shark was settled in the 1970s when J. E. Randall examined the shark's jaws and "found that the Port Fairy shark was of the order of 5 m (16 ft) in length and suggested that a mistake had been made in the original record, in 1870, of the shark's length".[43] These wrong measurements would make the alleged shark more than five times heavier than it really was.
+While these measurements have not been confirmed, some great white sharks caught in modern times have been estimated to be more than 7 m (23 ft) long,[48] but these claims have received some criticism.[41][48] However, J. E. Randall believed that great white shark may have exceeded 6.1 m (20 ft) in length.[43] A great white shark was captured near Kangaroo Island in Australia on 1 April 1987. This shark was estimated to be more than 6.9 m (23 ft) long by Peter Resiley,[43][49] and has been designated as KANGA.[48] Another great white shark was caught in Malta by Alfredo Cutajar on 16 April 1987. This shark was also estimated to be around 7.13 m (23.4 ft) long by John Abela and has been designated as MALTA.[48] However, Cappo drew criticism because he used shark size estimation methods proposed by J. E. Randall to suggest that the KANGA specimen was 5.8–6.4 m (19–21 ft) long.[48] In a similar fashion, I. K. Fergusson also used shark size estimation methods proposed by J. E. Randall to suggest that the MALTA specimen was 5.3–5.7 m (17–19 ft) long.[48] However, photographic evidence suggested that these specimens were larger than the size estimations yielded through Randall's methods.[48] Thus, a team of scientists—H. F. Mollet, G. M. Cailliet, A. P. Klimley, D. A. Ebert, A. D. Testi, and L. J. V. Compagno—reviewed the cases of the KANGA and MALTA specimens in 1996 to resolve the dispute by conducting a comprehensive morphometric analysis of the remains of these sharks and re-examination of photographic evidence in an attempt to validate the original size estimations and their findings were consistent with them. The findings indicated that estimations by P. Resiley and J. Abela are reasonable and could not be ruled out.[48] A particularly large female great white nicknamed "Deep Blue", estimated measuring at 6.1 m (20 ft) was filmed off Guadalupe during shooting for the 2014 episode of Shark Week "Jaws Strikes Back". Deep Blue would also later gain significant attention when she was filmed interacting with researcher Mauricio Hoyas Pallida in a viral video that Mauricio posted on Facebook on 11 June 2015.[50] Deep Blue was later seen off Oahu in January 2019 while scavenging a sperm whale carcass, whereupon she was filmed swimming beside divers including dive tourism operator and model Ocean Ramsey in open water.[51][52][53] In July 2019, a fisherman, J. B. Currell, was on a trip to Cape Cod from Bermuda with Tom Brownell when they saw a large shark about 40 mi (64 km) southeast of Martha's Vineyard. Recording it on video, he said that it weighed about 5,000 lb (2,300 kg), and measured 25–30 ft (7.6–9.1 m), evoking a comparison with the fictional shark Jaws. The video was shared with the page "Troy Dando Fishing" on Facebook.[54][55] A particularly infamous great white shark, supposedly of record proportions, once patrolled the area that comprises False Bay, South Africa, was said to be well over 7 m (23 ft) during the early 1980s. This shark, known locally as the "Submarine", had a legendary reputation that was supposedly well founded. Though rumours have stated this shark was exaggerated in size or non-existent altogether, witness accounts by the then young Craig Anthony Ferreira, a notable shark expert in South Africa, and his father indicate an unusually large animal of considerable size and power (though it remains uncertain just how massive the shark was as it escaped capture each time it was hooked). Ferreira describes the four encounters with the giant shark he participated in with great detail in his book "Great White Sharks On Their Best Behavior".[56]
+One contender in maximum size among the predatory sharks is the tiger shark (Galeocerdo cuvier). While tiger sharks which are typically both a few feet smaller and have a leaner, less heavy body structure than white sharks, have been confirmed to reach at least 5.5 m (18 ft) in the length, an unverified specimen was reported to have measured 7.4 m (24 ft) in length and weighed 3,110 kg (6,860 lb), more than two times heavier than the largest confirmed specimen at 1,524 kg (3,360 lb).[40][57][58] Some other macropredatory sharks such as the Greenland shark (Somniosus microcephalus) and the Pacific sleeper shark (S. pacificus) are also reported to rival these sharks in length (but probably weigh a bit less since they are more slender in build than a great white) in exceptional cases.[59][60] The question of maximum weight is complicated by the unresolved question of whether or not to include the shark's stomach contents when weighing the shark. With a single bite a great white can take in up to 14 kg (31 lb) of flesh and can also consume several hundred kilograms of food.
+Great white sharks, like all other sharks, have an extra sense given by the ampullae of Lorenzini which enables them to detect the electromagnetic field emitted by the movement of living animals. Great whites are so sensitive they can detect variations of half a billionth of a volt. At close range, this allows the shark to locate even immobile animals by detecting their heartbeat. Most fish have a less-developed but similar sense using their body's lateral line.[75]
+To more successfully hunt fast and agile prey such as sea lions, the great white has adapted to maintain a body temperature warmer than the surrounding water. One of these adaptations is a "rete mirabile" (Latin for "wonderful net"). This close web-like structure of veins and arteries, located along each lateral side of the shark, conserves heat by warming the cooler arterial blood with the venous blood that has been warmed by the working muscles. This keeps certain parts of the body (particularly the stomach) at temperatures up to 14 °C (25 °F) [76] above that of the surrounding water, while the heart and gills remain at sea temperature. When conserving energy, the core body temperature can drop to match the surroundings. A great white shark's success in raising its core temperature is an example of gigantothermy. Therefore, the great white shark can be considered an endothermic poikilotherm or mesotherm because its body temperature is not constant but is internally regulated.[36][77] Great whites also rely on the fat and oils stored within their livers for long-distance migrations across nutrient-poor areas of the oceans.[78] Studies by Stanford University and the Monterey Bay Aquarium published on 17 July 2013 revealed that in addition to controlling the sharks' buoyancy, the liver of great whites is essential in migration patterns. Sharks that sink faster during drift dives were revealed to use up their internal stores of energy quicker than those which sink in a dive at more leisurely rates.[79]
+Toxicity from heavy metals seems to have little negative effects on great white sharks. Blood samples taken from forty-three individuals of varying size, age and sex off the South African coast led by biologists from the University of Miami in 2012 indicates that despite high levels of mercury, lead, and arsenic, there was no sign of raised white blood cell count and granulate to lymphocyte ratios, indicating the sharks had healthy immune systems. This discovery suggests a previously unknown physiological defence against heavy metal poisoning. Great whites are known to have a propensity for "self-healing and avoiding age-related ailments".[80]
+A 2007 study from the University of New South Wales in Sydney, Australia, used CT scans of a shark's skull and computer models to measure the shark's maximum bite force. The study reveals the forces and behaviours its skull is adapted to handle and resolves competing theories about its feeding behaviour.[81] In 2008, a team of scientists led by Stephen Wroe conducted an experiment to determine the great white shark's jaw power and findings indicated that a specimen massing 3,324 kg (7,328 lb) could exert a bite force of 18,216 newtons (4,095 lbf).[45]
+This shark's behaviour and social structure is complex.[82] In South Africa, white sharks have a dominance hierarchy depending on the size, sex and squatter's rights: Females dominate males, larger sharks dominate smaller sharks, and residents dominate newcomers. When hunting, great whites tend to separate and resolve conflicts with rituals and displays. White sharks rarely resort to combat although some individuals have been found with bite marks that match those of other white sharks. This suggests that when a great white approaches too closely to another, they react with a warning bite. Another possibility is that white sharks bite to show their dominance.
+The great white shark is one of only a few sharks known to regularly lift its head above the sea surface to gaze at other objects such as prey. This is known as spy-hopping. This behaviour has also been seen in at least one group of blacktip reef sharks, but this might be learned from interaction with humans (it is theorized that the shark may also be able to smell better this way because smell travels through air faster than through water). White sharks are generally very curious animals, display intelligence and may also turn to socializing if the situation demands it. At Seal Island, white sharks have been observed arriving and departing in stable "clans" of two to six individuals on a yearly basis. Whether clan members are related is unknown, but they get along peacefully enough. In fact, the social structure of a clan is probably most aptly compared to that of a wolf pack; in that each member has a clearly established rank and each clan has an alpha leader. When members of different clans meet, they establish social rank nonviolently through any of a variety of interactions.[83]
+Great white sharks are carnivorous and prey upon fish (e.g. tuna, rays, other sharks),[83] cetaceans (i.e., dolphins, porpoises, whales), pinnipeds (e.g. seals, fur seals,[83] and sea lions), sea turtles,[83] sea otters (Enhydra lutris) and seabirds.[84] Great whites have also been known to eat objects that they are unable to digest. Juvenile white sharks predominantly prey on fish, including other elasmobranchs, as their jaws are not strong enough to withstand the forces required to attack larger prey such as pinnipeds and cetaceans until they reach a length of 3 m (9.8 ft) or more, at which point their jaw cartilage mineralizes enough to withstand the impact of biting into larger prey species.[85] Upon approaching a length of nearly 4 m (13 ft), great white sharks begin to target predominantly marine mammals for food, though individual sharks seem to specialize in different types of prey depending on their preferences.[86][87] They seem to be highly opportunistic.[88][89] These sharks prefer prey with a high content of energy-rich fat. Shark expert Peter Klimley used a rod-and-reel rig and trolled carcasses of a seal, a pig, and a sheep from his boat in the South Farallons. The sharks attacked all three baits but rejected the sheep carcass.[90]
+Off California, sharks immobilize northern elephant seals (Mirounga angustirostris) with a large bite to the hindquarters (which is the main source of the seal's mobility) and wait for the seal to bleed to death. This technique is especially used on adult male elephant seals, which are typically larger than the shark, ranging between 1,500 and 2,000 kg (3,300 and 4,400 lb), and are potentially dangerous adversaries.[91][92] Most commonly though, juvenile elephant seals are the most frequently eaten at elephant seal colonies.[93] Prey is normally attacked sub-surface. Harbor seals (Phoca vitulina) are taken from the surface and dragged down until they stop struggling. They are then eaten near the bottom. California sea lions (Zalophus californianus) are ambushed from below and struck mid-body before being dragged and eaten.[94]
+In the Northwest Atlantic mature great whites are known to feed on both harbor and grey seals.[32] Unlike adults, juvenile white sharks in the area feed on smaller fish species until they are large enough to prey on marine mammals such as seals.[95]
+White sharks also attack dolphins and porpoises from above, behind or below to avoid being detected by their echolocation. Targeted species include dusky dolphins (Lagenorhynchus obscurus),[48] Risso's dolphins (Grampus griseus),[48] bottlenose dolphins (Tursiops ssp.),[48][96] humpback dolphins (Sousa ssp.),[96] harbour porpoises (Phocoena phocoena),[48] and Dall's porpoises (Phocoenoides dalli).[48] Groups of dolphins have occasionally been observed defending themselves from sharks with mobbing behaviour.[96] White shark predation on other species of small cetacean has also been observed. In August 1989, a 1.8 m (5.9 ft) juvenile male pygmy sperm whale (Kogia breviceps) was found stranded in central California with a bite mark on its caudal peduncle from a great white shark.[97] In addition, white sharks attack and prey upon beaked whales.[48][96] Cases where an adult Stejneger's beaked whale (Mesoplodon stejnegeri), with a mean mass of around 1,100 kg (2,400 lb),[98] and a juvenile Cuvier's beaked whale (Ziphius cavirostris), an individual estimated at 3 m (9.8 ft), were hunted and killed by great white sharks have also been observed.[99] When hunting sea turtles, they appear to simply bite through the carapace around a flipper, immobilizing the turtle. The heaviest species of bony fish, the oceanic sunfish (Mola mola), has been found in great white shark stomachs.[88]
+Off Seal Island, False Bay in South Africa, the sharks ambush brown fur seals (Arctocephalus pusillus) from below at high speeds, hitting the seal mid-body. They can go so fast that they completely leave the water. The peak burst speed is estimated to be above 40 km/h (25 mph).[100] They have also been observed chasing prey after a missed attack. Prey is usually attacked at the surface.[101] Shark attacks most often occur in the morning, within 2 hours of sunrise, when visibility is poor. Their success rate is 55% in the first 2 hours, falling to 40% in late morning after which hunting stops.[83]
+Whale carcasses comprise an important part of the diet of white sharks. However, this has rarely been observed due to whales dying in remote areas. It has been estimated that 30 kg (66 lb) of whale blubber could feed a 4.5 m (15 ft) white shark for 1.5 months. Detailed observations were made of four whale carcasses in False Bay between 2000 and 2010. Sharks were drawn to the carcass by chemical and odour detection, spread by strong winds. After initially feeding on the whale caudal peduncle and fluke, the sharks would investigate the carcass by slowly swimming around it and mouthing several parts before selecting a blubber-rich area. During feeding bouts of 15–20 seconds the sharks removed flesh with lateral headshakes, without the protective ocular rotation they employ when attacking live prey. The sharks were frequently observed regurgitating chunks of blubber and immediately returning to feed, possibly in order to replace low energy yield pieces with high energy yield pieces, using their teeth as mechanoreceptors to distinguish them. After feeding for several hours, the sharks appeared to become lethargic, no longer swimming to the surface; they were observed mouthing the carcass but apparently unable to bite hard enough to remove flesh, they would instead bounce off and slowly sink. Up to eight sharks were observed feeding simultaneously, bumping into each other without showing any signs of aggression; on one occasion a shark accidentally bit the head of a neighbouring shark, leaving two teeth embedded, but both continued to feed unperturbed. Smaller individuals hovered around the carcass eating chunks that drifted away. Unusually for the area, large numbers of sharks over five metres long were observed, suggesting that the largest sharks change their behaviour to search for whales as they lose the manoeuvrability required to hunt seals. The investigating team concluded that the importance of whale carcasses, particularly for the largest white sharks, has been underestimated.[102] In another documented incident, white sharks were observed scavenging on a whale carcass alongside tiger sharks.[103] In 2020, Marine biologists Dines and Gennari et al., published a documented incident in the journal "Marine and Freshwater Research" of a group of great white sharks exhibiting pack-like behaviour, successfully attacking and killing a live adult humpback whale. The sharks utilized the classic attack strategy utilized on pinnipeds when attacking the whale, even utilizing the bite-and-spit tactic they employ on smaller prey items. The whale was an entangled individual, heavily emaciated and thus more vulnerable to the sharks' attacks. The incident is the first known documentation of great whites actively killing a large baleen whale.[104][105] A second incident regarding great white sharks killing humpback whales involving a single large female great white nicknamed Helen was documented off the coast of South Africa. Working alone, the shark attacked a 33 ft (10 m) emaciated and entangled humpback whale by attacking the whale's tail to cripple it before she managed to drown the whale by biting onto its head and pulling it underwater. The attack was witnessed via aerial drone by marine biologist Ryan Johnson, who said the attack went on for roughly 50 minutes before the shark successfully killed the whale. Johnson suggested that the shark may have strategized its attack in order to kill such a large animal.[106][107]
+Stomach contents of great whites also indicates that whale sharks both juvenile and adult may also be included on the animal's menu, though whether this is active hunting or scavenging is not known at present.[108][109]
+Great white sharks were previously thought to reach sexual maturity at around 15 years of age, but are now believed to take far longer; male great white sharks reach sexual maturity at age 26, while females take 33 years to reach sexual maturity.[9][110][111] Maximum life span was originally believed to be more than 30 years, but a study by the Woods Hole Oceanographic Institution placed it at upwards of 70 years. Examinations of vertebral growth ring count gave a maximum male age of 73 years and a maximum female age of 40 years for the specimens studied. The shark's late sexual maturity, low reproductive rate, long gestation period of 11 months and slow growth make it vulnerable to pressures such as overfishing and environmental change.[8]
+Little is known about the great white shark's mating habits, and mating behaviour has not yet been observed in this species. It is possible that whale carcasses are an important location for sexually mature sharks to meet for mating.[102] Birth has never been observed, but pregnant females have been examined. Great white sharks are ovoviviparous, which means eggs develop and hatch in the uterus and continue to develop until birth.[112] The great white has an 11-month gestation period. The shark pup's powerful jaws begin to develop in the first month. The unborn sharks participate in oophagy, in which they feed on ova produced by the mother. Delivery is in spring and summer.[113] The largest number of pups recorded for this species is 14 pups from a single mother measuring 4.5 m (15 ft) that was killed incidentally off Taiwan in 2019.[114] The Northern Pacific population of great whites is suspected to breed off the Sea of Cortez, as evidenced by local fisherman who have said to have caught them and evidenced by teeth found at dump sites for discarded parts from their catches.[citation needed]
+A breach is the result of a high speed approach to the surface with the resulting momentum taking the shark partially or completely clear of the water. This is a hunting technique employed by great white sharks whilst hunting seals. This technique is often used on cape fur seals at Seal Island in False Bay, South Africa. Because the behaviour is unpredictable, it is very hard to document. It was first photographed by Chris Fallows and Rob Lawrence who developed the technique of towing a slow-moving seal decoy to trick the sharks to breach.[115] Between April and September, scientists may observe around 600 breaches. The seals swim on the surface and the great white sharks launch their predatory attack from the deeper water below. They can reach speeds of up to 40 km/h (25 mph) and can at times launch themselves more than 3 m (10 ft) into the air. Just under half of observed breach attacks are successful.[116] In 2011, a 3-m-long shark jumped onto a seven-person research vessel off Seal Island in Mossel Bay. The crew were undertaking a population study using sardines as bait, and the incident was judged not to be an attack on the boat but an accident.[117]
+Interspecific competition between the great white shark and the orca is probable in regions where dietary preferences of both species may overlap.[96] An incident was documented on 4 October 1997, in the Farallon Islands off California in the United States. An estimated 4.7–5.3 m (15–17 ft) female orca immobilized an estimated 3–4 m (9.8–13.1 ft) great white shark.[118] The orca held the shark upside down to induce tonic immobility and kept the shark still for fifteen minutes, causing it to suffocate. The orca then proceeded to eat the dead shark's liver.[96][118][119] It is believed that the scent of the slain shark's carcass caused all the great whites in the region to flee, forfeiting an opportunity for a great seasonal feed.[120] Another similar attack apparently occurred there in 2000, but its outcome is not clear.[121] After both attacks, the local population of about 100 great whites vanished.[119][121] Following the 2000 incident, a great white with a satellite tag was found to have immediately submerged to a depth of 500 m (1,600 ft) and swum to Hawaii.[121] In 2015, a pod of orcas was recorded to have killed a great white shark off South Australia.[122] In 2017, three great whites were found washed ashore near Gaansbai, South Africa, with their body cavities torn open and the livers removed by what is likely to have been killer whales.[123] Killer whales also generally impact great white distribution. Studies published in 2019 of killer whale and great white shark distribution and interactions around the Farallon Islands indicate that the cetaceans impact the sharks negatively, with brief appearances by killer whales causing the sharks to seek out new feeding areas until the next season.[124] Occasionally, however, some great whites have been seen to swim near orcas without fear.[125]
+Of all shark species, the great white shark is responsible for by far the largest number of recorded shark bite incidents on humans, with 272 documented unprovoked bite incidents on humans as of 2012.[18]
+More than any documented bite incident, Peter Benchley's best-selling novel Jaws and the subsequent 1975 film adaptation directed by Steven Spielberg provided the great white shark with the image of being a "man-eater" in the public mind.[126] While great white sharks have killed humans in at least 74 documented unprovoked bite incidents, they typically do not target them: for example, in the Mediterranean Sea there have been 31 confirmed bite incidents against humans in the last two centuries, most of which were non-fatal. Many of the incidents seemed to be "test-bites". Great white sharks also test-bite buoys, flotsam, and other unfamiliar objects, and they might grab a human or a surfboard to identify what it is.
+Contrary to popular belief, great white sharks do not mistake humans for seals.[127] Many bite incidents occur in waters with low visibility or other situations which impair the shark's senses. The species appears to not like the taste of humans, or at least finds the taste unfamiliar. Further research shows that they can tell in one bite whether or not the object is worth predating upon. Humans, for the most part, are too bony for their liking. They much prefer seals, which are fat and rich in protein.[128]
+Humans are not appropriate prey because the shark's digestion is too slow to cope with a human's high ratio of bone to muscle and fat. Accordingly, in most recorded shark bite incidents, great whites broke off contact after the first bite. Fatalities are usually caused by blood loss from the initial bite rather than from critical organ loss or from whole consumption. From 1990 to 2011 there have been a total of 139 unprovoked great white shark bite incidents, 29 of which were fatal.[129]
+However, some researchers have hypothesized that the reason the proportion of fatalities is low is not because sharks do not like human flesh, but because humans are often able to escape after the first bite. In the 1980s, John McCosker, chair of aquatic biology at the California Academy of Sciences, noted that divers who dove solo and were bitten by great whites were generally at least partially consumed, while divers who followed the buddy system were generally rescued by their companion. McCosker and Timothy C. Tricas, an author and professor at the University of Hawaii, suggest that a standard pattern for great whites is to make an initial devastating attack and then wait for the prey to weaken before consuming the wounded animal. Humans' ability to move out of reach with the help of others, thus foiling the attack, is unusual for a great white's prey.[130]
+Shark culling is the deliberate killing of sharks by a government in an attempt to reduce shark attacks; shark culling is often called "shark control".[131] These programs have been criticized by environmentalists and scientists—they say these programs harm the marine ecosystem; they also say such programs are "outdated, cruel, and ineffective".[132] Many different species (dolphins, turtles, etc.) are also killed in these programs (because of their use of shark nets and drum lines)—15,135 marine animals were killed in New South Wales' nets between 1950 and 2008,[131] and 84,000 marine animals were killed by Queensland authorities from 1962 to 2015.[133]
+Great white sharks are currently killed in both Queensland and New South Wales in "shark control" (shark culling) programs.[131] Queensland uses shark nets and drum lines with baited hooks, while New South Wales only uses nets. From 1962 to 2018, Queensland authorities killed about 50,000 sharks, many of which were great whites.[134] From 2013 to 2014 alone, 667 sharks were killed by Queensland authorities, including great white sharks.[131] In Queensland, great white sharks found alive on the drum lines are shot.[135] In New South Wales, between 1950 and 2008, a total of 577 great white sharks were killed in nets.[131] Between September 2017 and April 2018, fourteen great white sharks were killed in New South Wales.[136]
+KwaZulu-Natal (an area of South Africa) also has a "shark control" program that kills great white sharks and other marine life. In a 30-year period, more than 33,000 sharks were killed in KwaZulu-Natal's shark-killing program, including great whites.[137]
+In 2014 the state government of Western Australia led by Premier Colin Barnett implemented a policy of killing large sharks. The policy, colloquially referred to as the Western Australian shark cull, was intended to protect users of the marine environment from shark bite incidents, following the deaths of seven people on the Western Australian coastline in the years 2010–2013.[138] Baited drum lines were deployed near popular beaches using hooks designed to catch great white sharks, as well as bull and tiger sharks. Large sharks found hooked but still alive were shot and their bodies discarded at sea.[139] The government claimed they were not culling the sharks, but were using a "targeted, localised, hazard mitigation strategy".[140] Barnett described opposition as "ludicrous" and "extreme", and said that nothing could change his mind.[141] This policy was met with widespread condemnation from the scientific community, which showed that species responsible for bite incidents were notoriously hard to identify, that the drum lines failed to capture white sharks, as intended, and that the government also failed to show any correlation between their drum line policy and a decrease in shark bite incidents in the region.[142]
+Great white sharks infrequently bite and sometimes even sink boats. Only five of the 108 authenticated unprovoked shark bite incidents reported from the Pacific Coast during the 20th century involved kayakers.[143] In a few cases they have bitten boats up to 10 m (33 ft) in length. They have bumped or knocked people overboard, usually biting the boat from the stern. In one case in 1936, a large shark leapt completely into the South African fishing boat Lucky Jim, knocking a crewman into the sea. Tricas and McCosker's underwater observations suggest that sharks are attracted to boats by the electrical fields they generate, which are picked up by the ampullae of Lorenzini and confuse the shark about whether or not wounded prey might be near-by.[144]
+Prior to August 1981, no great white shark in captivity lived longer than 11 days. In August 1981, a great white survived for 16 days at SeaWorld San Diego before being released.[145] The idea of containing a live great white at SeaWorld Orlando was used in the 1983 film Jaws 3-D.
+Monterey Bay Aquarium first attempted to display a great white in 1984, but the shark died after 11 days because it did not eat.[146] In July 2003, Monterey researchers captured a small female and kept it in a large netted pen near Malibu for five days. They had the rare success of getting the shark to feed in captivity before its release.[147] Not until September 2004 was the aquarium able to place a great white on long-term exhibit. A young female, which was caught off the coast of Ventura, was kept in the aquarium's 3.8 million l (1 million US gal) Outer Bay exhibit for 198 days before she was released in March 2005. She was tracked for 30 days after release.[148] On the evening of 31 August 2006, the aquarium introduced a juvenile male caught outside Santa Monica Bay.[149] His first meal as a captive was a large salmon steak on 8 September 2006, and as of that date, he was estimated to be 1.72 m (68 in) in length and to weigh approximately 47 kg (104 lb). He was released on 16 January 2007, after 137 days in captivity.
+Monterey Bay Aquarium housed a third great white, a juvenile male, for 162 days between 27 August 2007, and 5 February 2008. On arrival, he was 1.4 m (4.6 ft) long and weighed 30.6 kg (67 lb). He grew to 1.8 m (5.9 ft) and 64 kg (141 lb) before release. A juvenile female came to the Outer Bay Exhibit on 27 August 2008. While she did swim well, the shark fed only one time during her stay and was tagged and released on 7 September 2008. Another juvenile female was captured near Malibu on 12 August 2009, introduced to the Outer Bay exhibit on 26 August 2009, and was successfully released into the wild on 4 November 2009.[150] The Monterey Bay Aquarium introduced a 1.4-m-long male into their redesigned "Open Sea" exhibit on 31 August 2011. He was exhibited for 55 days, and was released into the wild on the 25th October the same year. However, the shark was determined to have died shortly after release via an attached electronic tag. The cause of death is not known.[151][152][153]
+The Monterey Bay Aquarium does not plan to exhibit any more great whites, as the main purpose of containing them was scientific. As data from captive great whites were no longer needed, the institute has instead shifted its focus to study wild sharks.[154]
+One of the largest adult great whites ever exhibited was at Japan's Okinawa Churaumi Aquarium in 2016, where a 3.5 m (11 ft) male was exhibited for three days before dying.[155][156] Probably the most famous captive was a 2.4 m (7.9 ft) female named Sandy, which in August 1980 became the only great white to be housed at the California Academy of Sciences' Steinhart Aquarium in San Francisco, California. She was released because she would not eat and constantly bumped against the walls.[157]
+Cage diving is most common at sites where great whites are frequent including the coast of South Africa, the Neptune Islands in South Australia,[158] and Guadalupe Island in Baja California. The popularity of cage diving and swimming with sharks is at the focus of a booming tourist industry.[159][160] A common practice is to chum the water with pieces of fish to attract the sharks. These practices may make sharks more accustomed to people in their environment and to associate human activity with food; a potentially dangerous situation. By drawing bait on a wire towards the cage, tour operators lure the shark to the cage, possibly striking it, exacerbating this problem. Other operators draw the bait away from the cage, causing the shark to swim past the divers.
+At present, hang baits are illegal off Isla Guadalupe and reputable dive operators do not use them. Operators in South Africa and Australia continue to use hang baits and pinniped decoys.[161] In South Australia, playing rock music recordings underwater, including the AC/DC album Back in Black has also been used experimentally to attract sharks.[162]
+Companies object to being blamed for shark bite incidents, pointing out that lightning tends to strike humans more often than sharks bite humans.[163] Their position is that further research needs to be done before banning practices such as chumming, which may alter natural behaviour.[164] One compromise is to only use chum in areas where whites actively patrol anyway, well away from human leisure areas. Also, responsible dive operators do not feed sharks. Only sharks that are willing to scavenge follow the chum trail and if they find no food at the end then the shark soon swims off and does not associate chum with a meal. It has been suggested that government licensing strategies may help enforce these responsible tourism.[161]
+The shark tourist industry has some financial leverage in conserving this animal. A single set of great white jaws can fetch up to £20,000. That is a fraction of the tourism value of a live shark; tourism is a more sustainable economic activity than shark fishing. For example, the dive industry in Gansbaai, South Africa consists of six boat operators with each boat guiding 30 people each day. With fees between £50 and £150 per person, a single live shark that visits each boat can create anywhere between £9,000 and £27,000 of revenue daily.[citation needed]
+Putting chum in the water
+A great white shark approaches divers in a cage off Dyer Island, Western Cape, South Africa
+A great white shark approaches a cage
+Tourists in a cage near Gansbaai
+It is unclear how much of a concurrent increase in fishing for great white sharks has caused the decline of great white shark populations from the 1970s to the present. No accurate global population numbers are available, but the great white shark is now considered vulnerable.[2] Sharks taken during the long interval between birth and sexual maturity never reproduce, making population recovery and growth difficult.
+The IUCN notes that very little is known about the actual status of the great white shark, but as it appears uncommon compared to other widely distributed species, it is considered vulnerable.[2] It is included in Appendix II of CITES,[15] meaning that international trade in the species requires a permit.[165] As of March 2010, it has also been included in Annex I of the CMS Migratory Sharks MoU, which strives for increased international understanding and coordination for the protection of certain migratory sharks.[166] A February 2010 study by Barbara Block of Stanford University estimated the world population of great white sharks to be lower than 3,500 individuals, making the species more vulnerable to extinction than the tiger, whose population is in the same range.[167] According to another study from 2014 by George H. Burgess, Florida Museum of Natural History, University of Florida, there are about 2,000 great white sharks near the California coast, which is 10 times higher than the previous estimate of 219 by Barbara Block.[168][169]
+Fishermen target many sharks for their jaws, teeth, and fins, and as game fish in general. The great white shark, however, is rarely an object of commercial fishing, although its flesh is considered valuable. If casually captured (it happens for example in some tonnare in the Mediterranean), it is misleadingly sold as smooth-hound shark.[170]
+The great white shark was declared vulnerable by the Australian Government in 1999 because of significant population decline and is currently protected under the Environmental Protection and Biodiversity Conservation (EPBC) Act.[171] The causes of decline prior to protection included mortality from sport fishing harvests as well as being caught in beach protection netting.[172]
+The national conservation status of the great white shark is reflected by all Australian states under their respective laws, granting the species full protection throughout Australia regardless of jurisdiction.[171] Many states had prohibited the killing or possession of great white sharks prior to national legislation coming into effect. The great white shark is further listed as threatened in Victoria under the Flora and Fauna Guarantee Act, and as rare or likely to become extinct under Schedule 5 of the Wildlife Conservation Act in Western Australia.[171]
+In 2002, the Australian government created the White Shark Recovery Plan, implementing government-mandated conservation research and monitoring for conservation in addition to federal protection and stronger regulation of shark-related trade and tourism activities.[172] An updated recovery plan was published in 2013 to review progress, research findings, and to implement further conservation actions.[16] A study in 2012 revealed that Australia's white shark population was separated by Bass Strait into genetically distinct eastern and western populations, indicating a need for the development of regional conservation strategies.[173]
+Presently, human-caused shark mortality is continuing, primarily from accidental and illegal catching in commercial and recreational fishing as well as from being caught in beach protection netting, and the populations of great white shark in Australia are yet to recover.[16]
+In spite of official protections in Australia, great white sharks continue to be killed in state "shark control" programs within Australia. For example, the government of Queensland has a "shark control" program (shark culling) which kills great white sharks (as well as other marine life) using shark nets and drum lines with baited hooks.[174][131] In Queensland, great white sharks that are found alive on the baited hooks are shot.[135] The government of New South Wales also kills great white sharks in its "shark control" program.[131] Partly because of these programs, shark numbers in eastern Australia have decreased.[134]
+The Australasian population of great white sharks is believed to be in excess of 8,000–10,000 individuals according to genetic research studies done by CSIRO, with an adult population estimated to be around 2,210 individuals in both Eastern and Western Australia. The annual survival rate for juveniles in these two separate populations was estimated in the same study to be close to 73 percent, while adult sharks had a 93 percent annual survival rate. Whether or not mortality rates in great white sharks have declined, or the population has increased as a result of the protection of this species in Australian waters is as yet unknown due to the slow growth rates of this species.[175]
+As of April 2007, great white sharks were fully protected within 370 km (230 mi) of New Zealand and additionally from fishing by New Zealand-flagged boats outside this range. The maximum penalty is a $250,000 fine and up to six months in prison.[176] In June 2018 the New Zealand Department of Conservation classified the great white shark under the New Zealand Threat Classification System as "Nationally Endangered". The species meets the criteria for this classification as there exists a moderate, stable population of between 1000 and 5000 mature individuals. This classification has the qualifiers "Data Poor" and "Threatened Overseas".[177]
+In 2013, great white sharks were added to California's Endangered Species Act. From data collected, the population of great whites in the North Pacific was estimated to be fewer than 340 individuals. Research also reveals these sharks are genetically distinct from other members of their species elsewhere in Africa, Australia, and the east coast of North America, having been isolated from other populations.[178]
+A 2014 study estimated the population of great white sharks along the California coastline to be approximately 2,400.[179][180]
+In 2015 Massachusetts banned catching, cage diving, feeding, towing decoys, or baiting and chumming for its significant and highly predictable migratory great white population without an appropriate research permit. The goal of these restrictions is to both protect the sharks and public health.[181]

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+Carcharhiniformes
+Heterodontiformes
+Hexanchiformes
+Lamniformes
+Orectolobiformes
+Pristiophoriformes
+Squaliformes
+Squatiniformes
+† Cladoselachiformes
+† Hybodontiformes
+† Symmoriida
+† Xenacanthida (Xenacantiformes)
+† Elegestolepis
+Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorpha (or Selachii) and are the sister group to the rays. However, the term "shark" has also been used for extinct members of the subclass Elasmobranchii outside the Selachimorpha, such as Cladoselache and Xenacanthus, as well as other Chondrichthyes such as the holocephalid eugenedontidans.
+Under this broader definition, the earliest known sharks date back to more than 420 million years ago.[2] Acanthodians are often referred to as "spiny sharks"; though they are not part of Chondrichthyes proper, they are a paraphyletic assemblage leading to cartilaginous fish as a whole.  Since then, sharks have diversified into over 500 species. They range in size from the small dwarf lanternshark (Etmopterus perryi), a deep sea species of only 17 centimetres (6.7 in) in length, to the whale shark (Rhincodon typus), the largest fish in the world, which reaches approximately 12 metres (40 ft) in length.[3] Sharks are found in all seas and are common to depths of 2,000 metres (6,600 ft). They generally do not live in freshwater although there are a few known exceptions, such as the bull shark and the river shark, which can be found in both seawater and freshwater.[4] Sharks have a covering of dermal denticles that protects their skin from damage and parasites in addition to improving their fluid dynamics. They have numerous sets of replaceable teeth.[5]
+Well-known species such as the tiger shark, blue shark, great white shark, mako shark, thresher shark, and hammerhead shark are apex predators—organisms at the top of their underwater food chain. Many shark populations are threatened by human activities.
+Until the 16th century,[6] sharks were known to mariners as "sea dogs".[7] This is still evidential in several species termed "dogfish," or the porbeagle.
+The etymology of the word "shark" is uncertain, the most likely etymology states that the original sense of the word was that of "predator, one who preys on others" from the Dutch schurk, meaning "villain, scoundrel" (cf. card shark, loan shark, etc.), which was later applied to the fish due to its predatory behaviour.[8]
+A now disproven theory is that it derives from the Yucatec Maya word xok (pronounced 'shok'), meaning "fish".[9]
+Evidence for this etymology came from the Oxford English Dictionary, which notes shark first came into use after Sir John Hawkins' sailors exhibited one in London in 1569 and posted "sharke" to refer to the large sharks of the Caribbean Sea. However, the Middle English Dictionary records an isolated occurrence of the word shark (referring to a sea fish) in a letter written by Thomas Beckington in 1442, which rules out a New World etymology.[10]
+Evidence for the existence of sharks dates from the Ordovician period, 450–420 million years ago, before land vertebrates existed and before a variety of plants had colonized the continents.[2] Only scales have been recovered from the first sharks and not all paleontologists agree that these are from true sharks, suspecting that these scales are actually those of thelodont agnathans.[11] The oldest generally accepted shark scales are from about 420 million years ago, in the Silurian period.[11] The first sharks looked very different from modern sharks.[12] At this time the most common shark tooth is the cladodont, a style of thin tooth with three tines like a trident, apparently to help catch fish. The majority of modern sharks can be traced back to around 100 million years ago.[13] Most fossils are of teeth, often in large numbers. Partial skeletons and even complete fossilized remains have been discovered. Estimates suggest that sharks grow tens of thousands of teeth over a lifetime, which explains the abundant fossils. The teeth consist of easily fossilized calcium phosphate, an apatite. When a shark dies, the decomposing skeleton breaks up, scattering the apatite prisms. Preservation requires rapid burial in bottom sediments.
+Among the most ancient and primitive sharks is Cladoselache, from about 370 million years ago,[12] which has been found within Paleozoic strata in Ohio, Kentucky, and Tennessee. At that point in Earth's history these rocks made up the soft bottom sediments of a large, shallow ocean, which stretched across much of North America. Cladoselache was only about 1 metre (3.3 ft) long with stiff triangular fins and slender jaws.[12] Its teeth had several pointed cusps, which wore down from use. From the small number of teeth found together, it is most likely that Cladoselache did not replace its teeth as regularly as modern sharks. Its caudal fins had a similar shape to the great white sharks and the pelagic shortfin and longfin makos. The presence of whole fish arranged tail-first in their stomachs suggest that they were fast swimmers with great agility.
+Most fossil sharks from about 300 to 150 million years ago can be assigned to one of two groups. The Xenacanthida was almost exclusive to freshwater environments.[14][15] By the time this group became extinct about 220 million years ago, they had spread worldwide. The other group, the hybodonts, appeared about 320 million years ago and lived mostly in the oceans, but also in freshwater.[citation needed] The results of a 2014 study of the gill structure of an unusually well preserved 325-million-year-old fossil suggested that sharks are not "living fossils", but rather have evolved more extensively than previously thought over the hundreds of millions of years they have been around.[16]
+Modern sharks began to appear about 100 million years ago.[13] Fossil mackerel shark teeth date to the Early Cretaceous. One of the most recently evolved families is the hammerhead shark (family Sphyrnidae), which emerged in the Eocene.[17] The oldest white shark teeth date from 60 to 66 million years ago, around the time of the extinction of the dinosaurs. In early white shark evolution there are at least two lineages: one lineage is of white sharks with coarsely serrated teeth and it probably gave rise to the modern great white shark, and another lineage is of white sharks with finely serrated teeth. These sharks attained gigantic proportions and include the extinct megatoothed shark, C. megalodon. Like most extinct sharks, C. megalodon is also primarily known from its fossil teeth and vertebrae. This giant shark reached a total length (TL) of more than 16 metres (52 ft).[18][19] C. megalodon may have approached a maxima of 20.3 metres (67 ft) in total length and 103 metric tons (114 short tons) in mass.[20] Paleontological evidence suggests that this shark was an active predator of large cetaceans.[20]
+Sharks belong to the superorder Selachimorpha in the subclass Elasmobranchii in the class Chondrichthyes. The Elasmobranchii also include rays and skates; the Chondrichthyes also include Chimaeras. It was thought that the sharks form a polyphyletic group: some sharks are more closely related to rays than they are to some other sharks,[21] but current molecular studies support monophyly of both groups of sharks and batoids.[22][23]
+The superorder Selachimorpha is divided into Galea (or Galeomorphii), and Squalea (or Squalomorphii). The Galeans are the Heterodontiformes, Orectolobiformes, Lamniformes, and Carcharhiniformes. Lamnoids and Carcharhinoids are usually placed in one clade, but recent studies show the Lamnoids and Orectoloboids are a clade. Some scientists now think that Heterodontoids may be Squalean. The Squaleans are divided into Hexanchiformes and Squalomorpha. The former includes cow shark and frilled shark, though some authors propose both families to be moved to separate orders. The Squalomorpha contains the Squaliformes and the Hypnosqualea. The Hypnosqualea may be invalid. It includes the Squatiniformes, and the Pristorajea, which may also be invalid, but includes the Pristiophoriformes and the Batoidea.[21][24]
+There are more than 470 species of sharks split across twelve orders, including four orders of sharks that have gone extinct:[24]
+Shark teeth are embedded in the gums rather than directly affixed to the jaw, and are constantly replaced throughout life. Multiple rows of replacement teeth grow in a groove on the inside of the jaw and steadily move forward in comparison to a conveyor belt; some sharks lose 30,000 or more teeth in their lifetime. The rate of tooth replacement varies from once every 8 to 10 days to several months. In most species, teeth are replaced one at a time as opposed to the simultaneous replacement of an entire row, which is observed in the cookiecutter shark.[25]
+Tooth shape depends on the shark's diet: those that feed on mollusks and crustaceans have dense and flattened teeth used for crushing, those that feed on fish have needle-like teeth for gripping, and those that feed on larger prey such as mammals have pointed lower teeth for gripping and triangular upper teeth with serrated edges for cutting. The teeth of plankton-feeders such as the basking shark are small and non-functional.[26]
+Shark skeletons are very different from those of bony fish and terrestrial vertebrates. Sharks and other cartilaginous fish (skates and rays) have skeletons made of cartilage and connective tissue. Cartilage is flexible and durable, yet is about half the normal density of bone. This reduces the skeleton's weight, saving energy.[27] Because sharks do not have rib cages, they can easily be crushed under their own weight on land.[28]
+The jaws of sharks, like those of rays and skates, are not attached to the cranium. The jaw's surface (in comparison to the shark's vertebrae and gill arches) needs extra support due to its heavy exposure to physical stress and its need for strength. It has a layer of tiny hexagonal plates called "tesserae", which are crystal blocks of calcium salts arranged as a mosaic.[29] This gives these areas much of the same strength found in the bony tissue found in other animals.
+Generally sharks have only one layer of tesserae, but the jaws of large specimens, such as the bull shark, tiger shark, and the great white shark, have two to three layers or more, depending on body size. The jaws of a large great white shark may have up to five layers.[27] In the rostrum (snout), the cartilage can be spongy and flexible to absorb the power of impacts.
+Fin skeletons are elongated and supported with soft and unsegmented rays named ceratotrichia, filaments of elastic protein resembling the horny keratin in hair and feathers.[30] Most sharks have eight fins. Sharks can only drift away from objects directly in front of them because their fins do not allow them to move in the tail-first direction.[28]
+Unlike bony fish, sharks have a complex dermal corset made of flexible collagenous fibers and arranged as a helical network surrounding their body. This works as an outer skeleton, providing attachment for their swimming muscles and thus saving energy.[31] Their dermal teeth give them hydrodynamic advantages as they reduce turbulence when swimming.[32]
+Tails provide thrust, making speed and acceleration dependent on tail shape. Caudal fin shapes vary considerably between shark species, due to their evolution in separate environments. Sharks possess a heterocercal caudal fin in which the dorsal portion is usually noticeably larger than the ventral portion. This is because the shark's vertebral column extends into that dorsal portion, providing a greater surface area for muscle attachment. This allows more efficient locomotion among these negatively buoyant cartilaginous fish. By contrast, most bony fish possess a homocercal caudal fin.[33]
+Tiger sharks have a large upper lobe, which allows for slow cruising and sudden bursts of speed. The tiger shark must be able to twist and turn in the water easily when hunting to support its varied diet, whereas the porbeagle shark, which hunts schooling fish such as mackerel and herring, has a large lower lobe to help it keep pace with its fast-swimming prey.[34] Other tail adaptations help sharks catch prey more directly, such as the thresher shark's usage of its powerful, elongated upper lobe to stun fish and squid.
+Unlike bony fish, sharks do not have gas-filled swim bladders for buoyancy. Instead, sharks rely on a large liver filled with oil that contains squalene, and their cartilage, which is about half the normal density of bone.[31] Their liver constitutes up to 30% of their total body mass.[35] The liver's effectiveness is limited, so sharks employ dynamic lift to maintain depth while swimming. Sand tiger sharks store air in their stomachs, using it as a form of swim bladder.  Bottom-dwelling sharks, like the nurse shark, have negative buoyancy, allowing them to rest on the ocean floor.
+Some sharks, if inverted or stroked on the nose, enter a natural state of tonic immobility. Researchers use this condition to handle sharks safely.[36]
+Like other fish, sharks extract oxygen from seawater as it passes over their gills. Unlike other fish, shark gill slits are not covered, but lie in a row behind the head. A modified slit called a spiracle lies just behind the eye, which assists the shark with taking in water during respiration and plays a major role in bottom–dwelling sharks. Spiracles are reduced or missing in active pelagic sharks.[26] While the shark is moving, water passes through the mouth and over the gills in a process known as "ram ventilation". While at rest, most sharks pump water over their gills to ensure a constant supply of oxygenated water. A small number of species have lost the ability to pump water through their gills and must swim without rest. These species are obligate ram ventilators and would presumably asphyxiate if unable to move. Obligate ram ventilation is also true of some pelagic bony fish species.[37][38]
+The respiration and circulation process begins when deoxygenated blood travels to the shark's two-chambered heart. Here the shark pumps blood to its gills via the ventral aorta artery where it branches into afferent brachial arteries. Reoxygenation takes place in the gills and the reoxygenated blood flows into the efferent brachial arteries, which come together to form the dorsal aorta. The blood flows from the dorsal aorta throughout the body. The deoxygenated blood from the body then flows through the posterior cardinal veins and enters the posterior cardinal sinuses. From there blood enters the heart ventricle and the cycle repeats.[39]
+Most sharks are "cold-blooded" or, more precisely, poikilothermic, meaning that their internal body temperature matches that of their ambient environment. Members of the family Lamnidae (such as the shortfin mako shark and the great white shark) are homeothermic and maintain a higher body temperature than the surrounding water. In these sharks, a strip of aerobic red muscle located near the center of the body generates the heat, which the body retains via a countercurrent exchange mechanism by a system of blood vessels called the rete mirabile ("miraculous net"). The common thresher and bigeye thresher sharks have a similar mechanism for maintaining an elevated body temperature.[40]
+In contrast to bony fish, with the exception of the coelacanth,[41] the blood and other tissue of sharks and Chondrichthyes is generally isotonic to their marine environments because of the high concentration of urea (up to 2.5%[42]) and trimethylamine N-oxide (TMAO), allowing them to be in osmotic balance with the seawater. This adaptation prevents most sharks from surviving in freshwater, and they are therefore confined to marine environments. A few exceptions exist, such as the bull shark, which has developed a way to change its kidney function to excrete large amounts of urea.[35] When a shark dies, the urea is broken down to ammonia by bacteria, causing the dead body to gradually smell strongly of ammonia.[43][44]
+Research in 1930 by Homer W. Smith showed that sharks' urine doesn't contain sufficient sodium to avoid hypernatremia, and it was postulated that there must be an additional mechanism for salt secretion. In 1960 it was discovered at the Mount Desert Island Biological Laboratory in Salsbury Cove, Maine that sharks have a type of salt gland located at the end of the intestine, known as the "rectal gland", whose function is the secretion of chlorides.[45]
+Digestion can take a long time. The food moves from the mouth to a J-shaped stomach, where it is stored and initial digestion occurs.[46] Unwanted items may never get past the stomach, and instead the shark either vomits or turns its stomachs inside out and ejects unwanted items from its mouth.[47]
+One of the biggest differences between the digestive systems of sharks and mammals is that sharks have much shorter intestines. This short length is achieved by the spiral valve with multiple turns within a single short section instead of a long tube-like intestine. The valve provides a long surface area, requiring food to circulate inside the short gut until fully digested, when remaining waste products pass into the cloaca.[46]
+Biofluorescence is a characteristic of a few shark species, such as the swell shark and the chain catshark, the mechanism of which is unique among marine animals and depends upon a small molecule metabolite.[48]
+Sharks have keen olfactory senses, located in the short duct (which is not fused, unlike bony fish) between the anterior and posterior nasal openings, with some species able to detect as little as one part per million of blood in seawater.[49]
+Sharks have the ability to determine the direction of a given scent based on the timing of scent detection in each nostril.[50] This is similar to the method mammals use to determine direction of sound.
+They are more attracted to the chemicals found in the intestines of many species, and as a result often linger near or in sewage outfalls. Some species, such as nurse sharks, have external barbels that greatly increase their ability to sense prey.
+Shark eyes are similar to the eyes of other vertebrates, including similar lenses, corneas and retinas, though their eyesight is well adapted to the marine environment with the help of a tissue called tapetum lucidum. This tissue is behind the retina and reflects light back to it, thereby increasing visibility in the dark waters. The effectiveness of the tissue varies, with some sharks having stronger nocturnal adaptations. Many sharks can contract and dilate their pupils, like humans, something no teleost fish can do. Sharks have eyelids, but they do not blink because the surrounding water cleans their eyes. To protect their eyes some species have nictitating membranes. This membrane covers the eyes while hunting and when the shark is being attacked. However, some species, including the great white shark (Carcharodon carcharias), do not have this membrane, but instead roll their eyes backwards to protect them when striking prey. The importance of sight in shark hunting behavior is debated. Some believe that electro- and chemoreception are more significant, while others point to the nictating membrane as evidence that sight is important. Presumably, the shark would not protect its eyes were they unimportant. The use of sight probably varies with species and water conditions. The shark's field of vision can swap between monocular and stereoscopic at any time.[51] A micro-spectrophotometry study of 17 species of shark found 10 had only rod photoreceptors and no cone cells in their retinas giving them good night vision while making them colorblind. The remaining seven species had in addition to rods a single type of cone photoreceptor sensitive to green and, seeing only in shades of grey and green, are believed to be effectively colorblind. The study indicates that an object's contrast against the background, rather than colour, may be more important for object detection.[52]
+[53][54]
+Although it is hard to test the hearing of sharks, they may have a sharp sense of hearing and can possibly hear prey from many miles away.[55] A small opening on each side of their heads (not the spiracle) leads directly into the inner ear through a thin channel. The lateral line shows a similar arrangement, and is open to the environment via a series of openings called lateral line pores. This is a reminder of the common origin of these two vibration- and sound-detecting organs that are grouped together as the acoustico-lateralis system. In bony fish and tetrapods the external opening into the inner ear has been lost.
+The ampullae of Lorenzini are the electroreceptor organs. They number in the hundreds to thousands. Sharks use the ampullae of Lorenzini to detect the electromagnetic fields that all living things produce.[56] This helps sharks (particularly the hammerhead shark) find prey. The shark has the greatest electrical sensitivity of any animal. Sharks find prey hidden in sand by detecting the electric fields they produce. Ocean currents moving in the magnetic field of the Earth also generate electric fields that sharks can use for orientation and possibly navigation.[57]
+This system is found in most fish, including sharks. It is a tactile sensory system which allows the organism to detect water speed and pressure changes near by.[58] The main component of the system is the neuromast, a cell similar to hair cells present in the vertebrate ear that interact with the surrounding aquatic environment. This helps sharks distinguish between the currents around them, obstacles off on their periphery, and struggling prey out of visual view. The shark can sense frequencies in the range of 25 to 50 Hz.[59]
+Shark lifespans vary by species. Most live 20 to 30 years. The spiny dogfish has one of the longest lifespans at more than 100 years.[60] Whale sharks (Rhincodon typus) may also live over 100 years.[61] Earlier estimates suggested the Greenland shark (Somniosus microcephalus) could reach about 200 years, but a recent study found that a 5.02-metre-long (16.5 ft) specimen was 392 ± 120 years old (i.e., at least 272 years old), making it the longest-lived vertebrate known.[62][63]
+Unlike most bony fish, sharks are K-selected reproducers, meaning that they produce a small number of well-developed young as opposed to a large number of poorly developed young. Fecundity in sharks ranges from 2 to over 100 young per reproductive cycle.[64] Sharks mature slowly relative to many other fish. For example, lemon sharks reach sexual maturity at around age 13–15.[65]
+Sharks practice internal fertilization.[66] The posterior part of a male shark's pelvic fins are modified into a pair of intromittent organs called claspers, analogous to a mammalian penis, of which one is used to deliver sperm into the female.[67]
+Mating has rarely been observed in sharks.[68] The smaller catsharks often mate with the male curling around the female. In less flexible species the two sharks swim parallel to each other while the male inserts a clasper into the female's oviduct. Females in many of the larger species have bite marks that appear to be a result of a male grasping them to maintain position during mating. The bite marks may also come from courtship behavior: the male may bite the female to show his interest. In some species, females have evolved thicker skin to withstand these bites.[67]
+There have been a number of documented cases in which a female shark who has not been in contact with a male has conceived a pup on her own through parthenogenesis.[69][70] The details of this process are not well understood, but genetic fingerprinting showed that the pups had no paternal genetic contribution, ruling out sperm storage. The extent of this behavior in the wild is unknown. Mammals are now the only major vertebrate group in which asexual reproduction has not been observed.
+Scientists say that asexual reproduction in the wild is rare, and probably a last-ditch effort to reproduce when a mate is not present. Asexual reproduction diminishes genetic diversity, which helps build defenses against threats to the species. Species that rely solely on it risk extinction. Asexual reproduction may have contributed to the blue shark's decline off the Irish coast.[71]
+Sharks display three ways to bear their young, varying by species, oviparity, viviparity and ovoviviparity.[72][73]
+Most sharks are ovoviviparous, meaning that the eggs hatch in the oviduct within the mother's body and that the egg's yolk and fluids secreted by glands in the walls of the oviduct nourishes the embryos. The young continue to be nourished by the remnants of the yolk and the oviduct's fluids. As in viviparity, the young are born alive and fully functional. Lamniforme sharks practice oophagy, where the first embryos to hatch eat the remaining eggs.  Taking this a step further,  sand tiger shark pups cannibalistically consume neighboring embryos.  The survival strategy for ovoviviparous species is to brood the young to a comparatively large size before birth. The whale shark is now classified as ovoviviparous rather than oviparous, because extrauterine eggs are now thought to have been aborted. Most ovoviviparous sharks give birth in sheltered areas, including bays, river mouths and shallow reefs. They choose such areas for protection from predators (mainly other sharks) and the abundance of food. Dogfish have the longest known gestation period of any shark, at 18 to 24 months. Basking sharks and frilled sharks appear to have even longer gestation periods, but accurate data are lacking.[72]
+Some species are oviparous, laying their fertilized eggs in the water. In most oviparous shark species, an egg case with the consistency of leather protects the developing embryo(s). These cases may be corkscrewed into crevices for protection. The egg case is commonly called a mermaid's purse. Oviparous sharks include the horn shark, catshark, Port Jackson shark, and swellshark.[72][74]
+Viviparity is the gestation of young without the use of a traditional egg, and results in live birth.[75] Viviparity in sharks can be placental or aplacental.[75] Young are born fully formed and self-sufficient.[75] Hammerheads, the requiem sharks (such as the bull and blue sharks), and smoothhounds are viviparous.[64][72]
+The classic view describes a solitary hunter, ranging the oceans in search of food. However, this applies to only a few species. Most live far more social, sedentary, benthic lives, and appear likely to have their own distinct personalities.[76] Even solitary sharks meet for breeding or at rich hunting grounds, which may lead them to cover thousands of miles in a year.[77] Shark migration patterns may be even more complex than in birds, with many sharks covering entire ocean basins.
+Sharks can be highly social, remaining in large schools. Sometimes more than 100 scalloped hammerheads congregate around seamounts and islands, e.g., in the Gulf of California.[35] Cross-species social hierarchies exist. For example, oceanic whitetip sharks dominate silky sharks of comparable size during feeding.[64]
+When approached too closely some sharks perform a threat display. This usually consists of exaggerated swimming movements, and can vary in intensity according to the threat level.[78]
+In general, sharks swim ("cruise") at an average speed of 8 kilometres per hour (5.0 mph), but when feeding or attacking, the average shark can reach speeds upwards of 19 kilometres per hour (12 mph). The shortfin mako shark, the fastest shark and one of the fastest fish, can burst at speeds up to 50 kilometres per hour (31 mph).[79] The great white shark is also capable of speed bursts. These exceptions may be due to the warm-blooded, or homeothermic, nature of these sharks' physiology. Sharks can travel 70 to 80 km in a day.[80]
+Sharks possess brain-to-body mass ratios that are similar to mammals and birds,[81] and have exhibited apparent curiosity and behavior resembling play in the wild.[82][83]
+There is evidence that juvenile lemon sharks can use observational learning in their investigation of novel objects in their environment.[84]
+All sharks need to keep water flowing over their gills in order for them to breathe; however, not all species need to be moving to do this. Those that are able to breathe while not swimming do so by using their spiracles to force water over their gills, thereby allowing them to extract oxygen from the water. It has been recorded that their eyes remain open while in this state and actively follow the movements of divers swimming around them[85] and as such they are not truly asleep.
+Species that do need to swim continuously to breathe go through a process known as sleep swimming, in which the shark is essentially unconscious. It is known from experiments conducted on the spiny dogfish that its spinal cord, rather than its brain, coordinates swimming, so spiny dogfish can continue to swim while sleeping, and this also may be the case in larger shark species.[85] In 2016 a great white shark was captured on video for the first time in a state researchers believed was sleep swimming.[86]
+Most sharks are carnivorous.[87] Basking sharks, whale sharks, and megamouth sharks have independently evolved different strategies for filter feeding plankton: basking sharks practice ram feeding, whale sharks use suction to take in plankton and small fishes, and megamouth sharks make suction feeding more efficient by using the luminescent tissue inside of their mouths to attract prey in the deep ocean. This type of feeding requires gill rakers—long, slender filaments that form a very efficient sieve—analogous to the baleen plates of the great whales. The shark traps the plankton in these filaments and swallows from time to time in huge mouthfuls. Teeth in these species are comparatively small because they are not needed for feeding.[87]
+Other highly specialized feeders include cookiecutter sharks, which feed on flesh sliced out of other larger fish and marine mammals. Cookiecutter teeth are enormous compared to the animal's size. The lower teeth are particularly sharp. Although they have never been observed feeding, they are believed to latch onto their prey and use their thick lips to make a seal, twisting their bodies to rip off flesh.[35]
+Some seabed–dwelling species are highly effective ambush predators. Angel sharks and wobbegongs use camouflage to lie in wait and suck prey into their mouths.[88] Many benthic sharks feed solely on crustaceans which they crush with their flat molariform teeth.
+Other sharks feed on squid or fish, which they swallow whole. The viper dogfish has teeth it can point outwards to strike and capture prey that it then swallows intact. The great white and other large predators either swallow small prey whole or take huge bites out of large animals. Thresher sharks use their long tails to stun shoaling fishes, and sawsharks either stir prey from the seabed or slash at swimming prey with their tooth-studded rostra.
+Many sharks, including the whitetip reef shark are cooperative feeders and hunt in packs to herd and capture elusive prey. These social sharks are often migratory, traveling huge distances around ocean basins in large schools. These migrations may be partly necessary to find new food sources.[89]
+Sharks are found in all seas. They generally do not live in fresh water, with a few exceptions such as the bull shark and the river shark which can swim both in seawater and freshwater.[90] Sharks are common down to depths of 2,000 metres (7,000 ft), and some live even deeper, but they are almost entirely absent below 3,000 metres (10,000 ft). The deepest confirmed report of a shark is a Portuguese dogfish at 3,700 metres (12,100 ft).[91]
+In 2006 the International Shark Attack File (ISAF) undertook an investigation into 96 alleged shark attacks, confirming 62 of them as unprovoked attacks and 16 as provoked attacks. The average number of fatalities worldwide per year between 2001 and 2006 from unprovoked shark attacks is 4.3.[92]
+Contrary to popular belief, only a few sharks are dangerous to humans. Out of more than 470 species, only four have been involved in a significant number of fatal, unprovoked attacks on humans: the great white, oceanic whitetip, tiger, and bull sharks.[93][94] These sharks are large, powerful predators, and may sometimes attack and kill people. Despite being responsible for attacks on humans they have all been filmed without using a protective cage.[95]
+The perception of sharks as dangerous animals has been popularized by publicity given to a few isolated unprovoked attacks, such as the Jersey Shore shark attacks of 1916, and through popular fictional works about shark attacks, such as the Jaws film series. Jaws author Peter Benchley, as well as Jaws director Steven Spielberg, later attempted to dispel the image of sharks as man-eating monsters.[96]
+To help avoid an unprovoked attack, humans should not wear jewelry or metal that is shiny and refrain from splashing around too much.[97]
+In general, sharks show little pattern of attacking humans specifically. Research indicates that when humans do become the object of a shark attack, it is possible that the shark has mistaken the human for species that are its normal prey, such as seals.[98][99]
+Until recently, only a few benthic species of shark, such as hornsharks, leopard sharks and catsharks, had survived in aquarium conditions for a year or more. This gave rise to the belief that sharks, as well as being difficult to capture and transport, were difficult to care for. More knowledge has led to more species (including the large pelagic sharks) living far longer in captivity, along with safer transportation techniques that have enabled long distance transportation.[100] The great white shark had never been successfully held in captivity for long periods of time until September 2004, when the Monterey Bay Aquarium successfully kept a young female for 198 days before releasing her.
+Most species are not suitable for home aquaria, and not every species sold by pet stores are appropriate. Some species can flourish in home saltwater aquaria.[101] Uninformed or unscrupulous dealers sometimes sell juvenile sharks like the nurse shark, which upon reaching adulthood is far too large for typical home aquaria.[101] Public aquaria generally do not accept donated specimens that have outgrown their housing. Some owners have been tempted to release them.[101] Species appropriate to home aquaria represent considerable spatial and financial investments as they generally approach adult lengths of 3 feet (90 cm) and can live up to 25 years.[101]
+Sharks figure prominently in Hawaiian mythology. Stories tell of men with shark jaws on their back who could change between shark and human form. A common theme was that a shark-man would warn beach-goers of sharks in the waters. The beach-goers would laugh and ignore the warnings and get eaten by the shark-man who warned them. Hawaiian mythology also includes many shark gods. Among a fishing people, the most popular of all aumakua, or deified ancestor guardians, are shark aumakua. Kamaku describes in detail how to offer a corpse to become a shark. The body transforms gradually until the kahuna can point the awe-struck family to the markings on the shark's body that correspond to the clothing in which the beloved's body had been wrapped. Such a shark aumakua becomes the family pet, receiving food, and driving fish into the family net and warding off danger. Like all aumakua it had evil uses such as helping kill enemies. The ruling chiefs typically forbade such sorcery. Many Native Hawaiian families claim such an aumakua, who is known by name to the whole community.[102]
+Kamohoali'i is the best known and revered of the shark gods, he was the older and favored brother of Pele,[103] and helped and journeyed with her to Hawaii. He was able to assume all human and fish forms. A summit cliff on the crater of Kilauea is one of his most sacred spots. At one point he had a heiau (temple or shrine) dedicated to him on every piece of land that jutted into the ocean on the island of Molokai. Kamohoali'i was an ancestral god, not a human who became a shark and banned the eating of humans after eating one herself.[104][105] In Fijian mythology, Dakuwaqa was a shark god who was the eater of lost souls.
+On the island of Tutuila in American Samoa (a U.S. territory), there is a location called Turtle and Shark (Laumei ma Malie) which is important in Samoan culture — the location is the site of a legend called O Le Tala I Le Laumei Ma Le Malie, in which two humans are said to have transformed into a turtle and a shark.[106][107][108] According to the U.S. National Park Service, "Villagers from nearby Vaitogi continue to reenact an important aspect of the legend at Turtle and Shark by performing a ritual song intended to summon the legendary animals to the ocean surface, and visitors are frequently amazed to see one or both of these creatures emerge from the sea in apparent response to this call."[106]
+In contrast to the complex portrayals by Hawaiians and other Pacific Islanders, the European and Western view of sharks has historically been mostly of fear and malevolence.[109] Sharks are used in popular culture commonly as eating machines, notably in the Jaws novel and the film of the same name, along with its sequels.[110] Sharks are threats in other films such as Deep Blue Sea, The Reef, and others, although they are sometimes used for comedic effect such as in Finding Nemo and the Austin Powers series.
+Sharks tend to be seen quite often in cartoons whenever a scene involves the ocean. Such examples include the Tom and Jerry cartoons, Jabberjaw, and other shows produced by Hanna-Barbera. They also are used commonly as a clichéd means of killing off a character that is held up by a rope or some similar object as the sharks swim right below them, or the character may be standing on a plank above shark infested waters.[citation needed]
+A popular myth is that sharks are immune to disease and cancer, but this is not scientifically supported. Sharks have been known to get cancer.[111][112] Both diseases and parasites affect sharks. The evidence that sharks are at least resistant to cancer and disease is mostly anecdotal and there have been few, if any, scientific or statistical studies that show sharks to have heightened immunity to disease.[113]
+Other apparently false claims are that fins prevent cancer[114] and treat osteoarthritis.[115] No scientific proof supports these claims; at least one study has shown shark cartilage of no value in cancer treatment.[116]
+It is estimated that 100 million sharks are killed by people every year, due to commercial and recreational fishing.[117][118]
+Shark finning yields are estimated at 1.44 million metric tons for 2000, and 1.41 million tons for 2010. Based on an analysis of average shark weights, this translates into a total annual mortality estimate of about 100 million sharks in 2000, and about 97 million sharks in 2010, with a total range of possible values between 63 and 273 million sharks per year.[119][120] Sharks are a common seafood in many places, including Japan and Australia. In the Australian state of Victoria, shark is the most commonly used fish in fish and chips,[citation needed] in which fillets are battered and deep-fried or crumbed and grilled. In fish and chip shops, shark is called flake. In India, small sharks or baby sharks (called sora in Tamil language, Telugu language) are sold in local markets. Since the flesh is not developed, cooking the flesh breaks it into powder, which is then fried in oil and spices (called sora puttu/sora poratu). The soft bones can be easily chewed. They are considered a delicacy in coastal Tamil Nadu. Icelanders ferment Greenland sharks to produce a delicacy called hákarl.[121] During a four-year period from 1996 to 2000, an estimated 26 to 73 million sharks were killed and traded annually in commercial markets.[122]
+Sharks are often killed for shark fin soup. Fishermen capture live sharks, fin them, and dump the finless animal back into the water. Shark finning involves removing the fin with a hot metal blade.[118] The resulting immobile shark soon dies from suffocation or predators.[123] Shark fin has become a major trade within black markets all over the world. Fins sell for about $300/lb in 2009.[124] Poachers illegally fin millions each year. Few governments enforce laws that protect them.[120] In 2010 Hawaii became the first U.S. state to prohibit the possession, sale, trade or distribution of shark fins.[125] From 1996 to 2000, an estimated 38 million sharks had been killed per year for harvesting shark fins.[122] It is estimated by TRAFFIC that over 14,000 tonnes of shark fins were exported into Singapore between 2005–2007 and 2012–2014.[126]
+Shark fin soup is a status symbol in Asian countries and is erroneously considered healthy and full of nutrients. Scientific research has revealed, however, that high concentrations of BMAA are present in shark fins.[127] Because BMAA is a neurotoxin, consumption of shark fin soup and cartilage pills, therefore, may pose a health risk.[128] BMAA is under study for its pathological role in neurodegegerative diseases such as, ALS, Alzheimer's disease, and Parkinson's disease.
+Sharks are also killed for meat. European diners consume dogfishes, smoothhounds, catsharks, makos, porbeagle and also skates and rays.[129] However, the U.S. FDA lists sharks as one of four fish (with swordfish, king mackerel, and tilefish) whose high mercury content is hazardous to children and pregnant women.
+Sharks generally reach sexual maturity only after many years and produce few offspring in comparison to other harvested fish. Harvesting sharks before they reproduce severely impacts future populations. Capture induced premature birth and abortion (collectively called capture-induced parturition) occurs frequently in sharks/rays when fished.[66] Capture-induced parturition is rarely considered in fisheries management despite being shown to occur in at least 12% of live bearing sharks and rays (88 species to date).[66]
+The majority of shark fisheries have little monitoring or management. The rise in demand for shark products increases pressure on fisheries.[36] Major declines in shark stocks have been recorded—some species have been depleted by over 90% over the past 20–30 years with population declines of 70% not unusual.[130] A study by the International Union for Conservation of Nature suggests that one quarter of all known species of sharks and rays are threatened by extinction and 25 species were classified as critically endangered.[131][132]
+In 2014, a shark cull in Western Australia killed dozens of sharks (mostly tiger sharks) using drum lines,[133] until it was cancelled after public protests and a decision by the Western Australia EPA; from 2014 to 2017, there was an "imminent threat" policy in Western Australia in which sharks that "threatened" humans in the ocean were shot and killed.[134] This "imminent threat" policy was criticized by senator Rachel Siewart for killing endangered sharks.[135] The "imminent threat" policy was cancelled in March 2017.[136] In August 2018, the Western Australia government announced a plan to re-introduce drum lines (though, this time the drum lines are "SMART" drum lines).[137]
+From 1962 to the present,[138] the government of Queensland has targeted and killed sharks in large numbers by using drum lines, under a "shark control" program—this program has also inadvertently killed large numbers of other animals such as dolphins; it has also killed endangered hammerhead sharks.[139][140][141][142] Queensland's drum line program has been called "outdated, cruel and ineffective".[142] From 2001 to 2018, a total of 10,480 sharks were killed on lethal drum lines in Queensland, including in the Great Barrier Reef.[143] From 1962 to 2018, roughly 50,000 sharks were killed by Queensland authorities.[144]
+The government of New South Wales has a program that deliberately kills sharks using nets.[141][145] The current net program in New South Wales has been described as being "extremely destructive" to marine life, including sharks.[146] Between 1950 and 2008, 352 tiger sharks and 577 great white sharks were killed in the nets in New South Wales — also during this period, a total of 15,135 marine animals were killed in the nets, including dolphins, whales, turtles, dugongs, and critically endangered grey nurse sharks.[147] There has been a very large decrease in the number of sharks in eastern Australia, and the shark-killing programs in Queensland and New South Wales are partly responsible for this decrease.[144]
+Kwazulu-Natal, an area of South Africa, has a shark-killing program using nets and drum lines—these nets and drum lines have killed turtles and dolphins, and have been criticized for killing wildlife.[148] During a 30-year period, more than 33,000 sharks have been killed in KwaZulu-Natal's shark-killing program — during the same 30-year period, 2,211 turtles, 8,448 rays, and 2,310 dolphins were killed in KwaZulu-Natal.[148] Authorities on the French island of Réunion kill about 100 sharks per year.[149]
+Killing sharks negatively affects the marine ecosystem.[150][151] Jessica Morris of Humane Society International calls shark culling a "knee-jerk reaction" and says, "sharks are top order predators that play an important role in the functioning of marine ecosystems. We need them for healthy oceans."[152]
+George H. Burgess, the former[153] director of the International Shark Attack File, "describes [shark] culling as a form of revenge, satisfying a public demand for blood and little else";[154] he also said shark culling is a "retro-type move reminiscent of what people would have done in the 1940s and 50s, back when we didn't have an ecological conscience and before we knew the consequences of our actions."[154] Jane Williamson, an associate professor in marine ecology at Macquarie University, says "There is no scientific support for the concept that culling sharks in a particular area will lead to a decrease in shark attacks and increase ocean safety."[155]
+Other threats include habitat alteration, damage and loss from coastal development, pollution and the impact of fisheries on the seabed and prey species.[156] The 2007 documentary Sharkwater exposed how sharks are being hunted to extinction.[157]
+In 1991, South Africa was the first country in the world to declare Great White sharks a legally protected species[158] (however, the KwaZulu-Natal Sharks Board is allowed to kill great white sharks in its "shark control" program in eastern South Africa).[148]
+Intending to ban the practice of shark finning while at sea, the United States Congress passed the Shark Finning Prohibition Act in 2000.[159]  Two years later the Act saw its first legal challenge in United States v. Approximately 64,695 Pounds of Shark Fins.  In 2008 a Federal Appeals Court ruled that a loophole in the law allowed non-fishing vessels to purchase shark fins from fishing vessels while on the high seas.[160]  Seeking to close the loophole, the Shark Conservation Act was passed by Congress in December 2010, and it was signed into law in January 2011.[161][162]
+In 2003, the European Union introduced a general shark finning ban for all vessels of all nationalities in Union waters and for all vessels flying a flag of one of its member states.[163] This prohibition was amended in June 2013 to close remaining loopholes.[164]
+In 2009, the International Union for Conservation of Nature's IUCN Red List of Endangered Species named 64 species, one-third of all oceanic shark species, as being at risk of extinction due to fishing and shark finning.[165][166]
+In 2010, the Convention on International Trade in Endangered Species (CITES) rejected proposals from the United States and Palau that would have required countries to strictly regulate trade in several species of scalloped hammerhead, oceanic whitetip and spiny dogfish sharks. The majority, but not the required two-thirds of voting delegates, approved the proposal. China, by far the world's largest shark market, and Japan, which battles all attempts to extend the convention to marine species, led the opposition.[167][168] In March 2013, three endangered commercially valuable sharks, the hammerheads, the oceanic whitetip and porbeagle were added to Appendix 2 of CITES, bringing shark fishing and commerce of these species under licensing and regulation.[169]
+In 2010, Greenpeace International added the school shark, shortfin mako shark, mackerel shark, tiger shark and spiny dogfish to its seafood red list, a list of common supermarket fish that are often sourced from unsustainable fisheries.[170] Advocacy group Shark Trust campaigns to limit shark fishing. Advocacy group Seafood Watch directs American consumers to not eat sharks.[171]
+Under the auspices of the Convention on the Conservation of Migratory Species of Wild Animals (CMS), also known as the Bonn Convention, the Memorandum of Understanding on the Conservation of Migratory Sharks was concluded and came into effect in March 2010. It was the first global instrument concluded under CMS and aims at facilitating international coordination for the protection, conservation and management of migratory sharks, through multilateral, intergovernmental discussion and scientific research.
+In July 2013, New York state, a major market and entry point for shark fins, banned the shark fin trade joining
+seven other states of the United States and the three Pacific U.S territories in providing legal protection to sharks.[172]
+In the United States, and as of January 16, 2019, 12 states including (Massachusetts, Maryland, Delaware, California, Illinois, Hawaii, Oregon, Nevada, Rhode Island, Washington, New York and Texas) along with 3 U.S. territories (American Samoa, Guam and the Northern Mariana Islands) have passed laws against the sale or possession of shark fins.[173][174]
+Several regions now have shark sanctuaries or have banned shark fishing — these regions include American Samoa, the Bahamas, the Cook Islands, French Polynesia, Guam, the Maldives, the Marshall Islands, Micronesia, the Northern Mariana Islands, and Palau.[175][176]

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+The Rhine (Latin: Rhenus [ˈreːnus], Romansh: Rein, German: Rhein [ʁaɪ̯n], French: Rhin,[1] Italian: Reno, Dutch: Rijn, Alemannic German: Rhi(n) including Alsatian/Low Alemannic German) is one of the major European rivers, which has its sources in Switzerland and flows in a mostly northerly direction through Germany and the Netherlands, emptying into the North Sea. The river begins in the Swiss canton of Graubünden in the southeastern Swiss Alps, forms part of the Swiss-Liechtenstein, Swiss-Austrian, Swiss-German and then the Franco-German border, then flows through the German Rhineland and the Netherlands and eventually empties into the North Sea.
+It is the second-longest river in Central and Western Europe (after the Danube), at about 1,230 km (760 mi),[note 1][note 2] with an average discharge of about 2,900 m3/s (100,000 cu ft/s).
+The Rhine and the Danube formed most of the northern inland frontier of the Roman Empire and, since those days, the Rhine has been a vital and navigable waterway carrying trade and goods deep inland.
+Its importance as a waterway in the Holy Roman Empire is supported by the many castles and fortifications built along it. In the modern era, it has become a symbol of German nationalism.
+Among the largest and most important cities on the Rhine are Cologne, Düsseldorf, Rotterdam, Strasbourg and Basel.
+The variants of the name of the Rhine in modern languages are all derived from the Gaulish name Rēnos, which was adapted in Roman-era geography (1st century BC) as Greek Ῥῆνος (Rhēnos), Latin Rhenus.[note 3]
+The spelling with Rh- in English Rhine as well as in German Rhein and French Rhin is due to the influence of Greek orthography, while the vocalisation -i- is due to the Proto-Germanic adoption of the Gaulish name as *Rīnaz, via Old Frankish giving Old English Rín,[3]
+Old High German Rīn, early Middle Dutch (c. 1200) Rijn (then also spelled Ryn or Rin).[4]
+The diphthong in modern German Rhein (also adopted in Romansh Rein, Rain) is a Central German development of the early modern period, the Alemannic name Rī(n) retaining the older vocalism,[note 4] as does Ripuarian Rhing, while Palatine has diphthongized Rhei, Rhoi. Spanish is with French in adopting the Germanic vocalism Rin-, while Italian, Occitan and Portuguese retain the Latin Ren-.
+The Gaulish name Rēnos (Proto-Celtic or pre-Celtic[note 5] *Reinos) belongs to a class of river names built from the PIE root *rei- "to move, flow, run", also found in other names such as the Reno in Italy.[note 6]
+The grammatical gender of the Celtic name (as well as of its Greek and Latin adaptation) is masculine, and the name remains masculine in German, Dutch and French.
+The Old English river name was variously inflected as masculine or feminine; and its Old Icelandic adoption was inflected as feminine.[5]
+The length of the Rhine is conventionally measured in "Rhine-kilometers" (Rheinkilometer), a scale introduced in 1939 which runs from the Old Rhine Bridge at Constance (0 km) to Hoek van Holland (1036.20 km).
+The river is significantly shortened from its natural course due to a number of canalisation projects completed in the 19th and 20th century.[note 7] The "total length of the Rhine", to the inclusion of Lake Constance and the Alpine Rhine is more difficult to measure objectively; it was cited as 1,232 kilometres (766 miles) by the Dutch Rijkswaterstaat in 2010.[note 1]
+Its course is conventionally divided as follows:
+The Rhine carries its name without distinctive accessories only from the confluence of the Rein Anteriur/Vorderrhein and Rein Posteriur/Hinterrhein next to Reichenau in Tamins. Above this point is the extensive catchment of the headwaters of the Rhine. It belongs almost exclusively to the Swiss canton of Graubünden, ranging from Saint-Gotthard Massif in the west via one valley lying in Ticino and Italy in the south to the Flüela Pass in the east.
+Traditionally, Lake Toma near the Oberalp Pass in the Gotthard region is seen as the source of the Anterior Rhine and the Rhine as a whole. The Posterior Rhine rises in the Rheinwald below the Rheinwaldhorn.
+The source of the river is generally considered north of Lai da Tuma/Tomasee on Rein Anteriur/Vorderrhein,[13] although its southern tributary Rein da Medel is actually longer before its confluence with the Anterior Rhine near Disentis.
+The Anterior Rhine arises from numerous source streams in the upper Surselva and flows in an easterly direction. One source is Lai da Tuma (2,345 m (7,694 ft))[14] with the Rein da Tuma, which is usually indicated as source of the Rhine, flowing through it.
+Into it flow tributaries from the south, some longer, some equal in length, such as the Rein da Medel, the Rein da Maighels, and the Rein da Curnera. The Cadlimo Valley in the canton of Ticino is drained by the Reno di Medel, which crosses the geomorphologic Alpine main ridge from the south.[note 8] All streams in the source area are partially, sometimes completely, captured and sent to storage reservoirs for the local hydro-electric power plants.
+The culminating point of the Anterior Rhine's drainage basin is the Piz Russein of the Tödi massif of the Glarus Alps at 3,613 metres (11,854 ft) above sea level. It starts with the creek Aua da Russein (lit.: "Water of the Russein").[15]
+In its lower course the Anterior Rhine flows through a gorge named Ruinaulta (Flims Rockslide). The whole stretch of the Anterior Rhine to the Alpine Rhine confluence next to Reichenau in Tamins is accompanied by a long-distance hiking trail called Senda Sursilvana.[16]
+The Posterior Rhine flows first east-northeast, then north. It flows through the three valleys named Rheinwald, Schams and Domleschg-Heinzenberg. The valleys are separated by the Rofla Gorge and Viamala Gorge. Its sources are located in the Adula Alps (Rheinwaldhorn, Rheinquellhorn, and Güferhorn).
+The Avers Rhine joins from the south. One of its headwaters, the Reno di Lei (stowed in the Lago di Lei), is partially located in Italy.
+Near Sils the Posterior Rhine is joined by the Albula, from the east, from the Albula Pass region. The Albula draws its water mainly from the Landwasser with the Dischmabach as the largest source stream, but almost as much from the Gelgia, which comes down from the Julier Pass.
+Numerous larger and smaller tributary rivers bear the name of the Rhine or equivalent in various Romansh idioms like Rein or Ragn. Examples:
+Next to Reichenau in Tamins the Anterior Rhine and the Posterior Rhine join and form the Alpine Rhine. The river makes a distinctive turn to the north near Chur. This section is nearly 86 km long, and descends from a height of 599 m to 396 m. It flows through a wide glacial Alpine valley known as the Rhine Valley (German: Rheintal). Near Sargans a natural dam, only a few metres high, prevents it from flowing into the open Seeztal valley and then through Lake Walen and Lake Zurich into the Aare. The Alpine Rhine begins in the westernmost part of the Swiss canton of Graubünden, and later forms the border between Switzerland to the west and Liechtenstein and later Austria to the east.
+As an effect of human work, it empties into Lake Constance on Austrian territory and not on the border that follows its old natural river bed.
+The mouth of the Rhine into Lake Constance forms an inland delta. The delta is delimited in the west by the Alter Rhein ("Old Rhine") and in the east by a modern canalized section. Most of the delta is a nature reserve and bird sanctuary. It includes the Austrian towns of Gaißau, Höchst and Fußach. The natural Rhine originally branched into at least two arms and formed small islands by precipitating sediments. In the local Alemannic dialect, the singular is pronounced "Isel" and this is also the local pronunciation of Esel ("Donkey"). Many local fields have an official name containing this element.
+A regulation of the Rhine was called for, with an upper canal near Diepoldsau and a lower canal at Fußach, in order to counteract the constant flooding and strong sedimentation in the western Rhine Delta. The Dornbirner Ach had to be diverted, too, and it now flows parallel to the canalized Rhine into the lake. Its water has a darker color than the Rhine; the latter's lighter suspended load comes from higher up the mountains. It is expected that the continuous input of sediment into the lake will silt up the lake. This has already happened to the former Lake Tuggenersee.
+The cut-off Old Rhine at first formed a swamp landscape. Later an artificial ditch of about two km was dug. It was made navigable to the Swiss town of Rheineck.
+Lake Constance consists of three bodies of water: the Obersee ("upper lake"), the Untersee ("lower lake"), and a connecting stretch of the Rhine, called the Seerhein ("Lake Rhine"). The lake is situated in Germany, Switzerland and Austria near the Alps. Specifically, its shorelines lie in the German states of Bavaria and Baden-Württemberg, the Austrian state of Vorarlberg, and the Swiss cantons of Thurgau and St. Gallen. The Rhine flows into it from the south following the Swiss-Austrian border. It is located at approximately 47°39′N 9°19′E / 47.650°N 9.317°E / 47.650; 9.317.
+The flow of cold, grey mountain water continues for some distance into the lake. The cold water flows near the surface and at first doesn't mix with the warmer, green waters of Upper Lake. But then, at the so-called Rheinbrech, the Rhine water abruptly falls into the depths because of the greater density of cold water. The flow reappears on the surface at the northern (German) shore of the lake, off the island of Lindau. The water then follows the northern shore until Hagnau am Bodensee. A small fraction of the flow is diverted off the island of Mainau into Lake Überlingen. Most of the water flows via the Constance hopper into the Rheinrinne ("Rhine Gutter") and Seerhein. Depending on the water level, this flow of the Rhine water is clearly visible along the entire length of the lake.
+The Rhine carries very large amounts of debris into the lake.[note 9] In the mouth region, it is therefore necessary to permanently remove gravel by dredging. The large sediment loads are partly due to the extensive land improvements upstream.
+Three countries border the Obersee, namely Switzerland in the south, Austria in the southeast and the German states of Bavaria in the northeast and Baden-Württemberg in the north and northwest.
+The Seerhein is only four km long. It connects the Obersee with the 30 cm lower Untersee. Distance markers along the Rhine measure the distance from the bridge in the old city centre of Constance.
+For most of its length, the Seerhein forms the border between Germany and Switzerland. The exception is the old city centre of Constance, on the Swiss side of the river.
+The Seerhein emerged in the last thousands of years, when erosion caused the lake level to be lowered by about 10 metres. Previously, the two lakes formed a single lake, as the name still suggests.
+Like in the Obersee, the flow the Rhine can be traced in the Untersee. Here, too, the river water is hardly mixed with the lake water. The northern parts of the Untersee (Lake Zell and Gnadensee) remain virtually unaffected by the flow. The river traverses the southern, which, in isolation, is sometimes called Rhinesee ("Lake Rhine").
+The Radolfzeller Aach adds large amounts of water from the Danube system to the Untersee.
+Reichenau Island was formed at the same time as the Seerhein, when the water level was lowered to its current level.
+Lake Untersee is part of the border between Switzerland and Germany, with Germany on the north bank and Switzerland on the south, except both sides are Swiss in Stein am Rhein, where the High Rhine flows out of the lake.
+The Rhine emerges from Lake Constance, flows generally westward, as the Hochrhein, passes the Rhine Falls, and is joined by its major tributary, the Aare. The Aare more than doubles the Rhine's water discharge, to an average of nearly 1,000 m3/s (35,000 cu ft/s), and provides more than a fifth of the discharge at the Dutch border. The Aare also contains the waters from the 4,274 m (14,022 ft) summit of Finsteraarhorn, the highest point of the Rhine basin. The Rhine roughly forms the German-Swiss border from Lake Constance with the exceptions of the canton of Schaffhausen and parts of the cantons of Zürich and Basel-Stadt, until it turns north at the so-called Rhine knee at Basel, leaving Switzerland.
+The High Rhine begins in Stein am Rhein at the western end of the Untersee. Unlike the Alpine Rhine and Upper Rhine, it flows to the west. It falls from 395 m to 252 m.
+Some stretches of the High Rhine between Stein am Rhein and Eglisau form the border between Switzerland on the south bank and Germany in the north. On other stretches, both sides are Swiss; in fact most of the Canton of Schaffhausen is on the north bank. Between Eglisau and Basel, the High Rhine consistently forms the border.
+The Rhine Falls are situated below Schaffhausen. It has an average water flow of 373 m³/s (mean summer discharge 700 m³/s) and is the second largest waterfall in Europe in terms of potential energy, after Dettifoss in Iceland. The High Rhine is characterized by numerous dams. On the few remaining natural sections, there are still several rapids.
+Near Koblenz in the Aargau, the Aare joins the Rhine. With an average discharge of 557 m³/s, the Aare is more voluminous than the Rhine, which has an average discharge of 439 m³/s. Nevertheless, the Alpine Rhine is considered the main branch, because it is longer.
+In the centre of Basel, the first major city in the course of the stream, is located the "Rhine knee"; this is a major bend, where the overall direction of the Rhine changes from west to north. Here the High Rhine ends. Legally, the Central Bridge is the boundary between High and Upper Rhine. The river now flows north as Upper Rhine through the Upper Rhine Plain, which is about 300 km long and up to 40 km wide. The most important tributaries in this area are the Ill below of Strasbourg, the Neckar in Mannheim and the Main across from Mainz. In Mainz, the Rhine leaves the Upper Rhine Valley and flows through the Mainz Basin.
+The southern half of the Upper Rhine forms the border between France (Alsace) and Germany (Baden-Württemberg). The northern part forms the border between the German states of Rhineland-Palatinate in the west on the one hand, and Baden-Württemberg and Hesse on the other hand, in the east and north. A curiosity of this border line is that the parts of the city of Mainz on the right bank of the Rhine were given to Hesse by the occupying forces in 1945.
+The Upper Rhine was a significant cultural landscape in Central Europe already in antiquity and during the Middle Ages. Today, the Upper Rhine area hosts many important manufacturing and service industries, particularly in the centers Basel, Strasbourg and Mannheim-Ludwigshafen. Strasbourg is the seat of the European Parliament, and so one of the three European capitals is located on the Upper Rhine.
+The Upper Rhine region was changed significantly by a Rhine straightening program in the 19th Century. The rate of flow was increased and the ground water level fell significantly. Dead branches were removed by construction workers and the area around the river was made more habitable for humans on flood plains as the rate of flooding decreased sharply. On the French side, the Grand Canal d'Alsace was dug, which carries a significant part of the river water, and all of the traffic. In some places, there are large compensation pools, for example, the huge Bassin de compensation de Plobsheim in Alsace.
+The Upper Rhine has undergone significant human change since the 19th century. While it was slightly modified during the Roman occupation, it was not until the emergence of engineers such as Johann Gottfried Tulla that significant modernization efforts changed the shape of the river. Earlier work under Frederick the Great surrounded efforts to ease shipping and construct dams to serve coal transportation.[17] Tulla is considered to have domesticated the Upper Rhine, domestication that served goals such as reducing stagnant bogs that fostered waterborne diseases, making regions more habitable for human settlement, and reduce high frequency of flooded water. Not long before Tulla went to work on widening and straightening the river heavy floods had brought significant loss of life.[18] Four diplomatic treaties were signed among German state governments and French regions dealing with the changes proposed along the Rhine, one was "the Treaty for the Rectification of the Rhine flow from Neuberg to Dettenheim"(1817), which surrounded states such as Bourbon France and the Bavarian Palatinate. Loops, oxbows, branches and islands were removed along the Upper Rhine so that there would be a present uniformity to the river.[19] The engineering of the Rhine was not without protest, farmers and fishermen had grave concerns about valuable fishing areas and farmland being lost. While some areas lost ground, other areas saw swamps and bogs be drained and turned into arable land.[20] Johann Tulla had the goal of shortening and straightening the Upper Rhine. Early engineering projects the Upper Rhine also had issues, with Tulla's project at one part of the river creating rapids, after the Rhine cut down from erosion to sheer rock.[21] Engineering along the Rhine eased flooding and made transportation along the river less cumbersome. These state projects were part of the advanced and technical progress efforts going on in the country alongside the industrial revolution. For the German state, to make the river more predictable was to ensure development projects could easily commence.[22]
+The section of the Upper Rhine downstream from Mainz is also known as the "Island Rhine". Here a number of river islands occur, locally known as "Rheinauen".
+The Rhine is the longest river in Germany. It is here that the Rhine encounters some more of its main tributaries, such as the Neckar, the Main and, later, the Moselle, which contributes an average discharge of more than 300 m3/s (11,000 cu ft/s). Northeastern France drains to the Rhine via the Moselle; smaller rivers drain the Vosges and Jura Mountains uplands. Most of Luxembourg and a very small part of Belgium also drain to the Rhine via the Moselle. As it approaches the Dutch border, the Rhine has an annual mean discharge of 2,290 m3/s (81,000 cu ft/s) and an average width of 400 m (1,300 ft).
+Between Bingen am Rhein and Bonn, the Middle Rhine flows through the Rhine Gorge, a formation which was created by erosion. The rate of erosion equaled the uplift in the region, such that the river was left at about its original level while the surrounding lands raised. The gorge is quite deep and is the stretch of the river which is known for its many castles and vineyards. It is a UNESCO World Heritage Site (2002) and known as "the Romantic Rhine", with more than 40 castles and fortresses from the Middle Ages and many quaint and lovely country villages.
+Between Strasbourg and Kehl
+Aerial photo between Eltville and Bingen
+Marksburg near Koblenz, built in 1231
+Ludendorff Bridge at Remagen, showing damage before collapse during the Battle of Remagen
+The Rhine in Cologne
+Rhine at Düsseldorf
+Rhine at Hünxe, near the border of the Netherlands
+The Mainz Basin ends in Bingen am Rhein; the Rhine continues as "Middle Rhine" into the Rhine Gorge in the Rhenish Slate Mountains. In this sections the river falls from 77.4 m above sea level to 50.4 m. On the left, is located the mountain ranges of Hunsrück and Eifel, on the right Taunus and Westerwald. According to geologists, the characteristic narrow valley form was created by erosion by the river while the surrounding landscape was lifted (see water gap).
+Major tributaries in this section are the Lahn and the Moselle. They join the Rhine near Koblenz, for the right and left respectively. Almost the entire length of the Middle Rhine runs in the German state of Rhineland-Palatinate.
+The dominant economic sectors in the Middle Rhine area are viniculture and tourism. The Rhine Gorge between Rüdesheim am Rhein and Koblenz is listed as a UNESCO World Heritage Site. Near Sankt Goarshausen, the Rhine flows around the famous rock Lorelei. With its outstanding architectural monuments, the slopes full of vines, settlements crowded on the narrow river banks and scores of castles lined up along the top of the steep slopes, the Middle Rhine Valley can be considered the epitome of the Rhine romanticism.
+In Bonn, where the Sieg flows into the Rhine, the Rhine enters the North German Plain and turns into the Lower Rhine. The Lower Rhine falls from 50 m to 12 m. The main tributaries on this stretch are the Ruhr and the Lippe. Like the Upper Rhine, the Lower Rhine used to meander until engineering created a solid river bed. Because the levees are some distance from the river, at high tide the Lower Rhine has more room for widening than the Upper Rhine.
+The Lower Rhine flows through North Rhine-Westphalia. Its banks are usually heavily populated and industrialized, in particular the agglomerations Cologne, Düsseldorf and Ruhr area. Here the Rhine flows through the largest conurbation in Germany, the Rhine-Ruhr region. One of the most important cities in this region is Duisburg with the largest river port in Europe (Duisport). The region downstream of Duisburg is more agricultural. In Wesel, 30 km downstream of Duisburg, is located the western end of the second east–west shipping route, the Wesel-Datteln Canal, which runs parallel to the Lippe. Between Emmerich and Cleves the Emmerich Rhine Bridge, the longest suspension bridge in Germany, crosses the 400-metre-wide (1,300 ft) river. Near Krefeld, the river crosses the Uerdingen line, the line which separates the areas where Low German and High German are spoken.
+Until the early 1980s, industry was a major source of water pollution. Although many plants and factories can be found along the Rhine up into Switzerland, it is along the Lower Rhine that the bulk of them are concentrated, as the river passes the major cities of Cologne, Düsseldorf and Duisburg. Duisburg is the home of Europe's largest inland port and functions as a hub to the sea ports of Rotterdam, Antwerp and Amsterdam. The Ruhr, which joins the Rhine in Duisburg, is nowadays a clean river, thanks to a combination of stricter environmental controls, a transition from heavy industry to light industry and cleanup measures, such as the reforestation of Slag and brownfields. The Ruhr currently provides the region with drinking water. It contributes 70 m3/s (2,500 cu ft/s) to the Rhine. Other rivers in the Ruhr Area, above all, the Emscher, still carry a considerable degree of pollution.
+The Dutch name for Rhine is "Rijn". The Rhine turns west and enters the Netherlands, where, together with the rivers Meuse and Scheldt, it forms the extensive Rhine-Meuse-Scheldt delta, with 25,347 km2 (9,787 sq mi) the largest river delta in Europe.[23] Crossing the border into the Netherlands at Spijk, close to Nijmegen and Arnhem, the Rhine is at its widest, although the river then splits into three main distributaries: the Waal, Nederrijn ("Nether Rhine") and IJssel.
+From here, the situation becomes more complicated, as the Dutch name Rijn no longer coincides with the main flow of water. Two thirds of the water flow volume of the Rhine flows farther west, through the Waal and then, via the Merwede and Nieuwe Merwede (De Biesbosch), merging with the Meuse, through the Hollands Diep and Haringvliet estuaries, into the North Sea. The Beneden Merwede branches off, near Hardinxveld-Giessendam and continues as the Noord, to join the Lek, near the village of Kinderdijk, to form the Nieuwe Maas; then flows past Rotterdam and continues via Het Scheur and the Nieuwe Waterweg, to the North Sea. The Oude Maas branches off, near Dordrecht, farther down rejoining the Nieuwe Maas to form Het Scheur.
+The other third of the water flows through the Pannerdens Kanaal and redistributes in the IJssel and Nederrijn. The IJssel branch carries one ninth of the water flow of the Rhine north into the IJsselmeer (a former bay), while the Nederrijn carries approximately two ninths of the flow west along a route parallel to the Waal. However, at Wijk bij Duurstede, the Nederrijn changes its name and becomes the Lek. It flows farther west, to rejoin the Noord into the Nieuwe Maas and to the North Sea.
+The name Rijn, from here on, is used only for smaller streams farther to the north, which together formed the main river Rhine in Roman times. Though they retained the name, these streams no longer carry water from the Rhine, but are used for draining the surrounding land and polders. From Wijk bij Duurstede, the old north branch of the Rhine is called Kromme Rijn ("Bent Rhine") past Utrecht, first Leidse Rijn ("Rhine of Leiden") and then, Oude Rijn ("Old Rhine"). The latter flows west into a sluice at Katwijk, where its waters can be discharged into the North Sea. This branch once formed the line along which the Limes Germanicus were built. During periods of lower sea levels within the various ice ages, the Rhine took a left turn, creating the Channel River, the course of which now lies below the English Channel.
+The Rhine-Meuse Delta, the most important natural region of the Netherlands begins near Millingen aan de Rijn, close to the Dutch-German border with the division of the Rhine into Waal and Nederrijn. Since the Rhine contributes most of the water, the shorter term Rhine Delta is commonly used. However, this name is also used for the river delta where the Rhine flows into Lake Constance, so it is clearer to call the larger one Rhine-Meuse delta, or even Rhine–Meuse–Scheldt delta, as the Scheldt ends in the same delta.
+The shape of the Rhine delta is determined by two bifurcations: first, at Millingen aan de Rijn, the Rhine splits into Waal and Pannerdens Kanaal, which changes its name to Nederrijn at Angeren, and second near Arnhem, the IJssel branches off from the Nederrijn. This creates three main flows, two of which change names rather often. The largest and southern main branch begins as Waal and continues as Boven Merwede ("Upper Merwede"), Beneden Merwede ("Lower Merwede"), Noord ("the North"), Nieuwe Maas ("New Meuse"), Het Scheur ("the Rip") and Nieuwe Waterweg ("New Waterway"). The middle flow begins as Nederrijn, then changes into Lek, then joins the Noord, thereby forming Nieuwe Maas. The northern flow keeps the name IJssel until it flows into Lake IJsselmeer. Three more flows carry significant amounts of water: the Nieuwe Merwede ("New Merwede"), which branches off from the southern branch where it changes from Boven to Beneden Merwede; the Oude Maas ("Old Meuse"), which branches off from the southern branch where it changes from Beneden Merwede into Noord, and Dordtse Kil, which branches off from Oude Maas.
+Before the St. Elizabeth's flood (1421), the Meuse flowed just south of today's line Merwede-Oude Maas to the North Sea and formed an archipelago-like estuary with Waal and Lek. This system of numerous bays, estuary-like extended rivers, many islands and constant changes of the coastline, is hard to imagine today. From 1421 to 1904, the Meuse and Waal merged further upstream at Gorinchem to form Merwede. For flood protection reasons, the Meuse was separated from the Waal through a lock and diverted into a new outlet called "Bergse Maas", then Amer and then flows into the former bay Hollands Diep.
+The northwestern part of the estuary (around Hook of Holland), is still called Maasmond ("Meuse Mouth"), ignoring the fact that it now carries only water from the Rhine. This might explain the confusing naming of the various branches.
+The hydrography of the current delta is characterized by the delta's main arms, disconnected arms (Hollandse IJssel, Linge, Vecht, etc.) and smaller rivers and streams. Many rivers have been closed ("dammed") and now serve as drainage channels for the numerous polders. The construction of Delta Works changed the Delta in the second half of the 20th Century fundamentally. Currently Rhine water runs into the sea, or into former marine bays now separated from the sea, in five places, namely at the mouths of the Nieuwe Merwede, Nieuwe Waterway (Nieuwe Maas), Dordtse Kil, Spui and IJssel.
+The Rhine-Meuse Delta is a tidal delta, shaped not only by the sedimentation of the rivers, but also by tidal currents. This meant that high tide formed a serious risk because strong tidal currents could tear huge areas of land into the sea. Before the construction of the Delta Works, tidal influence was palpable up to Nijmegen, and even today, after the regulatory action of the Delta Works, the tide acts far inland. At the Waal, for example, the most landward tidal influence can be detected between Brakel and Zaltbommel.
+The Rhine flows from the Alps to the North Sea Basin; the geography and geology of its present-day watershed has been developing, since the Alpine orogeny began.
+In southern Europe, the stage was set in the Triassic Period of the Mesozoic Era, with the opening of the Tethys Ocean, between the Eurasian and African tectonic plates, between about 240 MBP and 220 MBP (million years before present). The present Mediterranean Sea descends from this somewhat larger Tethys sea. At about 180 MBP, in the Jurassic Period, the two plates reversed direction and began to compress the Tethys floor, causing it to be subducted under Eurasia and pushing up the edge of the latter plate in the Alpine Orogeny of the Oligocene and Miocene Periods. Several microplates were caught in the squeeze and rotated or were pushed laterally, generating the individual features of Mediterranean geography: Iberia pushed up the Pyrenees; Italy, the Alps, and Anatolia, moving west, the mountains of Greece and the islands. The compression and orogeny continue today, as shown by the ongoing raising of the mountains a small amount each year and the active volcanoes.
+In northern Europe, the North Sea Basin had formed during the Triassic and Jurassic periods and continued to be a sediment receiving basin since. In between the zone of Alpine orogeny and North Sea Basin subsidence, remained highlands resulting from an earlier orogeny (Variscan), such as the Ardennes, Eifel and Vosges.
+From the Eocene onwards, the ongoing Alpine orogeny caused a N–S rift system to develop in this zone. The main elements of this rift are the Upper Rhine Graben, in southwest Germany and eastern France and the Lower Rhine Embayment, in northwest Germany and the southeastern Netherlands. By the time of the Miocene, a river system had developed in the Upper Rhine Graben, that continued northward and is considered the first Rhine river. At that time, it did not yet carry discharge from the Alps; instead, the watersheds of the Rhone and Danube drained the northern flanks of the Alps.
+The watershed of the Rhine reaches into the Alps today, but it did not start out that way.[24] In the Miocene period, the watershed of the Rhine reached south, only to the Eifel and Westerwald hills, about 450 km (280 mi) north of the Alps. The Rhine then had the Sieg as a tributary, but not yet the Moselle. The northern Alps were then drained by the Danube.
+Through stream capture, the Rhine extended its watershed southward. By the Pliocene period, the Rhine had captured streams down to the Vosges Mountains, including  the Main and the Neckar. The northern Alps were then drained by the Rhone. By the early Pleistocene period, the Rhine had captured most of its current Alpine watershed from the Rhône, including the Aare. Since that time, the Rhine has added the watershed above Lake Constance (Vorderrhein, Hinterrhein, Alpenrhein; captured from the Rhône), the upper reaches of the Main, beyond Schweinfurt and the Moselle in the Vosges Mountains, captured during the Saale Ice-age from the Meuse, to its watershed.
+Around 2.5 million years ago (ending 11,600 years ago) was the geological period of the Ice Ages. Since approximately 600,000 years ago, six major Ice Ages have occurred, in which sea level dropped 120 m (390 ft) and much of the continental margins became exposed. In the Early Pleistocene, the Rhine followed a course to the northwest, through the present North Sea. During the so-called Anglian glaciation (~450,000 yr BP, marine oxygen isotope stage 12), the northern part of the present North Sea was blocked by the ice and a large lake developed, that overflowed through the English Channel. This caused the Rhine's course to be diverted through the English Channel. Since then, during glacial times, the river mouth was located offshore of Brest, France and rivers, like the River Thames and the Seine, became tributaries to the Rhine. During interglacials, when sea level rose to approximately the present level, the Rhine built deltas, in what is now the Netherlands.
+The last glacial ran from ~74,000 (BP = Before Present), until the end of the Pleistocene (~11,600 BP). In northwest Europe, it saw two very cold phases, peaking around 70,000 BP and around 29,000–24,000 BP. The last phase slightly predates the global last ice age maximum (Last Glacial Maximum). During this time, the lower Rhine flowed roughly west through the Netherlands and extended to the southwest, through the English Channel and finally, to the Atlantic Ocean. The English Channel, the Irish Channel and most of the North Sea were dry land, mainly because sea level was approximately 120 m (390 ft) lower than today.
+Most of the Rhine's current course was not under the ice during the last Ice Age; although, its source must still have been a glacier. A tundra, with Ice Age flora and fauna, stretched across middle Europe, from Asia to the Atlantic Ocean. Such was the case during the Last Glacial Maximum, ca. 22,000–14,000 yr BP, when ice-sheets covered Scandinavia, the Baltics, Scotland and the Alps, but left the space between as open tundra. Loess (wind-blown topsoil dust) arose from the south and North Sea plain settling on the slopes of the Alps, Urals and the Rhine Valley, rendering the valleys facing the prevailing winds especially fertile.
+As northwest Europe slowly began to warm up from 22,000 years ago onward, frozen subsoil and expanded alpine glaciers began to thaw and fall-winter snow covers melted in spring. Much of the discharge was routed to the Rhine and its downstream extension.[25] Rapid warming and changes of vegetation, to open forest, began about 13,000 BP. By 9000 BP, Europe was fully forested. With globally shrinking ice-cover, ocean water levels rose and the English Channel and North Sea re-inundated. Meltwater, adding to the ocean and land subsidence, drowned the former coasts of Europe transgressionally.
+About 11000 years ago, the Rhine estuary was in the Strait of Dover. There remained some dry land in the southern North Sea, known as Doggerland, connecting mainland Europe to Britain. About 9000 years ago, that last divide was overtopped / dissected. Man was already resident in the area when these events happened.
+Since 7500 years ago the situation of tides, currents and land-forms has resembled the present. Rates of sea level rise dropped such that natural sedimentation by the Rhine and coastal processes widely compensate for transgression by the sea. In the southern North Sea, due to ongoing tectonic subsidence, the coastline and sea bed are sinking at the rate of about 1–3 cm (0.39–1.18 in) per century (1 metre or 39 inches in last 3000 years).
+About 7000–5000 BP, a general warming encouraged migration of all former ice-locked areas, including up the Danube and down the Rhine by peoples to the east. A sudden massive expansion of the Black Sea as the Mediterranean Sea burst into it through the Bosporus may have occurred about 7500 BP.
+At the begin of the Holocene (~11,700 years ago), the Rhine occupied its Late-Glacial valley. As a meandering river, it reworked its ice-age floodplain. As sea-level rise continued in the Netherlands, the formation of the Holocene Rhine-Meuse delta began (~8,000 years ago). Coeval absolute sea-level rise and tectonic subsidence have strongly influenced delta evolution. Other factors of importance to the shape of the delta are the local tectonic activities of the Peel Boundary Fault, the substrate and geomorphology, as inherited from the Last Glacial and the coastal-marine dynamics, such as barrier and tidal inlet formations.[26]
+Since ~3000 yr BP (= years Before Present), human impact is seen in the delta. As a result of increasing land clearance (Bronze Age agriculture), in the upland areas (central Germany), the sediment load of the Rhine has strongly increased[27] and delta growth has speeded up.[28] This has caused increased flooding and sedimentation, ending peat formation in the delta. In the geologically recent past the main process distributing sediment across the delta has been the shifting of river channels to new locations on the floodplain (termed avulsion). Over the past 6000 years, approximately 80 avulsions have occurred.[24] Direct human impact in the delta began with the mining of peat for salt and fuel from Roman times onward. This was followed by embankment of the major distributaries and damming of minor distributaries, which took place in the 11–13th century AD. Thereafter, canals were dug, bends were straightened and groynes were built to prevent the river's channels from migrating or silting up.
+At present, the branches Waal and Nederrijn-Lek discharge to the North Sea through the former Meuse estuary, near Rotterdam. The river IJssel branch flows to the north and enters the IJsselmeer (formerly the Zuider Zee), initially a brackish lagoon but a freshwater lake since 1932. The discharge of the Rhine is divided into three branches: the Waal (6/9 of total discharge), the Nederrijn – Lek (2/9 of total discharge) and the IJssel (1/9 of total discharge). This discharge distribution has been maintained since 1709 by river engineering works including the digging of the Pannerdens canal and the installation, in the 20th century, of a series of weirs on the Nederrijn.
+The Rhine was not known to Herodotus and first enters the historical period in the 1st century BC in Roman-era geography. At that time, it formed the boundary between Gaul and Germania.
+The Upper Rhine had been part of the areal of the late Hallstatt culture since the 6th century BC, and by the 1st century BC, the areal of the La Tène culture covered almost its entire length, forming a contact zone with the Jastorf culture, i.e. the locus of early Celtic-Germanic cultural contact.
+In Roman geography, the Rhine formed the boundary between Gallia and Germania by definition; e.g. Maurus Servius Honoratus, Commentary on the Aeneid of Vergil (8.727) (Rhenus) fluvius Galliae, qui Germanos a Gallia dividit "(The Rhine is a) river of Gaul, which divides the Germanic people from Gaul."
+In Roman geography, the Rhine and Hercynia Silva were considered the boundary of the civilized world; as it was a wilderness, the Romans were eager to explore it. This view is typified by Res Gestae Divi Augusti, a long public inscription of Augustus, in which he boasts of his exploits; including, sending an expeditionary fleet north of the Rheinmouth, to Old Saxony and Jutland, which he claimed no Roman had ever done before.
+Augustus ordered his stepson Roman general Drusus to establish 50 military camps along the Rhine, starting the Germanic Wars in 12 BC. At this time, the plain of the Lower Rhine was the territory of the Ubii.
+The first urban settlement, on the grounds of what is today the centre of Cologne, along the Rhine, was Oppidum Ubiorum, which was founded in 38 BC by the Ubii. Cologne became acknowledged, as a city by the Romans in AD 50, by the name of Colonia Claudia Ara Agrippinensium.
+From the death of Augustus in AD 14 until after AD 70, Rome accepted as her Germanic frontier the water-boundary of the Rhine and upper Danube. Beyond these rivers she held only the fertile plain of Frankfurt, opposite the Roman border fortress of Moguntiacum (Mainz), the southernmost slopes of the Black Forest and a few scattered bridge-heads. The northern section of this frontier, where the Rhine is deep and broad, remained the Roman boundary until the empire fell. The southern part was different. The upper Rhine and upper Danube are easily crossed. The frontier which they form is inconveniently long, enclosing an acute-angled wedge of foreign territory between the modern Baden and Württemberg. The Germanic populations of these lands seem in Roman times to have been scanty, and Roman subjects from the modern Alsace-Lorraine had drifted across the river eastwards.
+The Romans kept eight legions in five bases along the Rhine. The number was reduced to four as more units were moved to the Danube. The actual number of legions present at any base or in all, depended on whether a state or threat of war existed. Between about AD 14 and 180, the assignment of legions was as follows: for the army of Germania Inferior, two legions at Vetera (Xanten), I Germanica and XX Valeria (Pannonian troops); two legions at oppidum Ubiorum ("town of the Ubii"), which was renamed to Colonia Agrippina, descending to Cologne, V Alaudae, a Celtic legion recruited from Gallia Narbonensis and XXI, possibly a Galatian legion from the other side of the empire.
+For the army of Germania Superior: one legion, II Augusta, at Argentoratum (Strasbourg); and one, XIII Gemina, at Vindonissa (Windisch). Vespasian had commanded II Augusta, before his promotion to imperator. In addition, were a double legion, XIV and XVI, at Moguntiacum (Mainz).
+The two original military districts of Germania Inferior and Germania Superior, came to influence the surrounding tribes, who later respected the distinction in their alliances and confederations. For example, the upper Germanic peoples combined into the Alemanni. For a time, the Rhine ceased to be a border, when the Franks crossed the river and occupied Roman-dominated Celtic Gaul, as far as Paris.
+Germanic tribes crossed the Rhine in the Migration period, by the 5th century establishing the kingdoms of Francia on the Lower Rhine, Burgundy on the Upper Rhine and Alemannia on the High Rhine. This "Germanic Heroic Age" is reflected in medieval legend, such as the Nibelungenlied which tells of the hero Siegfried killing a dragon on the Drachenfels (Siebengebirge) ("dragons rock"), near Bonn at the Rhine and of the Burgundians and their court at Worms, at the Rhine and Kriemhild's golden treasure, which was thrown into the Rhine by Hagen.
+By the 6th century, the Rhine was within the borders of Francia.
+In the 9th, it formed part of the border between Middle and Western Francia,
+but in the 10th century, it was fully within the Holy Roman Empire, flowing through Swabia, Franconia and Lower Lorraine. The mouths of the Rhine, in the county of Holland, fell to the Burgundian Netherlands in the 15th century; Holland remained contentious territory throughout the European wars of religion and the eventual collapse of the Holy Roman Empire, when the length of the Rhine fell to the First French Empire and its client states. The Alsace on the left banks of the Upper Rhine was sold to Burgundy by Archduke Sigismund of Austria in 1469 and eventually fell to France in the Thirty Years' War. The numerous historic castles in Rhineland-Palatinate attest to the importance of the river as a commercial route.
+Since the Peace of Westphalia, the Upper Rhine formed a contentious border between France and Germany. Establishing "natural borders" on the Rhine was a long-term goal of French foreign policy, since the Middle Ages, though the language border was – and is – far more to the west. French leaders, such as Louis XIV and Napoleon Bonaparte, tried with varying degrees of success to annex lands west of the Rhine. The Confederation of the Rhine was established by Napoleon, as a French client state, in 1806 and lasted until 1814, during which time it served as a significant source of resources and military manpower for the First French Empire. In 1840, the Rhine crisis, prompted by French prime minister Adolphe Thiers's desire to reinstate the Rhine as a natural border, led to a diplomatic crisis and a wave of nationalism in Germany.
+The Rhine became an important symbol in German nationalism during the formation of the German state in the 19th century (see Rhine romanticism).
+At the end of World War I, the Rhineland was subject to the Treaty of Versailles. This decreed that it would be occupied by the allies, until 1935 and after that, it would be a demilitarised zone, with the German army forbidden to enter. The Treaty of Versailles and this particular provision, in general, caused much resentment in Germany and is often cited as helping Adolf Hitler's rise to power. The allies left the Rhineland, in 1930 and the German army re-occupied it in 1936, which was enormously popular in Germany. Although the allies could probably have prevented the re-occupation, Britain and France were not inclined to do so, a feature of their policy of appeasement to Hitler.
+In World War II, it was recognised that the Rhine would present a formidable natural obstacle to the invasion of Germany, by the Western Allies. The Rhine bridge at Arnhem, immortalized in the book, A Bridge Too Far and the film, was a central focus of the battle for Arnhem, during the failed Operation Market Garden of September 1944. The bridges at Nijmegen, over the Waal distributary of the Rhine, were also an objective of Operation Market Garden. In a separate operation, the Ludendorff Bridge, crossing the Rhine at Remagen, became famous, when U.S. forces were able to capture it intact – much to their own surprise – after the Germans failed to demolish it. This also became the subject of a film, The Bridge at Remagen.
+Seven Days to the River Rhine was a Warsaw Pact war plan for an invasion of Western Europe during the Cold War.
+Until 1932 the generally accepted length of the Rhine was 1,230 kilometres (764 miles). In 1932 the German encyclopedia Knaurs Lexikon stated the length as 1,320 kilometres (820 miles), presumably a typographical error.
+After this number was placed into the authoritative Brockhaus Enzyklopädie, it became generally accepted and found its way into numerous textbooks and official publications. The error was discovered in 2010, and the Dutch Rijkswaterstaat confirms the length at 1,232 kilometres (766 miles).[note 1]
+Large cities that are situated on the Rhine:
+Switzerland:
+France:
+Germany:
+Netherlands:
+Smaller cities that are situated on the Rhine:
+Switzerland:
+Liechtenstein:
+Germany:
+Netherlands:
+During its course from the Alps to the North Sea, the Rhine passes through four countries and constitutes six different country borders. On the various parts:
+Order: panning north to south through the Western Netherlands:
+Order: upstream to downstream:

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+Illegitimate:
+Louis XIV (Louis Dieudonné; 5 September 1638 – 1 September 1715), known as  Louis the Great (Louis le Grand) or the Sun King (le Roi Soleil), was King of France from 14 May 1643 until his death in 1715. His reign of 72 years and 110 days is the longest recorded of any monarch of a sovereign country in European history.[1][a] Louis XIV's France was emblematic of the age of absolutism in Europe.[2]
+Louis began his personal rule of France in 1661, after the death of his chief minister, the Italian Cardinal Mazarin.[3] An adherent of the concept of the divine right of kings, Louis continued his predecessors' work of creating a centralised state governed from the capital. He sought to eliminate the remnants of feudalism persisting in parts of France and, by compelling many members of the nobility to inhabit his lavish Palace of Versailles, succeeded in pacifying the aristocracy, many members of which had participated in the Fronde rebellion during Louis' minority. By these means he became one of the most powerful French monarchs and consolidated a system of absolute monarchical rule in France that endured until the French Revolution. He also enforced uniformity of religion under the Gallican Catholic Church. His revocation of the Edict of Nantes abolished the rights of the Huguenot Protestant minority and subjected them to a wave of dragonnades, effectively forcing Huguenots to emigrate or convert, and virtually destroying the French Protestant community.
+The Sun King surrounded himself with a variety of significant political, military, and cultural figures such as Mazarin, Colbert, Louvois, the Grand Condé, Turenne, Vauban, Boulle, Molière, Racine, Boileau, La Fontaine, Lully, Charpentier, Marais, Le Brun, Rigaud, Bossuet, Le Vau, Mansart, Charles, Claude Perrault, and Le Nôtre.
+During Louis' long reign, France emerged as the leading European power, and regularly asserted its military strength. A conflict with Spain marked his entire childhood, while during his personal rule, the kingdom took part in three major continental conflicts, each against powerful foreign alliances: the Franco-Dutch War, the War of the League of Augsburg, and the War of the Spanish Succession. In addition, France also contested shorter wars such as the War of Devolution and the War of the Reunions. Warfare defined the foreign policy of Louis XIV and his personality shaped his approach. Impelled by "a mix of commerce, revenge, and pique", Louis sensed that warfare was the ideal way to enhance his glory. In peacetime he concentrated on preparing for the next war. He taught his diplomats that their job was to create tactical and strategic advantages for the French military.[4]
+Louis XIV was born on 5 September 1638 in the Château de Saint-Germain-en-Laye, to Louis XIII and Anne of Austria. He was named Louis Dieudonné (Louis the God-given)[5] and bore the traditional title of French heirs apparent: Dauphin.[6] At the time of his birth, his parents had been married for 23 years. His mother had experienced four stillbirths between 1619 and 1631. Leading contemporaries thus regarded him as a divine gift and his birth a miracle of God.
+Sensing imminent death, Louis XIII decided to put his affairs in order in the spring of 1643, when Louis XIV was four years old. In defiance of custom, which would have made Queen Anne the sole Regent of France, the king decreed that a regency council would rule on his son's behalf. His lack of faith in Queen Anne's political abilities was his primary rationale. He did, however, make the concession of appointing her head of the council.
+Louis' relationship with his mother was uncommonly affectionate for the time. Contemporaries and eyewitnesses claimed that the Queen would spend all her time with Louis. Both were greatly interested in food and theatre, and it is highly likely that Louis developed these interests through his close relationship with his mother. This long-lasting and loving relationship can be evidenced by excerpts in Louis' journal entries, such as:
+"Nature was responsible for the first knots which tied me to my mother. But attachments formed later by shared qualities of the spirit are far more difficult to break than those formed merely by blood."[7]
+It was his mother who gave Louis his belief in the absolute and divine power of his monarchical rule.[8]
+During his childhood, he was taken care of by the governesses Françoise de Lansac and Marie-Catherine de Senecey. In 1646, Nicolas V de Villeroy became the young king's tutor. Louis XIV became friends with Villeroy's young children, particularly François de Villeroy, and divided his time between the Palais-Royal and the nearby Hotel de Villeroy.
+On 14 May 1643, with Louis XIII dead, Queen Anne had her husband's will annulled by the Parlement de Paris (a judicial body comprising mostly nobles and high clergymen).[9] This action abolished the regency council and made Anne sole Regent of France. Anne exiled some of her husband's ministers (Chavigny, Bouthilier), and she nominated Brienne as her minister of foreign affairs.[10] Anne also nominated Saint Vincent de Paul as her spiritual adviser, which helped her deal with religious policy and the Jansenism question.[11]
+Anne kept the direction of religious policy strongly in her hand until 1661; her most important political decisions were to nominate Cardinal Mazarin as her chief minister and the continuation of her late husband's and Cardinal Richelieu's policy, despite their persecution of her, for the sake of her son. Anne wanted to give her son absolute authority and a victorious kingdom. Her rationales for choosing Mazarin were mainly his ability and his total dependence on her, at least until 1653 when she was no longer regent. Anne protected Mazarin by arresting and exiling her followers who conspired against him in 1643: the Duke of Beaufort and Marie de Rohan.[12] She left the direction of the daily administration of policy to Cardinal Mazarin.
+The best example of Anne's statesmanship and the partial change in her heart towards her native Spain is seen in her keeping of one of Richelieu's men, the Chancellor of France Pierre Séguier, in his post. Séguier was the person who had interrogated Anne in 1637, treating her like a "common criminal" as she described her treatment following the discovery that she was giving military secrets and information to Spain. Anne was virtually under house arrest for a number of years during her husband's rule. By keeping him in his post, Anne was giving a sign that the interests of France and her son Louis were the guiding spirit of all her political and legal actions. Though not necessarily opposed to Spain, she sought to end the war with a French victory, in order to establish a lasting peace between the Catholic nations.
+The Queen also gave a partial Catholic orientation to French foreign policy. This was felt by the Netherlands, France's Protestant ally, which negotiated a separate peace with Spain in 1648.[13]
+In 1648, Anne and Mazarin successfully negotiated the Peace of Westphalia, which ended the Thirty Years' War. Its terms ensured Dutch independence from Spain, awarded some autonomy to the various German princes of the Holy Roman Empire, and granted Sweden seats on the Imperial Diet and territories to control the mouths of the Oder, Elbe, and Weser rivers. France, however, profited most from the settlement. Austria, ruled by the Habsburg Emperor Ferdinand III, ceded all Habsburg lands and claims in Alsace to France and acknowledged her de facto sovereignty over the Three Bishoprics of Metz, Verdun, and Toul. Moreover, eager to emancipate themselves from Habsburg domination, petty German states sought French protection. This anticipated the formation of the 1658 League of the Rhine, leading to the further diminution of Imperial power.
+As the Thirty Years' War came to an end, a civil war known as the Fronde (after the slings used to smash windows) erupted in France. It effectively checked France's ability to exploit the Peace of Westphalia. Anne and Mazarin had largely pursued the policies of Cardinal Richelieu, augmenting the Crown's power at the expense of the nobility and the Parlements. Anne interfered much more in internal policy than foreign affairs; she was a very proud queen who insisted on the divine rights of the King of France.[citation needed]
+All this led her to advocate a forceful policy in all matters relating to the King's authority, in a manner that was much more radical than the one proposed by Mazarin. The Cardinal depended totally on Anne's support and had to use all his influence on the Queen to avoid nullifying, but to restrain some of her radical actions. Anne imprisoned any aristocrat or member of parliament who challenged her will; her main aim was to transfer to her son an absolute authority in the matters of finance and justice. One of the leaders of the Parlement of Paris, whom she had jailed, died in prison.[14]
+The Frondeurs, political heirs of the disaffected feudal aristocracy, sought to protect their traditional feudal privileges from the increasingly centralized royal government. Furthermore, they believed their traditional influence and authority was being usurped by the recently ennobled bureaucrats (the Noblesse de Robe, or "nobility of the robe"), who administered the kingdom and on whom the monarchy increasingly began to rely. This belief intensified the nobles' resentment.[citation needed]
+In 1648, Anne and Mazarin attempted to tax members of the Parlement de Paris. The members refused to comply and ordered all of the king's earlier financial edicts burned. Buoyed by the victory of Louis, duc d’Enghien (later known as le Grand Condé) at the Battle of Lens, Mazarin, on Queen Anne's insistence, arrested certain members in a show of force.[15] The most important arrest, from Anne's point of view, concerned Pierre Broussel, one of the most important leaders in the Parlement de Paris.
+People in France were complaining about the expansion of royal authority, the high rate of taxation, and the reduction of the authority of the Parlement de Paris and other regional representative entities. Paris erupted in rioting as a result, and Anne was forced, under intense pressure, to free Broussel. Moreover, a mob of angry Parisians broke into the royal palace and demanded to see their king. Led into the royal bedchamber, they gazed upon Louis, who was feigning sleep, were appeased, and then quietly departed. The threat to the royal family prompted Anne to flee Paris with the king and his courtiers.
+Shortly thereafter, the conclusion of the Peace of Westphalia allowed Condé's army to return to aid Louis and his court. Condé's family was close to Anne at that time, and he agreed to help her attempt to restore the king's authority.[16] The queen's army, headed by Condé, attacked the rebels in Paris; the rebels were under the political control of Anne's old friend Marie de Rohan. Beaufort, who had escaped from the prison where Anne had incarcerated him five years before, was the military leader in Paris, under the nominal control of Conti. After a few battles, a political compromise was reached; the Peace of Rueil was signed, and the court returned to Paris.
+Unfortunately for Anne, her partial victory depended on Condé, who wanted to control the queen and destroy Mazarin's influence. It was Condé's sister who pushed him to turn against the queen. After striking a deal with her old friend Marie de Rohan, who was able to impose the nomination of Charles de l'Aubespine, marquis de Châteauneuf as minister of justice, Anne arrested Condé, his brother Armand de Bourbon, Prince of Conti, and the husband of their sister Anne Genevieve de Bourbon, duchess of Longueville. This situation did not last long, and Mazarin's unpopularity led to the creation of a coalition headed mainly by Marie de Rohan and the duchess of Longueville. This aristocratic coalition was strong enough to liberate the princes, exile Mazarin, and impose a condition of virtual house arrest on Queen Anne.
+All these events were witnessed by Louis and largely explained his later distrust of Paris and the higher aristocracy.[17] "In one sense, Louis' childhood came to an end with the outbreak of the Fronde. It was not only that life became insecure and unpleasant – a fate meted out to many children in all ages – but that Louis had to be taken into the confidence of his mother and Mazarin and political and military matters of which he could have no deep understanding".[18] "The family home became at times a near-prison when Paris had to be abandoned, not in carefree outings to other chateaux but in humiliating flights".[18] The royal family was driven out of Paris twice in this manner, and at one point Louis XIV and Anne were held under virtual arrest in the royal palace in Paris. The Fronde years planted in Louis a hatred of Paris and a consequent determination to move out of the ancient capital as soon as possible, never to return.[19]
+Just as the first Fronde (the Fronde parlementaire of 1648–1649) ended, a second one (the Fronde des princes of 1650–1653) began. Unlike that which preceded it, tales of sordid intrigue and half-hearted warfare characterized this second phase of upper-class insurrection. To the aristocracy, this rebellion represented a protest against and a reversal of their political demotion from vassals to courtiers. It was headed by the highest-ranking French nobles, among them Louis' uncle Gaston, Duke of Orléans and first cousin Anne Marie Louise d'Orléans, Duchess of Montpensier, known as la Grande Mademoiselle; Princes of the Blood such as Condé, his brother Armand de Bourbon, Prince of Conti, and their sister the Duchess of Longueville; dukes of legitimised royal descent, such as Henri, Duke of Longueville, and François, Duke of Beaufort; so-called "foreign princes" such as Frédéric Maurice, Duke of Bouillon, his brother Marshal Turenne, and Marie de Rohan, Duchess of Chevreuse; and scions of France's oldest families, such as François de La Rochefoucauld.
+Queen Anne played the most important role in defeating the Fronde because she wanted to transfer absolute authority to her son. In addition, most of the princes refused to deal with Mazarin, who went into exile for a number of years. The Frondeurs claimed to act on Louis' behalf, and in his real interest against his mother and Mazarin.
+Queen Anne had a very close relationship with the Cardinal, and many observers believed that Mazarin became Louis XIV's stepfather by a secret marriage to Queen Anne.[20] However, Louis' coming-of-age and subsequent coronation deprived them of the Frondeurs' pretext for revolt. The Fronde thus gradually lost steam and ended in 1653, when Mazarin returned triumphantly from exile. From that time until his death, Mazarin was in charge of foreign and financial policy without the daily supervision of Anne, who was no longer regent.[21]
+During this period, Louis fell in love with Mazarin's niece Marie Mancini, but Anne and Mazarin ended the king's infatuation by sending Mancini away from court to be married in Italy. While Mazarin might have been tempted for a short period of time to marry his niece to the King of France, Queen Anne was absolutely against this; she wanted to marry her son to the daughter of her brother, Philip IV of Spain, for both dynastic and political reasons. Mazarin soon supported the Queen's position because he knew that her support for his power and his foreign policy depended on making peace with Spain from a strong position and on the Spanish marriage. Additionally, Mazarin's relations with Marie Mancini were not good, and he did not trust her to support his position. All of Louis' tears and his supplications to his mother did not make her change her mind; the Spanish marriage was very important both for its role in ending the war between France and Spain, and because many of the claims and objectives of Louis' foreign policy in the next 50 years would be based on this marriage.[22]
+Louis XIV was declared to have reached the age of majority on 7 September 1651. On the death of Mazarin, in March 1661, Louis assumed personal control of the reins of government and astonished his court by declaring that he would rule without a chief minister: "Up to this moment I have been pleased to entrust the government of my affairs to the late Cardinal. It is now time that I govern them myself. You [he was talking to the secretaries and ministers of state] will assist me with your counsels when I ask for them. I request and order you to seal no orders except by my command . . . I order you not to sign anything, not even a passport . . . without my command; to render account to me personally each day and to favor no one".[23] Louis was able to capitalize on the widespread public yearning for law and order, that resulted from prolonged foreign wars and domestic civil strife, to further consolidate central political authority and reform at the expense of the feudal aristocracy. Praising his ability to choose and encourage men of talent, the historian Chateaubriand noted: "it is the voice of genius of all kinds which sounds from the tomb of Louis".[24]
+Louis began his personal reign with administrative and fiscal reforms. In 1661, the treasury verged on bankruptcy. To rectify the situation, Louis chose Jean-Baptiste Colbert as Controller-General of Finances in 1665. However, Louis first had to neutralize Nicolas Fouquet, the Superintendent of Finances, in order to give Colbert a free hand. Although Fouquet's financial indiscretions were not very different from Mazarin's before him or Colbert's after him, his ambition was worrying to Louis. He had, for example, built an opulent château at Vaux-le-Vicomte where he entertained Louis and his court ostentatiously, as if he were wealthier than the king himself. The court was left with the impression that the vast sums of money needed to support his lifestyle could only have been obtained through embezzlement of government funds.
+Fouquet appeared eager to succeed Mazarin and Richelieu in assuming power, and he indiscreetly purchased and privately fortified the remote island of Belle Île. These acts sealed his doom. Fouquet was charged with embezzlement. The Parlement found him guilty and sentenced him to exile. However, Louis altered the sentence to life-imprisonment and abolished Fouquet's post.
+With Fouquet dismissed, Colbert reduced the national debt through more efficient taxation. The principal taxes included the aides and douanes (both customs duties), the gabelle (a tax on salt), and the taille (a tax on land). The taille was reduced at first; financial officials were forced to keep regular accounts, auctioning certain taxes instead of selling them privately to a favored few, revising inventories and removing unauthorized exemptions (for example, in 1661 only 10 per cent from the royal domain reached the King). Reform proved difficult because the taille was levied by officers of the Crown who had purchased their post at a high price: punishment of abuses necessarily lowered the value of the post. Nevertheless, excellent results were achieved: the deficit of 1661 turned into a surplus in 1666. The interest on the debt was reduced from 52 million to 24 million livres. The taille was reduced to 42 million in 1661 and 35 million in 1665; finally the revenue from indirect taxation progressed from 26 million to 55 million. The revenues of the royal domain were raised from 80,000 livres in 1661 to 5.5 million livres in 1671. In 1661, the receipts were equivalent to 26 million British pounds, of which 10 million reached the treasury. The expenditure was around 18 million pounds, leaving a deficit of 8 million. In 1667, the net receipts had risen to 20 million pounds sterling, while expenditure had fallen to 11 million, leaving a surplus of 9 million pounds.
+To support the reorganized and enlarged army, the panoply of Versailles, and the growing civil administration, the king needed a good deal of money. Finance had always been the weak spot in the French monarchy: methods of collecting taxes were costly and inefficient; direct taxes passed through the hands of many intermediate officials; and indirect taxes were collected by private concessionaries, called tax farmers, who made a substantial profit. Consequently, the state always received far less than what the taxpayers actually paid.
+The main weakness arose from an old bargain between the French crown and nobility: the king might raise taxes without consent if only he refrained from taxing the nobles. Only the "unprivileged" classes paid direct taxes, and this term came to mean the peasants only, since many bourgeois, in one way or another, obtained exemptions.
+The system was outrageously unjust in throwing a heavy tax burden on the poor and helpless. Later, after 1700, the French ministers who were supported by Louis' secret wife Madame De Maintenon, were able to convince the king to change his fiscal policy. Louis was willing enough to tax the nobles but was unwilling to fall under their control, and only towards the close of his reign, under extreme stress of war, was he able, for the first time in French history, to impose direct taxes on the aristocratic elements of the population. This was a step toward equality before the law and toward sound public finance, but so many concessions and exemptions were won by nobles and bourgeois that the reform lost much of its value.[25]
+Louis and Colbert also had wide-ranging plans to bolster French commerce and trade. Colbert's mercantilist administration established new industries and encouraged manufacturers and inventors, such as the Lyon silk manufacturers and the Gobelins manufactory, a producer of tapestries. He invited manufacturers and artisans from all over Europe to France, such as Murano glassmakers, Swedish ironworkers, and Dutch shipbuilders. In this way, he aimed to decrease foreign imports while increasing French exports, hence reducing the net outflow of precious metals from France.
+Louis instituted reforms in military administration through Michel le Tellier and the latter's son François-Michel le Tellier, Marquis de Louvois. They helped to curb the independent spirit of the nobility, imposing order on them at court and in the army. Gone were the days when generals protracted war at the frontiers while bickering over precedence and ignoring orders from the capital and the larger politico-diplomatic picture. The old military aristocracy (the Noblesse d'épée, or "nobility of the sword") ceased to have a monopoly over senior military positions and rank. Louvois, in particular, pledged to modernize the army and re-organize it into a professional, disciplined, well-trained force. He was devoted to the soldiers' material well-being and morale, and even tried to direct campaigns.
+Legal matters did not escape Louis' attention, as is reflected in the numerous "Great Ordinances" he enacted. Pre-revolutionary France was a patchwork of legal systems, with as many legal customs as there were provinces, and two co-existing legal traditions—customary law in the north and Roman civil law in the south.[26] The Grande Ordonnance de Procédure Civile of 1667, also known as the Code Louis, was a comprehensive legal code attempting a uniform regulation of civil procedure throughout legally irregular France. Among other things, it prescribed baptismal, marriage and death records in the state's registers, not the church's, and it strictly regulated the right of the Parlements to remonstrate.[27] The Code Louis played an important part in French legal history as the basis for the Napoleonic code, from which many modern legal codes are, in turn, derived.
+One of Louis' more infamous decrees was the Grande Ordonnance sur les Colonies of 1685, also known as the Code Noir ("black code"). Although it sanctioned slavery, it attempted to humanise the practice by prohibiting the separation of families. Additionally, in the colonies, only Roman Catholics could own slaves, and these had to be baptised.
+Louis ruled through a number of councils:
+The death of his maternal uncle King Philip IV of Spain, in 1665, precipitated the War of Devolution. In 1660, Louis had married Philip IV's eldest daughter, Maria Theresa, as one of the provisions of the 1659 Treaty of the Pyrenees.[29] The marriage treaty specified that Maria Theresa was to renounce all claims to Spanish territory for herself and all her descendants.[29] Mazarin and Lionne, however, made the renunciation conditional on the full payment of a Spanish dowry of 500,000 écus.[30] The dowry was never paid and would later play a part persuading his maternal first cousin Charles II of Spain to leave his empire to Philip, Duke of Anjou (later Philip V of Spain), the grandson of Louis XIV and Maria Theresa.
+The War of Devolution did not focus on the payment of the dowry; rather, the lack of payment was what Louis XIV used as a pretext for nullifying Maria Theresa's renunciation of her claims, allowing the land to "devolve" to him. In Brabant (the location of the land in dispute), children of first marriages traditionally were not disadvantaged by their parents' remarriages and still inherited property. Louis' wife was Philip IV's daughter by his first marriage, while the new king of Spain, Charles II, was his son by a subsequent marriage. Thus, Brabant allegedly "devolved" to Maria Theresa, giving France a justification to attack the Spanish Netherlands.
+During the Eighty Years' War with Spain, France supported the Dutch Republic as part of a general policy of opposing Habsburg power. Johan de Witt, Dutch Grand Pensionary from 1653 to 1672, viewed them as crucial for Dutch security and against his domestic Orangist opponents. Louis provided support in the 1665-1667 Second Anglo-Dutch War but used the opportunity to launch the War of Devolution in 1667. This captured Franche-Comté and much of the Spanish Netherlands; French expansion in this area was a direct threat to Dutch economic interests.[31]
+The Dutch opened talks with Charles II of England on a common diplomatic front against France, leading to the Triple Alliance, between England, the Dutch and Sweden. The threat of an escalation and a secret treaty to divide Spanish possessions with Emperor Leopold, the other major claimant to the throne of Spain, led Louis to relinquish many of his gains in the 1668 Treaty of Aix-la-Chapelle.[32]
+Louis placed little reliance on his agreement with Leopold and as it was now clear French and Dutch aims were in direct conflict, he decided to first defeat the Republic, then seize the Spanish Netherlands. This required breaking up the Triple Alliance; he paid Sweden to remain neutral and signed the 1670 Secret Treaty of Dover with Charles, an Anglo-French alliance against the Dutch Republic. In May 1672, France invaded the Republic, supported by Münster and the Electorate of Cologne.[33]
+Rapid French advance led to a coup that toppled De Witt and brought William III to power. Leopold viewed French expansion into the Rhineland as an increasing threat, especially after their seizure of the strategic Duchy of Lorraine in 1670. The prospect of Dutch defeat led Leopold to an alliance with Brandenburg-Prussia on 23 June, followed by another with the Republic on 25th.[34] Although Brandenburg was forced out of the war by the June 1673 Treaty of Vossem, in August an anti-French alliance was formed by the Dutch, Spain, Emperor Leopold and the Duke of Lorraine.[35]
+The French alliance was deeply unpopular in England, who made peace with the Dutch in the February 1674 Treaty of Westminster. However, French armies held significant advantages over their opponents; an undivided command, talented generals like Turenne, Condé and Luxembourg and vastly superior logistics. Reforms introduced by Louvois, the Secretary of War, helped maintain large field armies that could be mobilised much quicker, allowing them to mount offensives in early spring before their opponents were ready.[36]
+The French were forced to retreat from the Dutch Republic but these advantages allowed them to hold their ground in Alsace and the Spanish Netherlands, while retaking Franche-Comté. By 1678, mutual exhaustion led to the Treaty of Nijmegen, which was generally settled in France's favour and allowed Louis to intervene in the Scanian War. Despite military defeat, his ally Sweden regained much of their losses under the 1679 treaties of Saint-Germain-en-Laye, Fontainebleau and Lund imposed on Denmark-Norway and Brandenburg.[37]
+Louis was at the height of his power, but at the cost of uniting his opponents; this increased as he continued his expansion. In 1679, he dismissed his foreign minister Simon Arnauld, marquis de Pomponne, because he was seen as having compromised too much with the allies. Louis maintained the strength of his army, but in his next series of territorial claims avoided using military force alone. Rather, he combined it with legal pretexts in his efforts to augment the boundaries of his kingdom. Contemporary treaties were intentionally phrased ambiguously. Louis established the Chambers of Reunion to determine the full extent of his rights and obligations under those treaties.
+Cities and territories, such as Luxembourg and Casale, were prized for their strategic positions on the frontier and access to important waterways. Louis also sought Strasbourg, an important strategic crossing on the left bank of the Rhine and theretofore a Free Imperial City of the Holy Roman Empire, annexing it and other territories in 1681. Although a part of Alsace, Strasbourg was not part of Habsburg-ruled Alsace and was thus not ceded to France in the Peace of Westphalia.
+Following these annexations, Spain declared war, precipitating the War of the Reunions. However, the Spanish were rapidly defeated because the Emperor (distracted by the Great Turkish War) abandoned them, and the Dutch only supported them minimally. By the Truce of Ratisbon, in 1684, Spain was forced to acquiesce in the French occupation of most of the conquered territories, for 20 years.[38]
+Louis' policy of the Réunions may have raised France to its greatest size and power during his reign, but it alienated much of Europe. This poor public opinion was compounded by French actions off the Barbary Coast and at Genoa. First, Louis had Algiers and Tripoli, two Barbary pirate strongholds, bombarded to obtain a favourable treaty and the liberation of Christian slaves. Next, in 1684, a punitive mission was launched against Genoa in retaliation for its support for Spain in previous wars. Although the Genoese submitted, and the Doge led an official mission of apology to Versailles, France gained a reputation for brutality and arrogance. European apprehension at growing French might and the realisation of the extent of the dragonnades' effect (discussed below) led many states to abandon their alliance with France.[39] Accordingly, by the late 1680s, France became increasingly isolated in Europe.
+French colonies multiplied in Africa, the Americas, and Asia during Louis' reign, and French explorers made important discoveries in North America.  In 1673, Louis Jolliet and Jacques Marquette discovered the Mississippi River. In 1682, René-Robert Cavelier, Sieur de La Salle, followed the Mississippi to the Gulf of Mexico and claimed the vast Mississippi basin in Louis' name, calling it Louisiane. French trading posts were also established in India, at Chandernagore and Pondicherry, and in the Indian Ocean at Île Bourbon. Throughout these regions Louis and Colbert embarked on an extensive program of architecture and urbanism meant to reflect the styles of Versailles and Paris and the 'gloire' of the realm.[40]
+Meanwhile, diplomatic relations were initiated with distant countries. In 1669, Suleiman Aga led an Ottoman embassy to revive the old Franco-Ottoman alliance.[41] Then, in 1682, after the reception of the Moroccan embassy of Mohammed Tenim in France, Moulay Ismail, Sultan of Morocco, allowed French consular and commercial establishments in his country.[42] In 1699, Louis once again received a Moroccan ambassador, Abdallah bin Aisha, and in 1715, he received a Persian embassy led by Mohammad Reza Beg.
+From farther afield, Siam dispatched an embassy in 1684, reciprocated by the French magnificently the next year under Alexandre, Chevalier de Chaumont. This, in turn, was succeeded by another Siamese embassy under Kosa Pan, superbly received at Versailles in 1686. Louis then sent another embassy in 1687, under Simon de la Loubère, and French influence grew at the Siamese court, which granted Mergui as a naval base to France. However, the death of Narai, King of Ayutthaya, the execution of his pro-French minister Constantine Phaulkon, and the Siege of Bangkok in 1688 ended this era of French influence.[43]
+France also attempted to participate actively in Jesuit missions to China. To break the Portuguese dominance there, Louis sent Jesuit missionaries to the court of the Kangxi Emperor in 1685: Jean de Fontaney, Joachim Bouvet, Jean-François Gerbillon, Louis Le Comte, and Claude de Visdelou.[44] Louis also received a Chinese Jesuit, Michael Shen Fu-Tsung, at Versailles in 1684.[45] Furthermore, Louis' librarian and translator Arcadio Huang was Chinese.[46][47]
+By the early 1680s, Louis had greatly augmented French influence in the world. Domestically, he successfully increased the influence of the crown and its authority over the church and aristocracy, thus consolidating absolute monarchy in France.
+Louis initially supported traditional Gallicanism, which limited papal authority in France, and convened an Assembly of the French clergy in November 1681. Before its dissolution eight months later, the Assembly had accepted the Declaration of the Clergy of France, which increased royal authority at the expense of papal power. Without royal approval, bishops could not leave France, and appeals could not be made to the Pope. Additionally, government officials could not be excommunicated for acts committed in pursuance of their duties. Although the king could not make ecclesiastical law, all papal regulations without royal assent were invalid in France. Unsurprisingly, the pope repudiated the Declaration.[3]
+By attaching nobles to his court at Versailles, Louis achieved increased control over the French aristocracy. Apartments were built to house those willing to pay court to the king.[48] However, the pensions and privileges necessary to live in a style appropriate to their rank were only possible by waiting constantly on Louis.[49] For this purpose, an elaborate court ritual was created wherein the king became the centre of attention and was observed throughout the day by the public. With his excellent memory, Louis could then see who attended him at court and who was absent, facilitating the subsequent distribution of favours and positions. Another tool Louis used to control his nobility was censorship, which often involved the opening of letters to discern their author's opinion of the government and king.[48] Moreover, by entertaining, impressing, and domesticating them with extravagant luxury and other distractions, Louis not only cultivated public opinion of him, he also ensured the aristocracy remained under his scrutiny.
+Louis' extravagance at Versailles extended far beyond the scope of elaborate court rituals. Louis took delivery of an African elephant as a gift from the king of Portugal.[50]}} The king encouraged the leading nobles to live at Versailles. This, along with the prohibition of private armies, prevented them from passing time on their own estates and in their regional power bases, from which they historically waged local wars and plotted resistance to royal authority.  Louis thus compelled and seduced the old military aristocracy (the "nobility of the sword") into becoming his ceremonial courtiers, further weakening their power. In their place, Louis raised commoners or the more recently ennobled bureaucratic aristocracy (the "nobility of the robe"). He judged that royal authority thrived more surely by filling high executive and administrative positions with these men because they could be more easily dismissed than nobles of ancient lineage, with entrenched influence. It is believed that Louis' policies were rooted in his experiences during the Fronde, when men of high birth readily took up the rebel cause against their king, who was actually the kinsman of some. This victory of Louis' over the nobility may have then in fact ensured the end of major civil wars in France until the French Revolution about a century later.
+In 1648 France was the leading European power, and most of the wars pivoted around its aggressiveness.  Only poverty-stricken Russia exceeded it in population, and no one could match its wealth, central location, and very strong professional army. It had largely avoided the devastation of the Thirty Years' War.  Its weaknesses included an inefficient financial system that was hard-pressed to pay for all the military adventures, and the tendency of most other powers to gang up against it.
+During the very long reign of King Louis XIV (1643 – 1715), France fought three major wars: the Franco-Dutch War, the War of the League of Augsburg, and the War of the Spanish Succession. There were also two lesser conflicts: the War of Devolution and the War of the Reunions.[51] The wars were very expensive but they defined Louis XIV's foreign policies, and his personality shaped his approach. Impelled "by a mix of commerce, revenge, and pique," Louis sensed that warfare was the ideal way to enhance his glory. In peacetime he concentrated on preparing for the next war. He taught his diplomats that their job was to create tactical and strategic advantages for the French military.[4] By 1695, France retained much of its dominance, but had lost control of the seas to the combination of England and Holland.  What's more, most countries, both Protestant and Catholic, were in alliance against it. Vauban, France's leading military strategist, warned the king in 1689 that a hostile "Alliance" was too powerful at sea. He recommended the best way for France to fight back was to license French merchants ships to privateer and seize enemy merchant ships, while avoiding its navies:
+Vauban was pessimistic about France's so-called friends and allies:
+Louis decided to persecute Protestants and revoke the 1598 Edict of Nantes, which awarded Huguenots political and religious freedom. He saw the persistence of Protestantism as a disgraceful reminder of royal powerlessness. After all, the Edict was the pragmatic concession of his grandfather Henry IV to end the longstanding French Wars of Religion. An additional factor in Louis' thinking was the prevailing contemporary European principle to assure socio-political stability, cuius regio, eius religio ("whose realm, his religion"), the idea that the religion of the ruler should be the religion of the realm (as originally confirmed in central Europe in the Peace of Augsburg of 1555).[54]
+Responding to petitions, Louis initially excluded Protestants from office, constrained the meeting of synods, closed churches outside of Edict-stipulated areas, banned Protestant outdoor preachers, and prohibited domestic Protestant migration. He also disallowed Protestant-Catholic intermarriages to which third parties objected, encouraged missions to the Protestants, and rewarded converts to Catholicism.[55] This discrimination did not encounter much Protestant resistance, and a steady conversion of Protestants occurred, especially among the noble elites.
+In 1681, Louis dramatically increased his persecution of Protestants. The principle of cuius regio, eius religio generally had also meant that subjects who refused to convert could emigrate, but Louis banned emigration and effectively insisted that all Protestants must be converted. Secondly, following the proposal of René de Marillac and the Marquis of Louvois, he began quartering dragoons in Protestant homes. Although this was within his legal rights, the dragonnades inflicted severe financial strain on Protestants and atrocious abuse. Between 300,000 and 400,000 Huguenots converted, as this entailed financial rewards and exemption from the dragonnades.[56]
+On 15 October 1685, Louis issued the Edict of Fontainebleau, which cited the redundancy of privileges for Protestants given their scarcity after the extensive conversions. The Edict of Fontainebleau revoked the Edict of Nantes and repealed all the privileges that arose therefrom.[3] By his edict, Louis no longer tolerated the existence of Protestant groups, pastors, or churches in France. No further churches were to be constructed, and those already existing were to be demolished. Pastors could choose either exile or a secular life. Those Protestants who had resisted conversion were now to be baptised forcibly into the established church.[57]
+Historians have debated Louis' reasons for issuing the Edict of Fontainebleau. He may have been seeking to placate Pope Innocent XI, with whom relations were tense and whose aid was necessary to determine the outcome of a succession crisis in the Electorate of Cologne. He may also have acted to upstage Emperor Leopold I and regain international prestige after the latter defeated the Turks without Louis' help. Otherwise, he may simply have desired to end the remaining divisions in French society dating to the Wars of Religion by fulfilling his coronation oath to eradicate heresy.[58][59]
+Many historians have condemned the Edict of Fontainebleau as gravely harmful to France.[60] In support, they cite the emigration of about 200,000 highly skilled Huguenots (roughly one-fourth of the Protestant population, or 1% of the French population) who defied royal decrees and fled France for various Protestant states, weakening the French economy and enriching that of Protestant states.
+On the other hand, there are historians who view this as an exaggeration. They argue that most of France's preeminent Protestant businessmen and industrialists converted to Catholicism and remained.[61]
+What is certain is that reaction to the Edict was mixed. Even while French Catholic leaders exulted, Pope Innocent XI still argued with Louis over Gallicanism and criticised the use of violence. Protestants across Europe were horrified at the treatment of their co-religionists, but most Catholics in France applauded the move. Nonetheless, it is indisputable that Louis' public image in most of Europe, especially in Protestant regions, was dealt a severe blow.
+In the end, however, despite renewed tensions with the Camisards of south-central France at the end of his reign, Louis may have helped ensure that his successor would experience fewer instances of the religion-based disturbances that had plagued his forebears. French society would sufficiently change by the time of his descendant, Louis XVI, to welcome tolerance in the form of the 1787 Edict of Versailles, also known as the Edict of Tolerance. This restored to non-Catholics their civil rights and the freedom to worship openly.[62] With the advent of the French Revolution in 1789, Protestants were granted equal rights with their Roman Catholic counterparts.
+The War of the League of Augsburg, which lasted from 1688 to 1697, initiated a period of decline in Louis' political and diplomatic fortunes. The conflict arose from two events in the Rhineland. First, in 1685, the Elector Palatine Charles II died. All that remained of his immediate family was Louis' sister-in-law, Elizabeth Charlotte. German law ostensibly barred her from succeeding to her brother's lands and electoral dignity, but it was unclear enough for arguments in favour of Elizabeth Charlotte to have a chance of success. Conversely, the princess was clearly entitled to a division of the family's personal property. Louis pressed her claims to land and chattels, hoping the latter, at least, would be given to her.[63] Then, in 1688, Maximilian Henry of Bavaria, Archbishop of Cologne, an ally of France, died. The archbishopric had traditionally been held by the Wittelsbachs of Bavaria. However, the Bavarian claimant to replace Maximilian Henry, Prince Joseph Clemens of Bavaria, was at that time not more than 17 years old and not even ordained. Louis sought instead to install his own candidate, William Egon of Fürstenberg, to ensure the key Rhenish state remained an ally.[64]
+In light of his foreign and domestic policies during the early 1680s, which were perceived as aggressive, Louis' actions, fostered by the succession crises of the late 1680s, created concern and alarm in much of Europe. This led to the formation of the 1686 League of Augsburg by the Holy Roman Emperor, Spain, Sweden, Saxony, and Bavaria. Their stated intention was to return France to at least the borders agreed to in the Treaty of Nijmegen.[65] Emperor Leopold I's persistent refusal to convert the Truce of Ratisbon into a permanent treaty fed Louis' fears that the Emperor would turn on France and attack the Reunions after settling his affairs in the Balkans.[66]
+Another event that Louis found threatening was the Glorious Revolution of 1688, in England. Although King James II was Catholic, his two Anglican daughters, Mary and Anne, ensured the English people a Protestant succession. However, when James II's son James was born, he took precedence in the succession over his elder sisters. This seemed to herald an era of Catholic monarchs in England. Protestant lords called on the Dutch Prince William III of Orange, grandson of Charles I of England, to come to their aid. He sailed for England with troops despite Louis' warning that France would regard it as a provocation. Witnessing numerous desertions and defections, even among those closest to him, James II fled England. Parliament declared the throne vacant, and offered it to James's daughter Mary II and his son-in-law and nephew William. Vehemently anti-French, William (now William III of England) pushed his new kingdoms into war, thus transforming the League of Augsburg into the Grand Alliance. Before this happened, Louis expected William's expedition to England to absorb his energies and those of his allies, so he dispatched troops to the Rhineland after the expiry of his ultimatum to the German princes requiring confirmation of the Truce of Ratisbon and acceptance of his demands about the succession crises. This military manoeuvre was also intended to protect his eastern provinces from Imperial invasion by depriving the enemy army of sustenance, thus explaining the pre-emptive scorched earth policy pursued in much of southwestern Germany (the "Devastation of the Palatinate").[67]
+French armies were generally victorious throughout the war because of Imperial commitments in the Balkans, French logistical superiority, and the quality of French generals such as Condé's famous pupil, François Henri de Montmorency-Bouteville, duc de Luxembourg. His triumphs at the Battles of Fleurus in 1690, Steenkerque in 1692, and Landen in 1693 preserved northern France from invasion.[68]
+Although an attempt to restore James II failed at the Battle of the Boyne in 1690, France accumulated a string of victories from Flanders in the north, Germany in the east, and Italy and Spain in the south, to the high seas and the colonies. Louis personally supervised the captures of Mons in 1691 and Namur in 1692. Luxembourg gave France the defensive line of the Sambre by capturing Charleroi in 1693. France also overran most of the Duchy of Savoy after the battles of Marsaglia and Staffarde in 1693. While naval stalemate ensued after the French victory at the Battle of Beachy Head in 1690 and the Allied victory at Barfleur-La Hougue in 1692, the Battle of Torroella in 1694 exposed Catalonia to French invasion, culminating in the capture of Barcelona. Although the Dutch captured Pondichéry in 1693, a French raid on the Spanish treasure port of Cartagena, Spain in 1697 yielded a fortune of 10,000,000 livres.
+In July 1695, the city of Namur, occupied for three years by the French, was besieged by an allied army led by William III. Louis XIV ordered the surprise destruction of a Flemish city to divert the attention of these troops. This led to the bombardment of Brussels, in which 4-5000 buildings were destroyed, including the entire city-center. The strategy failed, as Namur fell three weeks later, but harmed Louis XIV's reputation: a century later, Napoleon deemed the bombardment "as barbarous as it was useless."[69]
+Peace was broached by Sweden in 1690. By 1692, both sides evidently wanted peace, and secret bilateral talks began, but to no avail.[70] Louis tried to break up the alliance against him by dealing with individual opponents, but this did not achieve its aim until 1696, when the Savoyards agreed to the Treaty of Turin and switched sides. Thereafter, members of the League of Augsburg rushed to the peace table, and negotiations for a general peace began in earnest, culminating in the Treaty of Ryswick of 1697.[71]
+The Treaty of Ryswick ended the War of the League of Augsburg and disbanded the Grand Alliance. By manipulating their rivalries and suspicions, Louis divided his enemies and broke their power.
+The treaty yielded many benefits for France. Louis secured permanent French sovereignty over all of Alsace, including Strasbourg, and established the Rhine as the Franco-German border (which persists to this day). Pondichéry and Acadia were returned to France, and Louis' de facto possession of Saint-Domingue was recognised as lawful. However, he returned Catalonia and most of the Reunions.
+French military superiority might have allowed him to press for more advantageous terms. Thus, his generosity to Spain with regard to Catalonia has been read as a concession to foster pro-French sentiment and may ultimately have induced King Charles II to name Louis' grandson Philip, Duke of Anjou, as heir to the throne of Spain.[72] In exchange for financial compensation, France renounced its interests in the Electorate of Cologne and the Palatinate. Lorraine, which had been occupied by the French since 1670, was returned to its rightful Duke Leopold, albeit with a right of way to the French military. William and Mary were recognised as joint sovereigns of the British Isles, and Louis withdrew support for James II. The Dutch were given the right to garrison forts in the Spanish Netherlands that acted as a protective barrier against possible French aggression. Though in some respects, the Treaty of Ryswick may appear a diplomatic defeat for Louis since he failed to place client rulers in control of the Palatinate or the Electorate of Cologne, he did in fact fulfill many of the aims laid down in his 1688 ultimatum.[73] In any case, peace in 1697 was desirable to Louis, since France was exhausted from the costs of the war.
+By the time of the Treaty of Ryswick, the Spanish succession had been a source of concern to European leaders for well over forty years. King Charles II ruled a vast empire comprising Spain, Naples, Sicily, Milan, the Spanish Netherlands, and numerous Spanish colonies. He produced no children, however, and consequently had no direct heirs.
+The principal claimants to the throne of Spain belonged to the ruling families of France and Austria. The French claim derived from Louis XIV's mother Anne of Austria (the older sister of Philip IV of Spain) and his wife Maria Theresa (Philip IV's eldest daughter). Based on the laws of primogeniture, France had the better claim as it originated from the eldest daughters in two generations. However, their renunciation of succession rights complicated matters. In the case of Maria Theresa, nonetheless, the renunciation was considered null and void owing to Spain's breach of her marriage contract with Louis. In contrast, no renunciations tainted the claims of the Emperor Leopold I's son Charles, Archduke of Austria, who was a grandson of Philip III's youngest daughter Maria Anna. The English and Dutch feared that a French or Austrian-born Spanish king would threaten the balance of power and thus preferred the Bavarian Prince Joseph Ferdinand, a grandson of Leopold I through his first wife Margaret Theresa of Spain (the younger daughter of Philip IV).
+In an attempt to avoid war, Louis signed the Treaty of the Hague with William III of England in 1698. This agreement divided Spain's Italian territories between Louis's son le Grand Dauphin and the Archduke Charles, with the rest of the empire awarded to Joseph Ferdinand. William III consented to permitting the Dauphin's new territories to become part of France when the latter succeeded to his father's throne.[74] The signatories, however, omitted to consult the ruler of these lands, and Charles II was passionately opposed to the dismemberment of his empire. In 1699, he re-confirmed his 1693 will that named Joseph Ferdinand as his sole successor.[75]
+Six months later, Joseph Ferdinand died. Therefore, in 1700, Louis and William III concluded a fresh partitioning agreement, the Treaty of London. This allocated Spain, the Low Countries, and the Spanish colonies to the Archduke. The Dauphin would receive all of Spain's Italian territories.[76] Charles II acknowledged that his empire could only remain undivided by bequeathing it entirely to a Frenchman or an Austrian. Under pressure from his German wife, Maria Anna of Neuburg, Charles II named the Archduke Charles as his sole heir.
+On his deathbed in 1700, Charles II unexpectedly changed his will. The clear demonstration of French military superiority for many decades before this time, the pro-French faction at the court of Spain, and even Pope Innocent XII convinced him that France was more likely to preserve his empire intact. He thus offered the entire empire to the Dauphin's second son Philip, Duke of Anjou, provided it remained undivided. Anjou was not in the direct line of French succession, thus his accession would not cause a Franco-Spanish union.[76] If Anjou refused, the throne would be offered to his younger brother Charles, Duke of Berry. If the Duke of Berry declined it, it would go to the Archduke Charles, then to the distantly related House of Savoy if Charles declined it.[77]
+Louis was confronted with a difficult choice. He might agree to a partition of the Spanish possessions and avoid a general war, or accept Charles II's will and alienate much of Europe. Initially, Louis may have been inclined to abide by the partition treaties. However, the Dauphin's insistence persuaded Louis otherwise.[78] Moreover, Louis's foreign minister, Jean-Baptiste Colbert, marquis de Torcy, pointed out that war with the Emperor would almost certainly ensue whether Louis accepted the partition treaties or Charles II's will. He emphasised that, should it come to war, William III was unlikely to stand by France since he "made a treaty to avoid war and did not intend to go to war to implement the treaty".[75] Indeed, in the event of a war, it might be preferable to be already in control of the disputed lands. Eventually, therefore, Louis decided to accept Charles II's will. Philip, Duke of Anjou, thus became Philip V, King of Spain.
+Most European rulers accepted Philip as king, though some only reluctantly. Depending on one's views of the war as inevitable or not, Louis acted reasonably or arrogantly.[79] He confirmed that Philip V retained his French rights despite his new Spanish position. Admittedly, he may only have been hypothesising a theoretical eventuality and not attempting a Franco-Spanish union. But his actions were certainly not read as being disinterested. Moreover, Louis sent troops to the Spanish Netherlands to evict Dutch garrisons and secure Dutch recognition of Philip V. In 1701, Philip transferred the asiento (the right to supply slaves to Spanish colonies) to France, alienating English traders. As tensions mounted, Louis decided to acknowledge James Stuart, the son of James II, as king of England on the latter's death, infuriating William III. These actions enraged Britain and the Dutch Republic.[80] With the Holy Roman Emperor and the petty German states, they formed another Grand Alliance and declared war on France in 1702. French diplomacy, however, secured Bavaria, Portugal, and Savoy as Franco-Spanish allies.[81]
+Even before war was officially declared, hostilities began with Imperial aggression in Italy. When finally declared, the War of the Spanish Succession would last almost until Louis's death, at great cost to him and the kingdom of France.
+The war began with French successes, however the joint talents of John Churchill, Duke of Marlborough, and Eugene of Savoy checked these victories and broke the myth of French invincibility. The duo allowed the Palatinate and Austria to occupy Bavaria after their victory at the Battle of Blenheim. Maximilian II Emanuel, Elector of Bavaria, had to flee to the Spanish Netherlands. The impact of this victory won the support of Portugal and Savoy. Later, the Battle of Ramillies delivered the Low Countries up to the Allies, and the Battle of Turin forced Louis to evacuate Italy, leaving it open to Allied forces. Marlborough and Eugene of Savoy met again at the Battle of Oudenarde, which enabled them to mount an invasion of France.
+France established contact with Francis II Rákóczi and promised support if he took up the cause of Hungarian independence.
+Defeats, famine, and mounting debt greatly weakened France. Between 1693 and 1710, over two million people died in two famines, made worse as foraging armies seized food supplies from the villages.[82][83] In his desperation, Louis XIV even ordered a disastrous invasion of the English island of Guernsey in the autumn of 1704 with the aim of raiding their successful harvest. By the winter of 1708–1709, Louis was willing to accept peace at nearly any cost. He agreed that the entire Spanish empire should be surrendered to the Archduke Charles, and he also consented to return to the frontiers of the Peace of Westphalia, giving up all the territories he had acquired over sixty years of his reign. He could not speak for his grandson, however, and could not promise that Philip V would accept these terms. Thus, the Allies demanded that Louis single-handedly attack his own grandson to force these terms on him. If he could not achieve this within the year, the war would resume. Louis could not accept these terms.[84]
+The final phases of the War of the Spanish Succession demonstrated that the Allies could not maintain the Archduke Charles in Spain just as surely as France could not retain the entire Spanish inheritance for King Philip V. The Allies were definitively expelled from central Spain by the Franco-Spanish victories at the Battles of Villaviciosa and Brihuega in 1710. French forces elsewhere remained obdurate despite their defeats. The Allies suffered a Pyrrhic victory at the Battle of Malplaquet with 21,000 casualties, twice that of the French.[85] Eventually, France recovered its military pride with the decisive victory at Denain in 1712.
+French military successes near the end of the war took place against the background of a changed political situation in Austria. In 1705, the Emperor Leopold I died. His elder son and successor, Joseph I, followed him in 1711. His heir was none other than the Archduke Charles, who secured control of all of his brother's Austrian land holdings. If the Spanish empire then fell to him, it would have resurrected a domain as vast as that of Holy Roman Emperor Charles V in the sixteenth century. To the maritime powers of Great Britain and the Dutch Republic, this would have been as undesirable as a Franco-Spanish union.[86]
+As a result of the fresh British perspective on the European balance of power, Anglo-French talks began that culminated in the 1713 Treaty of Utrecht between Louis, Philip V of Spain, Anne, Queen of Great Britain, and the Dutch Republic. In 1714, after losing Landau and Freiburg, the Holy Roman Emperor also made peace with France in the Treaties of Rastatt and Baden.
+In the general settlement, Philip V retained Spain and its colonies, whereas Austria received the Spanish Netherlands and divided Spanish Italy with Savoy. Britain kept Gibraltar and Menorca. Louis agreed to withdraw his support for James Stuart, son of James II and pretender to the throne of Great Britain, and ceded Newfoundland, Rupert's Land, and Acadia in the Americas to Anne. Britain gained most from the Treaty of Utrecht, but the final terms were much more favourable to France than those which were being discussed in peace negotiations in 1709 and 1710.[citation needed] France retained Île-Saint-Jean and Île Royale, and Louis did acquire a few minor European territories, such as the Principality of Orange and the Ubaye Valley, which covered transalpine passes into Italy. Thanks to Louis, his allies the Electors of Bavaria and Cologne were restored to their pre-war status and returned their lands.[87]
+Louis and his wife Maria Theresa of Spain had six children from the marriage contracted for them in 1660. However, only one child, the eldest, survived to adulthood: Louis, le Grand Dauphin, known as Monseigneur. Maria Theresa died in 1683, whereupon Louis remarked that she had never caused him unease on any other occasion.
+Despite evidence of affection early on in their marriage, Louis was never faithful to Maria Theresa. He took a series of mistresses, both official and unofficial. Among the better documented are Louise de La Vallière (with whom he had 5 children; 1661–67), Bonne de Pons d'Heudicourt (1665), Catherine Charlotte de Gramont (1665), Françoise-Athénaïs, Marquise de Montespan (with whom he had 7 children; 1667–80), Anne de Rohan-Chabot (1669–75), Claude de Vin des Œillets (1 child born in 1676), Isabelle de Ludres (1675–78), and Marie Angélique de Scorailles (1679–81), who died at age 19 in childbirth. Through these liaisons, he produced numerous illegitimate children, most of whom he married to members of cadet branches of the royal family.
+Louis proved relatively more faithful to his second wife, Françoise d'Aubigné, Marquise de Maintenon.  He first met her through her work caring for his children by Madame de Montespan, noting the care she gave to his favorite, Louis Auguste, Duke of Maine.[88]  The king was, at first, put off by her strict religious practice, but he warmed to her through her care for his children.[88]
+When he legitimized his children by Madame de Montespan on 20 December 1673, Françoise d'Aubigné became the royal governess at Saint-Germain.[88]  As governess, she was one of very few people permitted to speak to him as an equal, without limits.[88]  It is believed that they were married secretly at Versailles on or around 10 October 1683[89] or January 1684.[90] This marriage, though never announced or publicly discussed, was an open secret and lasted until his death.[91]
+Louis was a pious and devout king who saw himself as the head and protector of the Gallican Church. Louis made his devotions daily regardless of where he was, following the liturgical calendar regularly.[92] Under the influence of his very religious second wife, he became much stronger in the practice of his Catholic faith.[93]  This included the banning of opera and comedy performances during Lent.[93]
+Towards the middle and the end of his reign, the centre for the King's religious observances was usually the Chapelle Royale at Versailles. Ostentation was a distinguishing feature of daily Mass, annual celebrations, such as those of Holy Week, and special ceremonies.[94] Louis established the Paris Foreign Missions Society, but his informal alliance with the Ottoman Empire was criticised for undermining Christendom.[95]
+Louis generously supported the royal court of France and those who worked under him. He brought the Académie Française under his patronage and became its "Protector". He allowed Classical French literature to flourish by protecting such writers as Molière, Racine, and La Fontaine, whose works remain greatly influential to this day. Louis also patronised the visual arts by funding and commissioning various artists, such as Charles Le Brun, Pierre Mignard, Antoine Coysevox, and Hyacinthe Rigaud, whose works became famous throughout Europe. In music, composers and musicians such as Jean-Baptiste Lully, Jacques Champion de Chambonnières, and François Couperin thrived. In 1661, Louis founded the Académie Royale de Danse, and in 1669, the Académie d'Opéra, important driving events in the evolution of ballet. The King also attracted, supported and patronized such artists as André Charles Boulle who revolutionised marquetry with his art of inlay, today known as "Boulle Work".
+Over the course of four building campaigns, Louis converted a hunting lodge built by Louis XIII into the spectacular Palace of Versailles. With the exception of the current Royal Chapel (built near the end of Louis' reign), the palace achieved much of its current appearance after the third building campaign, which was followed by an official move of the royal court to Versailles on 6 May 1682. Versailles became a dazzling, awe-inspiring setting for state affairs and the reception of foreign dignitaries. At Versailles, the king alone commanded attention.
+Several reasons have been suggested for the creation of the extravagant and stately palace, as well as the relocation of the monarchy's seat. For example, the memoirist Saint-Simon speculated that Louis viewed Versailles as an isolated power center where treasonous cabals could be more readily discovered and foiled.[49] Alternatively, there has been speculation that the revolt of the Fronde caused Louis to hate Paris, which he abandoned for a country retreat. However, his sponsorship of many public works in Paris, such as the establishment of a police force and of street-lighting,[96] lend little credence to this theory. As a further example of his continued care for the capital, Louis constructed the Hôtel des Invalides, a military complex and home to this day for officers and soldiers rendered infirm either by injury or old age. While pharmacology was still quite rudimentary in his day, the Invalides pioneered new treatments and set new standards for hospice treatment. The conclusion of the Treaty of Aix-la-Chapelle, in 1668, also induced Louis to demolish the northern walls of Paris in 1670 and replace them with wide tree-lined boulevards.[97]
+Louis also renovated and improved the Louvre and other royal residences. Gian Lorenzo Bernini was originally to plan additions to the Louvre; however, his plans would have meant the destruction of much of the existing structure, replacing it with an Italian summer villa in the centre of Paris. Bernini's plans were eventually shelved in favour of the elegant Louvre Colonnade designed by three Frenchmen: Louis Le Vau, Charles Le Brun, and Claude Perrault. With the relocation of the court to Versailles, the Louvre was given over to the arts and the public.[98]
+During his visit from Rome, Bernini also executed a renowned portrait bust of the king.
+Few rulers in world history have commemorated themselves in as grand a manner as Louis.[99] Louis used court ritual and the arts to validate and augment his control over France. With his support, Colbert established from the beginning of Louis' personal reign a centralised and institutionalised system for creating and perpetuating the royal image. The King was thus portrayed largely in majesty or at war, notably against Spain. This portrayal of the monarch was to be found in numerous media of artistic expression, such as painting, sculpture, theatre, dance, music, and the almanacs that diffused royal propaganda to the population at large.
+Over his lifetime, Louis commissioned numerous works of art to portray himself, among them over 300 formal portraits. The earliest portrayals of Louis already followed the pictorial conventions of the day in depicting the child king as the majestically royal incarnation of France. This idealisation of the monarch continued in later works, which avoided depictions of the effect of the smallpox that Louis contracted in 1647. In the 1660s, Louis began to be shown as a Roman emperor, the god Apollo, or Alexander the Great, as can be seen in many works of Charles Le Brun, such as sculpture, paintings, and the decor of major monuments.
+The depiction of the king in this manner focused on allegorical or mythological attributes, instead of attempting to produce a true likeness. As Louis aged, so too did the manner in which he was depicted. Nonetheless, there was still a disparity between realistic representation and the demands of royal propaganda. There is no better illustration of this than in Hyacinthe Rigaud's frequently-reproduced Portrait of Louis XIV of 1701, in which a 63-year-old Louis appears to stand on a set of unnaturally young legs.[100]
+Rigaud's portrait exemplified the height of royal portraiture during Louis' reign. Although Rigaud crafted a credible likeness of Louis, the portrait was neither meant as an exercise in realism nor to explore Louis' personal character. Certainly, Rigaud was concerned with detail and depicted the king's costume with great precision, down to his shoe buckle.[101]
+However, Rigaud's intention was to glorify the monarchy. Rigaud's original, now housed in the Louvre, was originally meant as a gift to Louis' grandson, Philip V of Spain. However, Louis was so pleased with the work that he kept the original and commissioned a copy to be sent to his grandson. That became the first of many copies, both in full and half-length formats, to be made by Rigaud, often with the help of his assistants. The portrait also became a model for French royal and imperial portraiture down to the time of Charles X over a century later. In his work, Rigaud proclaims Louis' exalted royal status through his elegant stance and haughty expression, the royal regalia and throne, rich ceremonial fleur-de-lys robes, as well as the upright column in the background, which, together with the draperies, serves to frame this image of majesty.
+In addition to portraits, Louis commissioned at least 20 statues of himself in the 1680s, to stand in Paris and provincial towns as physical manifestations of his rule. He also commissioned "war artists" to follow him on campaigns to document his military triumphs. To remind the people of these triumphs, Louis erected permanent triumphal arches in Paris and the provinces for the first time since the decline of the Roman Empire.
+Louis' reign marked the birth and infancy of the art of medallions. Sixteenth-century rulers had often issued medals in small numbers to commemorate the major events of their reigns. Louis, however, struck more than 300 to celebrate the story of the king in bronze, that were enshrined in thousands of households throughout France.
+He also used tapestries as a medium of exalting the monarchy. Tapestries could be allegorical, depicting the elements or seasons, or realist, portraying royal residences or historical events. They were among the most significant means to spread royal propaganda prior to the construction of the Hall of Mirrors at Versailles.[102]
+Louis loved ballet and frequently danced in court ballets during the early half of his reign. In general, Louis was an eager dancer who performed 80 roles in 40 major ballets. This approaches the career of a professional ballet dancer.[103]
+His choices were strategic and varied. He danced four parts in three of Molière's comédies-ballets, which are plays accompanied by music and dance. Louis played an Egyptian in Le Mariage forcé in 1664, a Moorish gentleman in Le Sicilien in 1667, and both Neptune and Apollo in Les Amants magnifiques in 1670.
+He sometimes danced leading roles which were suitably royal or godlike (such as Neptune, Apollo, or the Sun).[103] At other times, he would adopt mundane roles before appearing at the end in the lead role. It is considered that, at all times, he provided his roles with sufficient majesty and drew the limelight with his flair for dancing.[103] For Louis, ballet may not have merely been a tool for manipulation in his propaganda machinery. The sheer number of performances he gave as well as the diversity of roles he played may serve to indicate a deeper understanding and interest in the art form.[104][105]
+Ballet dancing was actually used by Louis as a political tool to hold power over his state. [106] He integrated ballet deeply in court social functions and fixated his nobles' attention on upholding standards in ballet dancing, effectively distracting them from political activities. [107] In 1661, Royal of the Academy was founded by Louis to further his ambition. Pierre Beauchamp, his private dance instructor, was ordered by Louis to come up with a notation system to record ballet performances, which he did with great success. His work was adopted and published by Feuillet in 1700. This major development in ballet played an important role in promoting French culture and ballet throughout Europe during Louis' time. [108]
+Louis greatly emphasized etiquettes in ballet dancing, evidently seen in "La belle danse" (the French noble style). More challenging skills were required to perform this dance with movements very much resembling court behaviors, as a way to remind the nobles of the king's absolute power and their own status. All the details and rules were compressed in five positions of the bodies codified by Beauchamp. [109]
+Besides the official depiction and image of Louis, his subjects also followed a non-official discourse consisting mainly of clandestine publications, popular songs, and rumors that provided an alternative interpretation of Louis and his government. They often focused on the miseries arising from poor government, but also carried the hope for a better future when Louis escaped the malignant influence of his ministers and mistresses, and took the government into his own hands. On the other hand, petitions addressed either directly to Louis or to his ministers exploited the traditional imagery and language of monarchy. These varying interpretations of Louis abounded in self-contradictions that reflected the people's amalgamation of their everyday experiences with the idea of monarchy.[110]
+Despite the image of a healthy and virile king that Louis sought to project, evidence exists to suggest that his health was not very good. He had many ailments: for example, symptoms of diabetes, as confirmed in reports of suppurating periostitis in 1678, dental abscesses in 1696, along with recurring boils, fainting spells, gout, dizziness, hot flushes, and headaches.
+From 1647 to 1711, the three chief physicians to the king (Antoine Vallot, Antoine d'Aquin, and Guy-Crescent Fagon) recorded all of his health problems in the Journal de Santé du Roi (Journal of the King's Health), a daily report of his health. On 18 November 1686, Louis underwent a painful operation for an anal fistula that was performed by the surgeon Charles Felix de Tassy, who prepared a specially shaped curved scalpel for the occasion. The wound took more than two months to heal.[111]
+Louis died of gangrene at Versailles on 1 September 1715, four days before his 77th birthday, after 72 years on the throne. Enduring much pain in his last days, he finally "yielded up his soul without any effort, like a candle going out", while reciting the psalm Domine, ad adjuvandum me festina (O Lord, make haste to help me).[112] His body was laid to rest in Saint-Denis Basilica outside Paris. It remained there undisturbed for about 80 years, until revolutionaries exhumed and destroyed all of the remains found in the Basilica.[113]
+Louis outlived most of his immediate legitimate family. His last surviving in-wedlock son, the Dauphin, died in 1711. Barely a year later, the Duke of Burgundy, the eldest of the Dauphin's three sons and then heir to Louis, followed his father. Burgundy's elder son, Louis, Duke of Brittany, joined them a few weeks later. Thus, on his deathbed, Louis' heir was his five-year-old great-grandson, Louis, Duke of Anjou, Burgundy's younger son.
+Louis foresaw an underaged heir and sought to restrict the power of his nephew Philip II, Duke of Orléans, who, as his closest surviving legitimate relative in France, would likely become regent to the prospective Louis XV. Accordingly, the king created a regency council as Louis XIII had in anticipation of Louis XIV's own minority, with some power vested in his illegitimate son Louis-Auguste de Bourbon, Duke of Maine.[114] Orléans, however, had Louis' will annulled by the Parlement of Paris after his death and made himself sole regent. He stripped Maine and his brother, Louis-Alexandre, Count of Toulouse, of the rank of Prince of the Blood, which Louis had granted them, and significantly reduced Maine's power and privileges.[115]
+Line of succession to the French throne upon the death of Louis XIV in 1715. Louis XIV's only surviving legitimate grandson, Philip V, was not included in the line of succession due to having renounced the French throne after the war of the Spanish succession, which lasted for 13 years after the death of Charles II of Spain in 1700.[116]
+According to Philippe de Dangeau's Journal, Louis on his deathbed advised his heir with these words:
+Do not follow the bad example which I have set you; I have often undertaken war too lightly and have sustained it for vanity. Do not imitate me, but be a peaceful prince, and may you apply yourself principally to the alleviation of the burdens of your subjects.[117]
+Some historians point out that it was a customary demonstration of piety in those days to exaggerate one's sins. Thus they do not place much emphasis on Louis' deathbed declarations in assessing his accomplishments. Rather, they focus on military and diplomatic successes, such as how he placed a French prince on the Spanish throne. This, they contend, ended the threat of an aggressive Spain that historically interfered in domestic French politics. These historians also emphasise the effect of Louis' wars in expanding France's boundaries and creating more defensible frontiers that preserved France from invasion until the Revolution.[117]
+Arguably, Louis also applied himself indirectly to "the alleviation of the burdens of [his] subjects." For example, he patronised the arts, encouraged industry, fostered trade and commerce, and sponsored the founding of an overseas empire. Moreover, the significant reduction in civil wars and aristocratic rebellions during his reign are seen by these historians as the result of Louis' consolidation of royal authority over feudal elites.[118] In their analysis, his early reforms centralised France and marked the birth of the modern French state. They regard the political and military victories as well as numerous cultural achievements as the means by which Louis helped raise France to a preeminent position in Europe.[119] Europe came to admire France for its military and cultural successes, power, and sophistication. Europeans generally began to emulate French manners, values, goods, and deportment. French became the universal language of the European elite.
+Louis' detractors have argued that his considerable foreign, military, and domestic expenditure impoverished and bankrupted France. His supporters, however, distinguish the state, which was impoverished, from France, which was not. As supporting evidence, they cite the literature of the time, such as the social commentary in Montesquieu's Persian Letters.[120]
+Alternatively, Louis' critics attribute the social upheaval culminating in the French Revolution to his failure to reform French institutions while the monarchy was still secure. Other scholars counter that there was little reason to reform institutions that largely worked well under Louis. They also maintain that events occurring almost 80 years after his death were not reasonably foreseeable to Louis, and that in any case, his successors had sufficient time to initiate reforms of their own.[121]
+Louis has often been criticised for his vanity. The memoirist Saint-Simon, who claimed that Louis slighted him, criticised him thus:
+There was nothing he liked so much as flattery, or, to put it more plainly, adulation; the coarser and clumsier it was, the more he relished it.
+For his part, Voltaire saw Louis' vanity as the cause for his bellicosity:
+It is certain that he passionately wanted glory, rather than the conquests themselves. In the acquisition of Alsace and half of Flanders, and of all of Franche-Comté, what he really liked was the name he made for himself.[122]
+Nonetheless, Louis has also received praise. The anti-Bourbon Napoleon described him not only as "a great king", but also as "the only King of France worthy of the name".[123] Leibniz, the German Protestant philosopher, commended him as "one of the greatest kings that ever was".[124] And Lord Acton admired him as "by far the ablest man who was born in modern times on the steps of a throne".[125] The historian and philosopher Voltaire wrote: "His name can never be pronounced without respect and without summoning the image of an eternally memorable age".[126] Voltaire's history, The Age of Louis XIV, named Louis' reign as not only one of the four great ages in which reason and culture flourished, but the greatest ever.[127][128]
+In 1848, at Nuneham House, a piece of Louis' mummified heart, taken from his tomb and kept in a silver locket by Lord Harcourt, Archbishop of York, was shown to the Dean of Westminster, William Buckland, who ate it.[129]
+Numerous quotes have been attributed to Louis XIV by legend.
+The well-known "I am the state" ("L'état, c'est moi.") was reported from at least the late 18th century.[130]
+It was widely repeated but also denounced as apocryphal by the early 19th century.[131][b][132]
+He did say, "Every time I appoint someone to a vacant position, I make a hundred unhappy and one ungrateful."[133][134] Louis is recorded by numerous eyewitnesses as having said on his deathbed: "Je m'en vais, mais l'État demeurera toujours." ("I depart, but the State shall always remain.")[135]
+Louis's formal style was "Louis XIV, par la grâce de Dieu, roi de France et de Navarre", or "Louis XIV, by the Grace of God, King of France and of Navarre".
+On 5 April 1693, Louis also founded the Royal and Military Order of Saint Louis (French: Ordre Royal et Militaire de Saint-Louis), a military order of chivalry.[137][138] He named it after Louis IX and intended it as a reward for outstanding officers. It is notable as the first decoration that could be granted to non-nobles and is roughly the forerunner of the Légion d'honneur, with which it shares the red ribbon (though the Légion d'honneur is awarded to military personnel and civilians alike).
+Louis' patriline is the line from which he is descended father to son.
+Patrilineal descent is the principle behind membership in royal houses, as it can be traced back through the generations - which means that if King Louis were to choose an historically accurate house name it would be Robertian, as all his male-line ancestors have been of that house.
+Louis is a member of the House of Bourbon, a branch of the Capetian dynasty and of the Robertians.
+Louis' patriline is the line from which he is descended father to son. It follows the Bourbon, Kings of France, and the Counts of Paris and Worms. This line can be traced back more than 1,200 years from Robert of Hesbaye to the present day, through Kings of France & Navarre, Spain and Two-Sicilies, Dukes of Parma and Grand-Dukes of Luxembourg, Princes of Orléans and Emperors of Brazil. It is one of the oldest in Europe.
+This is an incomplete list of Louis XIV's illegitimate children. He reputedly had more, but the difficulty in fully documenting all such births restricts the list only to the better-known and/or legitimised.

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+Romanticism (also known as the Romantic era) was an artistic, literary, musical and intellectual movement that originated in Europe towards the end of the 18th century, and in most areas was at its peak in the approximate period from 1800 to 1890. Romanticism was characterized by its emphasis on emotion and individualism as well as glorification of all the past and nature, preferring the medieval rather than the classical.   It was partly a reaction to the Industrial Revolution,[1] the aristocratic social and political norms of the Age of Enlightenment, and the scientific rationalization of nature—all components of modernity.[2] It was embodied most strongly in the visual arts, music, and literature, but had a major impact on historiography,[3] education,[4] chess, social sciences, and the natural sciences.[5][failed verification] It had a significant and complex effect on politics, with romantic thinkers influencing liberalism, radicalism, conservatism, and nationalism.[6]
+The movement emphasized intense emotion as an authentic source of aesthetic experience, placing new emphasis on such emotions as apprehension, horror and terror, and awe—especially that experienced in confronting the new aesthetic categories of the sublimity and beauty of nature. It elevated folk art and ancient custom to something noble, but also spontaneity as a desirable characteristic (as in the musical impromptu). In contrast to the Rationalism and Classicism of the Enlightenment, Romanticism revived medievalism[7] and elements of art and narrative perceived as authentically medieval in an attempt to escape population growth, early urban sprawl, and industrialism.
+Although the movement was rooted in the German Sturm und Drang movement, which preferred intuition and emotion to the rationalism of the Enlightenment, the events and ideologies of the French Revolution were also proximate factors. Romanticism assigned a high value to the achievements of "heroic" individualists and artists, whose examples, it maintained, would raise the quality of society. It also promoted the individual imagination as a critical authority allowed of freedom from classical notions of form in art. There was a strong recourse to historical and natural inevitability, a Zeitgeist, in the representation of its ideas. In the second half of the 19th century, Realism was offered as a polar opposite to Romanticism.[8] The decline of Romanticism during this time was associated with multiple processes, including social and political changes and the spread of nationalism.[9]
+The nature of Romanticism may be approached from the primary importance of the free expression of the feelings of the artist. The importance the Romantics placed on emotion is summed up in the remark of the German painter Caspar David Friedrich, "the artist's feeling is his law".[10] For William Wordsworth, poetry should begin as "the spontaneous overflow of powerful feelings", which the poet then "recollect[s] in tranquility", evoking a new but corresponding emotion the poet can then mold into art.[11]
+To express these feelings, it was considered the content of art had to come from the imagination of the artist, with as little interference as possible from "artificial" rules dictating what a work should consist of. Samuel Taylor Coleridge and others believed there were natural laws the imagination—at least of a good creative artist—would unconsciously follow through artistic inspiration if left alone.[12] As well as rules, the influence of models from other works was considered to impede the creator's own imagination, so that originality was essential. The concept of the genius, or artist who was able to produce his own original work through this process of creation from nothingness, is key to Romanticism, and to be derivative was the worst sin.[13][14][15] This idea is often called "romantic originality".[16] Translator and prominent Romantic August Wilhelm Schlegel argued in his Lectures on Dramatic Arts and Letters that the most phenomenal power of human nature is its capacity to divide and diverge into opposite directions.[17]
+Not essential to Romanticism, but so widespread as to be normative, was a strong belief and interest in the importance of nature. This particularly in the effect of nature upon the artist when he is surrounded by it, preferably alone. In contrast to the usually very social art of the Enlightenment, Romantics were distrustful of the human world, and tended to believe a close connection with nature was mentally and morally healthy. Romantic art addressed its audiences with what was intended to be felt as the personal voice of the artist. So, in literature, "much of romantic poetry invited the reader to identify the protagonists with the poets themselves".[18]
+According to Isaiah Berlin, Romanticism embodied "a new and restless spirit, seeking violently to burst through old and cramping forms, a nervous preoccupation with perpetually changing inner states of consciousness, a longing for the unbounded and the indefinable, for perpetual movement and change, an effort to return to the forgotten sources of life, a passionate effort at self-assertion both individual and collective, a search after means of expressing an unappeasable yearning for unattainable goals".[19]
+The group of words with the root "Roman" in the various European languages, such as "romance" and "Romanesque", has a complicated history. By the 1700s, European languages – notably German, French and Russian – were using the term "Roman" in the sense of the English word "novel", i.e. a work of popular narrative fiction.[20] This usage derived from the term "Romance languages", which referred to vernacular (or popular) language in contrast to formal Latin.[20] Most such novels took the form of "chivalric romance", tales of adventure, devotion and honour.[21]
+The founders of Romanticism, critics August and Friedrich, began to speak of romantische Poesie ("romantic poetry") in the 1790s, contrasting it with "classic" but in terms of spirit rather than merely dating. Friedrich Schlegel wrote in his 1800 essay Gespräch über die Poesie ("Dialogue on Poetry"): "I seek and find the romantic among the older moderns, in Shakespeare, in Cervantes, in Italian poetry, in that age of chivalry, love and fable, from which the phenomenon and the word itself are derived."[22][23]
+The modern sense of the term spread more widely in France by its persistent use by Germaine de Staël in her De l'Allemagne (1813), recounting her travels in Germany.[24] In England Wordsworth wrote in a preface to his poems of 1815 of the "romantic harp" and "classic lyre",[24] but in 1820 Byron could still write, perhaps slightly disingenuously, "I perceive that in Germany, as well as in Italy, there is a great struggle about what they call 'Classical' and 'Romantic', terms which were not subjects of classification in England, at least when I left it four or five years ago".[25] It is only from the 1820s that Romanticism certainly knew itself by its name, and in 1824 the Académie française took the wholly ineffective step of issuing a decree condemning it in literature.[26]
+The period typically called Romantic varies greatly between different countries and different artistic media or areas of thought. Margaret Drabble described it in literature as taking place "roughly between 1770 and 1848",[27] and few dates much earlier than 1770 will be found. In English literature, M. H. Abrams placed it between 1789, or 1798, this latter a very typical view, and about 1830, perhaps a little later than some other critics.[28] Others have proposed 1780–1830.[29] In other fields and other countries the period denominated as Romantic can be considerably different; musical Romanticism, for example, is generally regarded as only having ceased as a major artistic force as late as 1910, but in an extreme extension the Four Last Songs of Richard Strauss are described stylistically as "Late Romantic" and were composed in 1946–48.[30] However, in most fields the Romantic period is said to be over by about 1850, or earlier.
+The early period of the Romantic era was a time of war, with the French Revolution (1789–1799) followed by the Napoleonic Wars until 1815. These wars, along with the political and social turmoil that went along with them, served as the background for Romanticism.[31] The key generation of French Romantics born between 1795–1805 had, in the words of one of their number, Alfred de Vigny, been "conceived between battles, attended school to the rolling of drums".[32] According to Jacques Barzun, there were three generations of Romantic artists. The first emerged in the 1790s and 1800s, the second in the 1820s, and the third later in the century.[33]
+The more precise characterization and specific definition of Romanticism has been the subject of debate in the fields of intellectual history and literary history throughout the 20th century, without any great measure of consensus emerging. That it was part of the Counter-Enlightenment, a reaction against the Age of Enlightenment, is generally accepted in current scholarship. Its relationship to the French Revolution, which began in 1789 in the very early stages of the period, is clearly important, but highly variable depending on geography and individual reactions. Most Romantics can be said to be broadly progressive in their views, but a considerable number always had, or developed, a wide range of conservative views,[34] and nationalism was in many countries strongly associated with Romanticism, as discussed in detail below.
+In philosophy and the history of ideas, Romanticism was seen by Isaiah Berlin as disrupting for over a century the classic Western traditions of rationality and the idea of moral absolutes and agreed values, leading "to something like the melting away of the very notion of objective truth",[35] and hence not only to nationalism, but also fascism and totalitarianism, with a gradual recovery coming only after World War II.[36] For the Romantics, Berlin says,
+in the realm of ethics, politics, aesthetics it was the authenticity and sincerity of the pursuit of inner goals that mattered; this applied equally to individuals and groups—states, nations, movements. This is most evident in the aesthetics of romanticism, where the notion of eternal models, a Platonic vision of ideal beauty, which the artist seeks to convey, however imperfectly, on canvas or in sound, is replaced by a passionate belief in spiritual freedom, individual creativity. The painter, the poet, the composer do not hold up a mirror to nature, however ideal, but invent; they do not imitate (the doctrine of mimesis), but create not merely the means but the goals that they pursue; these goals represent the self-expression of the artist's own unique, inner vision, to set aside which in response to the demands of some "external" voice—church, state, public opinion, family friends, arbiters of taste—is an act of betrayal of what alone justifies their existence for those who are in any sense creative.[37]
+Arthur Lovejoy attempted to demonstrate the difficulty of defining Romanticism in his seminal article "On The Discrimination of Romanticisms" in his Essays in the History of Ideas (1948); some scholars see Romanticism as essentially continuous with the present, some like Robert Hughes see in it the inaugural moment of modernity,[38] and some like Chateaubriand, Novalis and Samuel Taylor Coleridge see it as the beginning of a tradition of resistance to Enlightenment rationalism—a "Counter-Enlightenment"—[39][40] to be associated most closely with German Romanticism. An earlier definition comes from Charles Baudelaire: "Romanticism is precisely situated neither in choice of subject nor exact truth, but in the way of feeling."[41]
+The end of the Romantic era is marked in some areas by a new style of Realism, which affected literature, especially the novel and drama, painting, and even music, through Verismo opera. This movement was led by France, with Balzac and Flaubert in literature and Courbet in painting; Stendhal and Goya were important precursors of Realism in their respective media. However, Romantic styles, now often representing the established and safe style against which Realists rebelled, continued to flourish in many fields for the rest of the century and beyond. In music such works from after about 1850 are referred to by some writers as "Late Romantic" and by others as "Neoromantic" or "Postromantic", but other fields do not usually use these terms; in English literature and painting the convenient term "Victorian" avoids having to characterise the period further.
+In northern Europe, the Early Romantic visionary optimism and belief that the world was in the process of great change and improvement had largely vanished, and some art became more conventionally political and polemical as its creators engaged polemically with the world as it was. Elsewhere, including in very different ways the United States and Russia, feelings that great change was underway or just about to come were still possible. Displays of intense emotion in art remained prominent, as did the exotic and historical settings pioneered by the Romantics, but experimentation with form and technique was generally reduced, often replaced with meticulous technique, as in the poems of Tennyson or many paintings. If not realist, late 19th-century art was often extremely detailed, and pride was taken in adding authentic details in a way that earlier Romantics did not trouble with. Many Romantic ideas about the nature and purpose of art, above all the pre-eminent importance of originality, remained important for later generations, and often underlie modern views, despite opposition from theorists.
+In literature, Romanticism found recurrent themes in the evocation or criticism of the past, the cult of "sensibility" with its emphasis on women and children, the isolation of the artist or narrator, and respect for nature. Furthermore, several romantic authors, such as Edgar Allan Poe and Nathaniel Hawthorne, based their writings on the supernatural/occult and human psychology. Romanticism tended to regard satire as something unworthy of serious attention, a prejudice still influential today.[42] The Romantic movement in literature was preceded by the Enlightenment and succeeded by Realism.
+Some authors cite 16th-century poet Isabella di Morra as an early precursor of Romantic literature. Her lyrics covering themes of isolation and loneliness, which reflected the tragic events of her life, are considered "an impressive prefigurement of Romanticism",[43] differing from the Petrarchist fashion of the time based on the philosophy of love.
+The precursors of Romanticism in English poetry go back to the middle of the 18th century, including figures such as Joseph Warton (headmaster at Winchester College) and his brother Thomas Warton, Professor of Poetry at Oxford University.[44] Joseph maintained that invention and imagination were the chief qualities of a poet. The Scottish poet James Macpherson influenced the early development of Romanticism with the international success of his Ossian cycle of poems published in 1762, inspiring both Goethe and the young Walter Scott. Thomas Chatterton is generally considered the first Romantic poet in English.[45] Both Chatterton and Macpherson's work involved elements of fraud, as what they claimed was earlier literature that they had discovered or compiled was, in fact, entirely their own work. The Gothic novel, beginning with Horace Walpole's The Castle of Otranto (1764), was an important precursor of one strain of Romanticism, with a delight in horror and threat, and exotic picturesque settings, matched in Walpole's case by his role in the early revival of Gothic architecture. Tristram Shandy, a novel by Laurence Sterne (1759–67), introduced a whimsical version of the anti-rational sentimental novel to the English literary public.
+An early German influence came from Johann Wolfgang von Goethe, whose 1774 novel The Sorrows of Young Werther had young men throughout Europe emulating its protagonist, a young artist with a very sensitive and passionate temperament. At that time Germany was a multitude of small separate states, and Goethe's works would have a seminal influence in developing a unifying sense of nationalism. Another philosophic influence came from the German idealism of Johann Gottlieb Fichte and Friedrich Schelling, making Jena (where Fichte lived, as well as Schelling, Hegel, Schiller and the brothers Schlegel) a centre for early German Romanticism (see Jena Romanticism). Important writers were Ludwig Tieck, Novalis (Heinrich von Ofterdingen, 1799), Heinrich von Kleist and Friedrich Hölderlin. Heidelberg later became a centre of German Romanticism, where writers and poets such as Clemens Brentano, Achim von Arnim, and Joseph Freiherr von Eichendorff (Aus dem Leben eines Taugenichts) met regularly in literary circles.
+Important motifs in German Romanticism are travelling, nature, for example the German Forest, and Germanic myths. The later German Romanticism of, for example E. T. A. Hoffmann's Der Sandmann (The Sandman), 1817, and Joseph Freiherr von Eichendorff's Das Marmorbild (The Marble Statue), 1819, was darker in its motifs and has gothic elements.  The significance to Romanticism of childhood innocence, the importance of imagination, and racial theories all combined to give an unprecedented importance to folk literature, non-classical mythology and children's literature, above all in Germany.  Brentano and von Arnim were significant literary figures who together published Des Knaben Wunderhorn ("The Boy's Magic Horn" or cornucopia), a collection of versified folk tales, in 1806–08.  The first collection of Grimms' Fairy Tales by the Brothers Grimm was published in 1812.[46] Unlike the much later work of Hans Christian Andersen, who was publishing his invented tales in Danish from 1835, these German works were at least mainly based on collected folk tales, and the Grimms remained true to the style of the telling in their early editions, though later rewriting some parts. One of the brothers, Jacob, published in 1835 Deutsche Mythologie, a long academic work on Germanic mythology.[47] Another strain is exemplified by Schiller's highly emotional language and the depiction of physical violence in his play The Robbers of 1781.
+In English literature, the key figures of the Romantic movement are considered to be the group of poets including William Wordsworth, Samuel Taylor Coleridge, John Keats, Lord Byron, Percy Bysshe Shelley and the much older William Blake, followed later by the isolated figure of John Clare; also such novelists as Walter Scott from Scotland and Mary Shelley, and the essayists William Hazlitt and Charles Lamb. The publication in 1798 of Lyrical Ballads, with many of the finest poems by Wordsworth and Coleridge, is often held to mark the start of the movement. The majority of the poems were by Wordsworth, and many dealt with the lives of the poor in his native Lake District, or his feelings about nature—which he more fully developed in his long poem The Prelude, never published in his lifetime. The longest poem in the volume was Coleridge's The Rime of the Ancient Mariner, which showed the Gothic side of English Romanticism, and the exotic settings that many works featured. In the period when they were writing, the Lake Poets were widely regarded as a marginal group of radicals, though they were supported by the critic and writer William Hazlitt and others.
+In contrast, Lord Byron and Walter Scott achieved enormous fame and influence throughout Europe with works exploiting the violence and drama of their exotic and historical settings; Goethe called Byron "undoubtedly the greatest genius of our century".[48] Scott achieved immediate success with his long narrative poem The Lay of the Last Minstrel in 1805, followed by the full epic poem Marmion in 1808. Both were set in the distant Scottish past, already evoked in Ossian; Romanticism and Scotland were to have a long and fruitful partnership. Byron had equal success with the first part of Childe Harold's Pilgrimage in 1812, followed by four "Turkish tales", all in the form of long poems, starting with The Giaour in 1813, drawing from his Grand Tour, which had reached Ottoman Europe, and orientalizing the themes of the Gothic novel in verse. These featured different variations of the "Byronic hero", and his own life contributed a further version. Scott meanwhile was effectively inventing the historical novel, beginning in 1814 with Waverley, set in the 1745 Jacobite rising, which was an enormous and highly profitable success, followed by over 20 further Waverley Novels over the next 17 years, with settings going back to the Crusades that he had researched to a degree that was new in literature.[49]
+In contrast to Germany, Romanticism in English literature had little connection with nationalism, and the Romantics were often regarded with suspicion for the sympathy many felt for the ideals of the French Revolution, whose collapse and replacement with the dictatorship of Napoleon was, as elsewhere in Europe, a shock to the movement. Though his novels celebrated Scottish identity and history, Scott was politically a firm Unionist, but admitted to Jacobite sympathies. Several spent much time abroad, and a famous stay on Lake Geneva with Byron and Shelley in 1816 produced the hugely influential novel Frankenstein by Shelley's wife-to-be Mary Shelley and the novella The Vampyre by Byron's doctor John William Polidori. The lyrics of Robert Burns in Scotland, and Thomas Moore from Ireland, reflected in different ways their countries and the Romantic interest in folk literature, but neither had a fully Romantic approach to life or their work.
+Though they have modern critical champions such as György Lukács, Scott's novels are today more likely to be experienced in the form of the many operas that composers continued to base on them over the following decades, such as Donizetti's Lucia di Lammermoor and Vincenzo Bellini's I puritani (both 1835). Byron is now most highly regarded for his short lyrics and his generally unromantic prose writings, especially his letters, and his unfinished satire Don Juan.[50] Unlike many Romantics, Byron's widely publicised personal life appeared to match his work, and his death at 36 in 1824 from disease when helping the Greek War of Independence appeared from a distance to be a suitably Romantic end, entrenching his legend.[51] Keats in 1821 and Shelley in 1822 both died in Italy, Blake (at almost 70) in 1827, and Coleridge largely ceased to write in the 1820s. Wordsworth was by 1820 respectable and highly regarded, holding a government sinecure, but wrote relatively little. In the discussion of English literature, the Romantic period is often regarded as finishing around the 1820s, or sometimes even earlier, although many authors of the succeeding decades were no less committed to Romantic values.
+The most significant novelist in English during the peak Romantic period, other than Walter Scott, was Jane Austen, whose essentially conservative world-view had little in common with her Romantic contemporaries, retaining a strong belief in decorum and social rules, though critics such as Claudia L. Johnson have detected tremors under the surface of many works, such as Northanger Abbey (1817), Mansfield Park (1814) and Persuasion (1817).[52] But around the mid-century the undoubtedly Romantic novels of the Yorkshire-based Brontë family appeared. Most notably Charlotte's Jane Eyre and Emily's Wuthering Heights, both published in 1847, which also introduced more Gothic themes. While these two novels were written and published after the Romantic period is said to have ended, their novels were heavily influenced by Romantic literature they'd read as children.
+Byron, Keats and Shelley all wrote for the stage, but with little success in England, with Shelley's The Cenci perhaps the best work produced, though that was not played in a public theatre in England until a century after his death. Byron's plays, along with dramatizations of his poems and Scott's novels, were much more popular on the Continent, and especially in France, and through these versions several were turned into operas, many still performed today. If contemporary poets had little success on the stage, the period was a legendary one for performances of Shakespeare, and went some way to restoring his original texts and removing the Augustan "improvements" to them. The greatest actor of the period, Edmund Kean, restored the tragic ending to King Lear;[53] Coleridge said that, "Seeing him act was like reading Shakespeare by flashes of lightning."[54]
+Although after union with England in 1707 Scotland increasingly adopted English language and wider cultural norms, its literature developed a distinct national identity and began to enjoy an international reputation. Allan Ramsay (1686–1758) laid the foundations of a reawakening of interest in older Scottish literature, as well as leading the trend for pastoral poetry, helping to develop the Habbie stanza as a poetic form.[55] James Macpherson (1736–96) was the first Scottish poet to gain an international reputation. Claiming to have found poetry written by the ancient bard Ossian, he published translations that acquired international popularity, being proclaimed as a Celtic equivalent of the Classical epics. Fingal, written in 1762, was speedily translated into many European languages, and its appreciation of natural beauty and treatment of the ancient legend has been credited more than any single work with bringing about the Romantic movement in European, and especially in German literature, through its influence on Johann Gottfried von Herder and Johann Wolfgang von Goethe.[56] It was also popularised in France by figures that included Napoleon.[57] Eventually it became clear that the poems were not direct translations from Scottish Gaelic, but flowery adaptations made to suit the aesthetic expectations of his audience.[58]
+Robert Burns (1759–96) and Walter Scott (1771–1832) were highly influenced by the Ossian cycle. Burns, an Ayrshire poet and lyricist, is widely regarded as the national poet of Scotland and a major influence on the Romantic movement. His poem (and song) "Auld Lang Syne" is often sung at Hogmanay (the last day of the year), and "Scots Wha Hae" served for a long time as an unofficial national anthem of the country.[59] Scott began as a poet and also collected and published Scottish ballads. His first prose work, Waverley in 1814, is often called the first historical novel.[60] It launched a highly successful career, with other historical novels such as Rob Roy (1817), The Heart of Midlothian (1818) and Ivanhoe (1820). Scott probably did more than any other figure to define and popularise Scottish cultural identity in the nineteenth century.[61] Other major literary figures connected with Romanticism include the poets and novelists James Hogg (1770–1835), Allan Cunningham (1784–1842) and John Galt (1779–1839).[62] One of the most significant figures of the Romantic movement, Lord Byron, was brought up in Scotland until he inherited his family's English peerage.[63]
+Scotland was also the location of two of the most important literary magazines of the era, The Edinburgh Review (founded in 1802) and Blackwood's Magazine (founded in 1817), which had a major impact on the development of British literature and drama in the era of Romanticism.[64][65] Ian Duncan and Alex Benchimol suggest that publications like the novels of Scott and these magazines were part of a highly dynamic Scottish Romanticism that by the early nineteenth century, caused Edinburgh to emerge as the cultural capital of Britain and become central to a wider formation of a "British Isles nationalism".[66]
+Scottish "national drama" emerged in the early 1800s, as plays with specifically Scottish themes began to dominate the Scottish stage. Theatres had been discouraged by the Church of Scotland and fears of Jacobite assemblies. In the later eighteenth century, many plays were written for and performed by small amateur companies and were not published and so most have been lost. Towards the end of the century there were "closet dramas", primarily designed to be read, rather than performed, including work by Scott, Hogg, Galt and Joanna Baillie (1762–1851), often influenced by the ballad tradition and Gothic Romanticism.[67]
+Romanticism was relatively late in developing in French literature, more so than in the visual arts. The 18th-century precursor to Romanticism, the cult of sensibility, had become associated with the Ancien Régime, and the French Revolution had been more of an inspiration to foreign writers than those experiencing it at first-hand. The first major figure was François-René de Chateaubriand, a minor aristocrat who had remained a royalist throughout the Revolution, and returned to France from exile in England and America under Napoleon, with whose regime he had an uneasy relationship. His writings, all in prose, included some fiction, such as his influential novella of exile René (1802), which anticipated Byron in its alienated hero, but mostly contemporary history and politics, his travels, a defence of religion and the medieval spirit (Génie du christianisme, 1802), and finally in the 1830s and 1840s his enormous autobiography Mémoires d'Outre-Tombe ("Memoirs from beyond the grave").[68]
+After the Bourbon Restoration, French Romanticism developed in the lively world of Parisian theatre, with productions of Shakespeare, Schiller (in France a key Romantic author), and adaptations of Scott and Byron alongside French authors, several of whom began to write in the late 1820s. Cliques of pro- and anti-Romantics developed, and productions were often accompanied by raucous vocalizing by the two sides, including the shouted assertion by one theatregoer in 1822 that "Shakespeare, c'est l'aide-de-camp de Wellington" ("Shakespeare is Wellington's aide-de-camp").[69] Alexandre Dumas began as a dramatist, with a series of successes beginning with Henri III et sa cour (1829) before turning to novels that were mostly historical adventures somewhat in the manner of Scott, most famously The Three Musketeers and The Count of Monte Cristo, both of 1844. Victor Hugo published as a poet in the 1820s before achieving success on the stage with Hernani—a historical drama in a quasi-Shakespearian style that had famously riotous performances on its first run in 1830.[70] Like Dumas, Hugo is best known for his novels, and was already writing The Hunchback of Notre-Dame (1831), one of the best known works, which became a paradigm of the French Romantic movement. The preface to his unperformed play Cromwell gives an important manifesto of French Romanticism, stating that "there are no rules, or models". The career of Prosper Mérimée followed a similar pattern; he is now best known as the originator of the story of Carmen, with his novella published 1845. Alfred de Vigny remains best known as a dramatist, with his play on the life of the English poet Chatterton (1835) perhaps his best work. George Sand was a central figure of the Parisian literary scene, famous both for her novels and criticism and her affairs with Chopin and several others;[71] she too was inspired by the theatre, and wrote works to be staged at her private estate.
+French Romantic poets of the 1830s to 1850s include Alfred de Musset, Gérard de Nerval, Alphonse de Lamartine and the flamboyant Théophile Gautier, whose prolific output in various forms continued until his death in 1872.
+Stendhal is today probably the most highly regarded French novelist of the period, but he stands in a complex relation with Romanticism, and is notable for his penetrating psychological insight into his characters and his realism, qualities rarely prominent in Romantic fiction. As a survivor of the French retreat from Moscow in 1812, fantasies of heroism and adventure had little appeal for him, and like Goya he is often seen as a forerunner of Realism. His most important works are Le Rouge et le Noir (The Red and the Black, 1830) and La Chartreuse de Parme (The Charterhouse of Parma, 1839).
+Romanticism in Poland is often taken to begin with the publication of Adam Mickiewicz's first poems in 1822, and end with the crushing of the January Uprising of 1863 against the Russians. It was strongly marked by interest in Polish history.[72] Polish Romanticism revived the old "Sarmatism" traditions of the szlachta or Polish nobility. Old traditions and customs were revived and portrayed in a positive light in the Polish messianic movement and in works of great Polish poets such as Adam Mickiewicz (Pan Tadeusz), Juliusz Słowacki and Zygmunt Krasiński, as well as prose writers such as Henryk Sienkiewicz. This close connection between Polish Romanticism and Polish history became one of the defining qualities of the literature of Polish Romanticism period, differentiating it from that of other countries. They had not suffered the loss of national statehood as was the case with Poland.[73] Influenced by the general spirit and main ideas of European Romanticism, the literature of Polish Romanticism is unique, as many scholars have pointed out, in having developed largely outside of Poland and in its emphatic focus upon the issue of Polish nationalism. The Polish intelligentsia, along with leading members of its government, left Poland in the early 1830s, during what is referred to as the "Great Emigration", resettling in France, Germany, Great Britain, Turkey, and the United States.
+Their art featured emotionalism and irrationality, fantasy and imagination, personality cults, folklore and country life, and the propagation of ideals of freedom. In the second period, many of the Polish Romantics worked abroad, often banished from Poland by the occupying powers due to their politically subversive ideas. Their work became increasingly dominated by the ideals of political struggle for freedom and their country's sovereignty. Elements of mysticism became more prominent. There developed the idea of the poeta wieszcz (the prophet). The wieszcz (bard) functioned as spiritual leader to the nation fighting for its independence. The most notable poet so recognized was Adam Mickiewicz.
+Zygmunt Krasiński also wrote to inspire political and religious hope in his countrymen. Unlike his predecessors, who called for victory at whatever price in Poland's struggle against Russia, Krasinski emphasized Poland's spiritual role in its fight for independence, advocating an intellectual rather than a military superiority. His works best exemplify the Messianic movement in Poland: in two early dramas, Nie-boska komedia (1835; The Undivine Comedy) and Irydion (1836; Iridion), as well as in the later Psalmy przyszłości (1845), he asserted that Poland was the Christ of Europe: specifically chosen by God to carry the world's burdens, to suffer, and eventually be resurrected.
+Early Russian Romanticism is associated with the writers Konstantin Batyushkov (A Vision on the Shores of the Lethe, 1809), Vasily Zhukovsky (The Bard, 1811; Svetlana, 1813) and Nikolay Karamzin (Poor Liza, 1792; Julia, 1796; Martha the Mayoress, 1802; The Sensitive and the Cold, 1803). However the principal exponent of Romanticism in Russia is Alexander Pushkin (The Prisoner of the Caucasus, 1820–1821; The Robber Brothers, 1822; Ruslan and Ludmila, 1820; Eugene Onegin, 1825–1832). Pushkin's work influenced many writers in the 19th century and led to his eventual recognition as Russia's greatest poet.[74] Other Russian Romantic poets include Mikhail Lermontov (A Hero of Our Time, 1839), Fyodor Tyutchev (Silentium!, 1830), Yevgeny Baratynsky (Eda, 1826), Anton Delvig, and Wilhelm Küchelbecker.
+Influenced heavily by Lord Byron, Lermontov sought to explore the Romantic emphasis on metaphysical discontent with society and self, while Tyutchev's poems often described scenes of nature or passions of love. Tyutchev commonly operated with such categories as night and day, north and south, dream and reality, cosmos and chaos, and the still world of winter and spring teeming with life. Baratynsky's style was fairly classical in nature, dwelling on the models of the previous century.
+Romanticism in Spanish literature developed a well-known literature with a huge variety of poets and playwrights. The most important Spanish poet during this movement was José de Espronceda. After him there were other poets like Gustavo Adolfo Bécquer, Mariano José de Larra and the dramatists Ángel de Saavedra and José Zorrilla, author of Don Juan Tenorio. Before them may be mentioned the pre-romantics José Cadalso and Manuel José Quintana.[76]  The plays of Antonio García Gutiérrez were adapted to produce Giuseppe Verdi's operas Il trovatore and Simon Boccanegra. Spanish Romanticism also influenced regional literatures. For example, in Catalonia and in Galicia there was a national boom of writers in the local languages, like the Catalan Jacint Verdaguer and the Galician Rosalía de Castro, the main figures of the national revivalist movements Renaixença and Rexurdimento, respectively.[77]
+There are scholars who consider Spanish Romanticism to be Proto-Existentialism because it is more anguished than the movement in other European countries. Foster et al., for example, say that the work of Spain's writers such as Espronceda, Larra, and other writers in the 19th century demonstrated a "metaphysical crisis".[78] These observers put more weight on the link between the 19th-century Spanish writers with the existentialist movement that emerged immediately after. According to Richard Caldwell, the writers that we now identify with Spain's romanticism were actually precursors to those who galvanized the literary movement that emerged in the 1920s.[79] This notion is the subject of debate for there are authors who stress that Spain's romanticism is one of the earliest in Europe,[80] while some assert that Spain really had no period of literary romanticism.[81] This controversy underscores a certain uniqueness to Spanish Romanticism in comparison to its European counterparts.
+Romanticism began in Portugal with the publication of the poem Camões (1825), by Almeida Garrett, who was raised by his uncle D. Alexandre, bishop of Angra, in the precepts of Neoclassicism, which can be observed in his early work. The author himself confesses (in Camões' preface) that he voluntarily refused to follow the principles of epic poetry enunciated by Aristotle in his Poetics, as he did the same to Horace's Ars Poetica. Almeida Garrett had participated in the 1820 Liberal Revolution, which caused him to exile himself in England in 1823 and then in France, after the Vila-Francada. While living in Great Britain, he had contacts with the Romantic movement and read authors such as Shakespeare, Scott, Ossian, Byron, Hugo, Lamartine and de Staël, at the same time visiting feudal castles and ruins of Gothic churches and abbeys, which would be reflected in his writings. In 1838, he presented Um Auto de Gil Vicente ("A Play by Gil Vicente"), in an attempt to create a new national theatre, free of Greco-Roman and foreign influence. But his masterpiece would be Frei Luís de Sousa (1843), named by himself as a "Romantic drama" and it was acclaimed as an exceptional work, dealing with themes as national independence, faith, justice and love. He was also deeply interested in Portuguese folkloric verse, which resulted in the publication of Romanceiro ("Traditional Portuguese Ballads") (1843), that recollect a great number of ancient popular ballads, known as "romances" or "rimances", in redondilha maior verse form, that contained stories of chivalry, life of saints, crusades, courtly love, etc. He wrote the novels Viagens na Minha Terra, O Arco de Sant'Ana and Helena.[82][83][84]
+Alexandre Herculano is, alongside Almeida Garrett, one of the founders of Portuguese Romanticism. He too was forced to exile to Great Britain and France because of his liberal ideals. All of his poetry and prose are (unlike Almeida Garrett's) entirely Romantic, rejecting Greco-Roman myth and history. He sought inspiration in medieval Portuguese poems and chronicles as in the Bible. His output is vast and covers many different genres, such as historical essays, poetry, novels, opuscules and theatre, where he brings back a whole world of Portuguese legends, tradition and history, especially in Eurico, o Presbítero ("Eurico, the Priest") and Lendas e Narrativas ("Legends and Narratives"). His work was influenced by Chateaubriand, Schiller, Klopstock, Walter Scott and the Old Testament Psalms.[85]
+António Feliciano de Castilho made the case for Ultra-Romanticism, publishing the poems A Noite no Castelo ("Night in the Castle") and Os Ciúmes do Bardo ("The Jealousy of the Bard"), both in 1836, and the drama Camões. He became an unquestionable master for successive Ultra-Romantic generations, whose influence would not be challenged until the famous Coimbra Question. He also created polemics by translating Goethe's Faust without knowing German, but using French versions of the play. Other notable figures of Portuguese Romanticism are the famous novelists Camilo Castelo Branco and Júlio Dinis, and Soares de Passos, Bulhão Pato and Pinheiro Chagas.[84]
+Romantic style would be revived in the beginning of the 20th century, notably through the works of poets linked to the Portuguese Renaissance, such as Teixeira de Pascoais, Jaime Cortesão, Mário Beirão, among others, who can be considered Neo-Romantics. An early Portuguese expression of Romanticism is found already in poets such as Manuel Maria Barbosa du Bocage (especially in his sonnets dated at the end of the 18th century) and Leonor de Almeida Portugal, Marquise of Alorna.[84]
+Romanticism in Italian literature was a minor movement although some important works were produced; it began officially in 1816 when Germaine de Staël wrote an article in the journal Biblioteca italiana called "Sulla maniera e l'utilità delle traduzioni", inviting Italian people to reject Neoclassicism and to study new authors from other countries. Before that date, Ugo Foscolo had already published poems anticipating Romantic themes. The most important Romantic writers were Ludovico di Breme, Pietro Borsieri and Giovanni Berchet.[87] Better known authors such as Alessandro Manzoni and Giacomo Leopardi were influenced by Enlightenment as well as by Romanticism and Classicism.[88]
+Spanish-speaking South American Romanticism was influenced heavily by Esteban Echeverría, who wrote in the 1830 and 1840s. His writings were influenced by his hatred for the Argentine dictator Juan Manuel de Rosas, and filled with themes of blood and terror, using the metaphor of a slaughterhouse to portray the violence of Rosas' dictatorship.
+Brazilian Romanticism is characterized and divided in three different periods. The first one is basically focused on the creation of a sense of national identity, using the ideal of the heroic Indian. Some examples include José de Alencar, who wrote Iracema and O Guarani, and Gonçalves Dias, renowned by the poem "Canção do exílio" (Song of the Exile). The second period, sometimes called Ultra-Romanticism, is marked by a profound influence of European themes and traditions, involving the melancholy, sadness and despair related to unobtainable love. Goethe and Lord Byron are commonly quoted in these works. Some of the most notable authors of this phase are Álvares de Azevedo, Casimiro de Abreu, Fagundes Varela and Junqueira Freire. The third cycle is marked by social poetry, especially the abolitionist movement, and it includes Castro Alves, Tobias Barreto and Pedro Luís Pereira de Sousa.[89]
+In the United States, at least by 1818 with William Cullen Bryant's "To a Waterfowl", Romantic poetry was being published. American Romantic Gothic literature made an early appearance with Washington Irving's The Legend of Sleepy Hollow (1820) and Rip Van Winkle (1819), followed from 1823 onwards by the Leatherstocking Tales of James Fenimore Cooper, with their emphasis on heroic simplicity and their fervent landscape descriptions of an already-exotic mythicized frontier peopled by "noble savages", similar to the philosophical theory of Rousseau, exemplified by Uncas, from The Last of the Mohicans. There are picturesque "local colour" elements in Washington Irving's essays and especially his travel books. Edgar Allan Poe's tales of the macabre and his balladic poetry were more influential in France than at home, but the romantic American novel developed fully with the atmosphere and melodrama of Nathaniel Hawthorne's The Scarlet Letter (1850). Later Transcendentalist writers such as Henry David Thoreau and Ralph Waldo Emerson still show elements of its influence and imagination, as does the romantic realism of Walt Whitman. The poetry of Emily Dickinson—nearly unread in her own time—and Herman Melville's novel Moby-Dick can be taken as epitomes of American Romantic literature. By the 1880s, however, psychological and social realism were competing with Romanticism in the novel.
+The European Romantic movement reached America in the early 19th century. American Romanticism was just as multifaceted and individualistic as it was in Europe. Like the Europeans, the American Romantics demonstrated a high level of moral enthusiasm, commitment to individualism and the unfolding of the self, an emphasis on intuitive perception, and the assumption that the natural world was inherently good, while human society was filled with corruption.[90]
+Romanticism became popular in American politics, philosophy and art. The movement appealed to the revolutionary spirit of America as well as to those longing to break free of the strict religious traditions of early settlement. The Romantics rejected rationalism and religious intellect. It appealed to those in opposition of Calvinism, which includes the belief that the destiny of each individual is preordained. The Romantic movement gave rise to New England Transcendentalism, which portrayed a less restrictive relationship between God and Universe. The new philosophy presented the individual with a more personal relationship with God. Transcendentalism and Romanticism appealed to Americans in a similar fashion, for both privileged feeling over reason, individual freedom of expression over the restraints of tradition and custom. It often involved a rapturous response to nature. It encouraged the rejection of harsh, rigid Calvinism, and promised a new blossoming of American culture.[90][91]
+American Romanticism embraced the individual and rebelled against the confinement of neoclassicism and religious tradition. The Romantic movement in America created a new literary genre that continues to influence American writers. Novels, short stories, and poems replaced the sermons and manifestos of yore. Romantic literature was personal, intense, and portrayed more emotion than ever seen in neoclassical literature. America's preoccupation with freedom became a great source of motivation for Romantic writers as many were delighted in free expression and emotion without so much fear of ridicule and controversy. They also put more effort into the psychological development of their characters, and the main characters typically displayed extremes of sensitivity and excitement.[92]
+The works of the Romantic Era also differed from preceding works in that they spoke to a wider audience, partly reflecting the greater distribution of books as costs came down during the period.[31]
+Romantic architecture appeared in the late 18th century in a reaction against the rigid forms of neoclassical architecture. Romantic architecture reached its peak in the mid-19th century, and continued to appear until the end of the 19th century. It was designed to evoke an emotional reaction, either respect for tradition or nostalgia for a bucolic past. It was frequently inspired by the architecture of the Middle Ages, especially Gothic architecture, It was strongly influenced by romanticism in literature, particularly the historical novels of Victor Hugo and Walter Scott. It sometimes moved into the domain of eclecticism, with features assembled from different historic periods and regions of the world.[93]
+Gothic Revival architecture was a popular variant of the romantic style, particularly in the construction of churches, Cathedrals, and university buildings. Notable examples include the completion of Cologne Cathedral in Germany, by Karl Friedrich Schinkel. The cathedral had been begun in 1248, but work was halted in 1473. The original plans for the façade were discovered in 1840, and it was decided to recommence. Schinkel followed the original design as much as possible, but used modern construction technology, including an iron frame for the roof. The building was finished in 1880.[94]
+In Britain, notable examples include the Royal Pavilion in Brighton, a romantic version of traditional Indian architecture by John Nash (1815–1823), and the Houses of Parliament in London, built in a Gothic revival style by Charles Barry between 1840 and 1876.[95]
+In France, one of the earliest examples of romantic architecture is the Hameau de la Reine, the small rustic hamlet created at the Palace of Versailles for Queen Marie Antoinette between 1783 and 1785 by the royal architect Richard Mique with the help of the romantic painter Hubert Robert. It consisted of twelve structures, ten of which still exist, in the style of villages in Normandy. It was designed for the Queen and her friends to amuse themselves by playing at being peasants, and included a farmhouse with a dairy, a mill, a boudoir, a pigeon loft, a tower in the form of a lighthouse from which one could fish in the pond, a belvedere, a cascade and grotto, and a luxuriously furnished cottage with a billiard room for the Queen.[96]
+French romantic architecture in the 19th century was strongly influenced by two writers; Victor Hugo, whose novel The Hunchback of Notre Dame inspired a resurgence in interest in the Middle Ages; and Prosper Mérimée, who wrote celebrated romantic novels and short stories and was also the first head of the commission of Historic Monuments in France, responsible for publicizing and restoring (and sometimes romanticizing) many French cathedrals and monuments desecrated and ruined after the French Revolution. His projects were carried out by the architect Eugène Viollet-le-Duc. These included the restoration (sometimes creative) of the Cathedral of Notre Dame de Paris, the fortified city of Carcassonne, and the unfinished medieval Château de Pierrefonds.[94][97]
+The romantic style continued in the second half of the 19th century. The Palais Garnier, the Paris opera house designed by Charles Garnier was a highly romantic and eclectic combination of artistic styles.  Another notable example of late 19th century romanticism is the Basilica of Sacré-Cœur by Paul Abadie, who drew upon the model of Byzantine architecture for his elongated domes (1875–1914).[95]
+Hameau de la Reine, Palace of Versailles (1783–1785)
+Royal Pavilion in Brighton by John Nash (1815–1823)
+Cologne Cathedral (1840–80)
+Grand Staircase of the Paris Opera by Charles Garnier (1861–75)
+Basilica of Sacré-Cœur by Paul Abadie (1875–1914)
+In the visual arts, Romanticism first showed itself in landscape painting, where from as early as the 1760s British artists began to turn to wilder landscapes and storms, and Gothic architecture, even if they had to make do with Wales as a setting. Caspar David Friedrich and J. M. W. Turner were born less than a year apart in 1774 and 1775 respectively and were to take German and English landscape painting to their extremes of Romanticism, but both their artistic sensibilities were formed when forms of Romanticism was already strongly present in art. John Constable, born in 1776, stayed closer to the English landscape tradition, but in his largest "six-footers" insisted on the heroic status of a patch of the working countryside where he had grown up—challenging the traditional hierarchy of genres, which relegated landscape painting to a low status. Turner also painted very large landscapes, and above all, seascapes. Some of these large paintings had contemporary settings and staffage, but others had small figures that turned the work into history painting in the manner of Claude Lorrain, like Salvator Rosa, a late Baroque artist whose landscapes had elements that Romantic painters repeatedly turned to. Friedrich often used single figures, or features like crosses, set alone amidst a huge landscape, "making them images of the transitoriness of human life and the premonition of death".[98]
+Other groups of artists expressed feelings that verged on the mystical, many largely abandoning classical drawing and proportions.  These included William Blake and Samuel Palmer and the other members of the Ancients in England, and in Germany Philipp Otto Runge. Like Friedrich, none of these artists had significant influence after their deaths for the rest of the 19th century, and were 20th-century rediscoveries from obscurity, though Blake was always known as a poet, and Norway's leading painter Johan Christian Dahl was heavily influenced by Friedrich. The Rome-based Nazarene movement of German artists, active from 1810, took a very different path, concentrating on medievalizing history paintings with religious and nationalist themes.[99]
+The arrival of Romanticism in French art was delayed by the strong hold of Neoclassicism on the academies, but from the Napoleonic period it became increasingly popular, initially in the form of history paintings propagandising for the new regime, of which Girodet's Ossian receiving the Ghosts of the French Heroes, for Napoleon's Château de Malmaison, was one of the earliest. Girodet's old teacher David was puzzled and disappointed by his pupil's direction, saying: "Either Girodet is mad or I no longer know anything of the art of painting".[100] A new generation of the French school,[101] developed personal Romantic styles, though still concentrating on history painting with a political message. Théodore Géricault (1791–1824) had his first success with The Charging Chasseur, a heroic military figure derived from Rubens, at the Paris Salon of 1812 in the years of the Empire, but his next major completed work, The Raft of the Medusa of 1821, remains the greatest achievement of the Romantic history painting, which in its day had a powerful anti-government message.
+Eugène Delacroix (1798–1863) made his first Salon hits with The Barque of Dante (1822), The Massacre at Chios (1824) and Death of Sardanapalus (1827). The second was a scene from the Greek War of Independence, completed the year Byron died there, and the last was a scene from one of Byron's plays. With Shakespeare, Byron was to provide the subject matter for many other works of Delacroix, who also spent long periods in North Africa, painting colourful scenes of mounted Arab warriors. His Liberty Leading the People (1830) remains, with the Medusa, one of the best-known works of French Romantic painting. Both reflected current events, and increasingly "history painting", literally "story painting", a phrase dating back to the Italian Renaissance meaning the painting of subjects with groups of figures, long considered the highest and most difficult form of art, did indeed become the painting of historical scenes, rather than those from religion or mythology.[102]
+Francisco Goya was called "the last great painter in whose art thought and observation were balanced and combined to form a faultless unity".[103]  But the extent to which he was a Romantic is a complex question. In Spain, there was still a struggle to introduce the values of the Enlightenment, in which Goya saw himself as a participant.  The demonic and anti-rational monsters thrown up by his imagination are only superficially similar to those of the Gothic fantasies of northern Europe, and in many ways he remained wedded to the classicism and realism of his training, as well as looking forward to the Realism of the later 19th century.[104] But he, more than any other artist of the period, exemplified the Romantic values of the expression of the artist's feelings and his personal imaginative world.[105] He also shared with many of the Romantic painters a more free handling of paint, emphasized in the new prominence of the brushstroke and impasto, which tended to be repressed in neoclassicism under a self-effacing finish.
+Sculpture remained largely impervious to Romanticism, probably partly for technical reasons, as the most prestigious material of the day, marble, does not lend itself to expansive gestures. The leading sculptors in Europe, Antonio Canova and Bertel Thorvaldsen, were both based in Rome and firm Neoclassicists, not at all tempted to allow influence from medieval sculpture, which would have been one possible approach to Romantic sculpture. When it did develop, true Romantic sculpture—with the exception of a few artists such as Rudolf Maison—[106] rather oddly was missing in Germany, and mainly found in France, with François Rude, best known from his group of the 1830s from the Arc de Triomphe in Paris, David d'Angers, and Auguste Préault. Préault's plaster relief entitled Slaughter, which represented the horrors of wars with exacerbated passion, caused so much scandal at the 1834 Salon that Préault was banned from this official annual exhibition for nearly twenty years.[107] In Italy, the most important Romantic sculptor was Lorenzo Bartolini.[108]
+Francisco Goya, The Third of May 1808, 1814
+Théodore Géricault, The Raft of the Medusa, 1819
+Eugène Delacroix, Liberty Leading the People 1830
+J. M. W. Turner, The Fighting Téméraire tugged to her last Berth to be broken up, 1839
+In France, historical painting on idealized medieval and Renaissance themes is known as the style Troubadour, a term with no equivalent for other countries, though the same trends occurred there. Delacroix, Ingres and Richard Parkes Bonington all worked in this style, as did lesser specialists such as Pierre-Henri Révoil (1776–1842) and Fleury-François Richard (1777–1852). Their pictures are often small, and feature intimate private and anecdotal moments, as well as those of high drama. The lives of great artists such as Raphael were commemorated on equal terms with those of rulers, and fictional characters were also depicted. Fleury-Richard's Valentine of Milan weeping for the death of her husband, shown in the Paris Salon of 1802, marked the arrival of the style, which lasted until the mid-century, before being subsumed into the increasingly academic history painting of artists like Paul Delaroche.[109]
+Another trend was for very large apocalyptic history paintings, often combining extreme natural events, or divine wrath, with human disaster, attempting to outdo The Raft of the Medusa, and now often drawing comparisons with effects from Hollywood. The leading English artist in the style was John Martin, whose tiny figures were dwarfed by enormous earthquakes and storms, and worked his way through the biblical disasters, and those to come in the final days. Other works such as Delacroix's Death of Sardanapalus included larger figures, and these often drew heavily on earlier artists, especially Poussin and Rubens, with extra emotionalism and special effects.
+Elsewhere in Europe, leading artists adopted Romantic styles: in Russia there were the portraitists Orest Kiprensky and Vasily Tropinin, with Ivan Aivazovsky specializing in marine painting, and in Norway Hans Gude painted scenes of fjords. In Italy Francesco Hayez (1791–1882) was the leading artist of Romanticism in mid-19th-century Milan. His long, prolific and extremely successful career saw him begin as a Neoclassical painter, pass right through the Romantic period, and emerge at the other end as a sentimental painter of young women. His Romantic period included many historical pieces of "Troubadour" tendencies, but on a very large scale, that are heavily influenced by Gian Battista Tiepolo and other late Baroque Italian masters.
+Literary Romanticism had its counterpart in the American visual arts, most especially in the exaltation of an untamed American landscape found in the paintings of the Hudson River School. Painters like Thomas Cole, Albert Bierstadt and Frederic Edwin Church and others often expressed Romantic themes in their paintings. They sometimes depicted ancient ruins of the old world, such as in Fredric Edwin Church's piece Sunrise in Syria. These works reflected the Gothic feelings of death and decay. They also show the Romantic ideal that Nature is powerful and will eventually overcome the transient creations of men. More often, they worked to distinguish themselves from their European counterparts by depicting uniquely American scenes and landscapes. This idea of an American identity in the art world is reflected in W. C. Bryant's poem To Cole, the Painter, Departing for Europe, where Bryant encourages Cole to remember the powerful scenes that can only be found in America.
+Some American paintings (such as Albert Bierstadt's The Rocky Mountains, Lander's Peak) promote the literary idea of the "noble savage" by portraying idealized Native Americans living in harmony with the natural world. Thomas Cole's paintings tend towards allegory, explicit in The Voyage of Life series painted in the early 1840s, showing the stages of life set amidst an awesome and immense nature.
+Thomas Cole, Childhood, one of the four scenes in The Voyage of Life, 1842
+William Blake, Albion Rose, 1794–95
+Louis Janmot, from his series The Poem of the Soul, before 1854
+Thomas Cole, 1842, The Voyage of LifeOld Age
+Musical Romanticism is predominantly a German phenomenon—so much so that one respected French reference work defines it entirely in terms of "The role of music in the aesthetics of German romanticism".[110] Another French encyclopedia holds that the German temperament generally "can be described as the deep and diverse action of romanticism on German musicians", and that there is only one true representative of Romanticism in French music, Hector Berlioz, while in Italy, the sole great name of musical Romanticism is Giuseppe Verdi, "a sort of [Victor] Hugo of opera, gifted with a real genius for dramatic effect". Similarly, in his analysis of Romanticism and its pursuit of harmony, Henri Lefebvre posits that, "But of course, German romanticism was more closely linked to music than French romanticism was, so it is there we should look for the direct expression of harmony as the central romantic idea."[111] Nevertheless, the huge popularity of German Romantic music led, "whether by imitation or by reaction", to an often nationalistically inspired vogue amongst Polish, Hungarian, Russian, Czech, and Scandinavian musicians, successful "perhaps more because of its extra-musical traits than for the actual value of musical works by its masters".[112]
+Although the term "Romanticism" when applied to music has come to imply the period roughly from 1800 until 1850, or else until around 1900, the contemporary application of "romantic" to music did not coincide with this modern interpretation. Indeed, one of the earliest sustained applications of the term to music occurs in 1789, in the Mémoires of André Grétry.[113] This is of particular interest because it is a French source on a subject mainly dominated by Germans, but also because it explicitly acknowledges its debt to Jean-Jacques Rousseau (himself a composer, amongst other things) and, by so doing, establishes a link to one of the major influences on the Romantic movement generally.[114] In 1810 E. T. A. Hoffmann named Haydn, Mozart and Beethoven as "the three masters of instrumental compositions" who "breathe one and the same romantic spirit". He justified his view on the basis of these composers' depth of evocative expression and their marked individuality. In Haydn's music, according to Hoffmann, "a child-like, serene disposition prevails", while Mozart (in the late E-flat major Symphony, for example) "leads us into the depths of the spiritual world", with elements of fear, love, and sorrow, "a presentiment of the infinite ... in the eternal dance of the spheres". Beethoven's music, on the other hand, conveys a sense of "the monstrous and immeasurable", with the pain of an endless longing that "will burst our breasts in a fully coherent concord of all the passions".[115] This elevation in the valuation of pure emotion resulted in the promotion of music from the subordinate position it had held in relation to the verbal and plastic arts during the Enlightenment. Because music was considered to be free of the constraints of reason, imagery, or any other precise concept, it came to be regarded, first in the writings of Wackenroder and Tieck and later by writers such as Schelling and Wagner, as preeminent among the arts, the one best able to express the secrets of the universe, to evoke the spirit world, infinity, and the absolute.[116]
+This chronologic agreement of musical and literary Romanticism continued as far as the middle of the 19th century, when Richard Wagner denigrated the music of Meyerbeer and Berlioz as "neoromantic": "The Opera, to which we shall now return, has swallowed down the Neoromanticism of Berlioz, too, as a plump, fine-flavoured oyster, whose digestion has conferred on it anew a brisk and well-to-do appearance."[117]
+It was only toward the end of the 19th century that the newly emergent discipline of Musikwissenschaft (musicology)—itself a product of the historicizing proclivity of the age—attempted a more scientific periodization of music history, and a distinction between Viennese Classical and Romantic periods was proposed. The key figure in this trend was Guido Adler, who viewed Beethoven and Franz Schubert as transitional but essentially Classical composers, with Romanticism achieving full maturity only in the post-Beethoven generation of Frédéric Chopin, Felix Mendelssohn, Robert Schumann, Hector Berlioz and Franz Liszt. From Adler's viewpoint, found in books like Der Stil in der Musik (1911), composers of the New German School and various late-19th-century nationalist composers were not Romantics but "moderns" or "realists" (by analogy with the fields of painting and literature), and this schema remained prevalent through the first decades of the 20th century.[114]
+By the second quarter of the 20th century, an awareness that radical changes in musical syntax had occurred during the early 1900s caused another shift in historical viewpoint, and the change of century came to be seen as marking a decisive break with the musical past. This in turn led historians such as Alfred Einstein[118] to extend the musical "Romantic era" throughout the 19th century and into the first decade of the 20th. It has continued to be referred to as such in some of the standard music references such as The Oxford Companion to Music[119] and Grout's History of Western Music[120] but was not unchallenged. For example, the prominent German musicologist Friedrich Blume, the chief editor of the first edition of Die Musik in Geschichte und Gegenwart (1949–86), accepted the earlier position that Classicism and Romanticism together constitute a single period beginning in the middle of the 18th century, but at the same time held that it continued into the 20th century, including such pre-World War II developments as expressionism and neoclassicism.[121] This is reflected in some notable recent reference works such as the New Grove Dictionary of Music and Musicians[114] and the new edition of Musik in Geschichte und Gegenwart.[122]
+Felix Mendelssohn, 1839
+Robert Schumann, 1839
+Franz Liszt, 1847
+Daniel Auber, c. 1868
+Hector Berlioz by Gustave Courbet, 1850
+Giovanni Boldini, Portrait of Giuseppe Verdi, 1886
+Richard Wagner, c. 1870s
+Giacomo Meyerbeer, 1847
+In the contemporary music culture, the romantic musician followed a public career depending on sensitive middle-class audiences rather than on a courtly patron, as had been the case with earlier musicians and composers. Public persona characterized a new generation of virtuosi who made their way as soloists, epitomized in the concert tours of Paganini and Liszt, and the conductor began to emerge as an important figure, on whose skill the interpretation of the increasingly complex music depended.[123]
+The Romantic movement affected most aspects of intellectual life, and Romanticism and science had a powerful connection, especially in the period 1800–1840. Many scientists were influenced by versions of the Naturphilosophie of Johann Gottlieb Fichte, Friedrich Wilhelm Joseph von Schelling and Georg Wilhelm Friedrich Hegel and others, and without abandoning empiricism, sought in their work to uncover what they tended to believe was a unified and organic Nature. The English scientist Sir Humphry Davy, a prominent Romantic thinker, said that understanding nature required "an attitude of admiration, love and worship, [...] a personal response".[124] He believed that knowledge was only attainable by those who truly appreciated and respected nature. Self-understanding was an important aspect of Romanticism. It had less to do with proving that man was capable of understanding nature (through his budding intellect) and therefore controlling it, and more to do with the emotional appeal of connecting himself with nature and understanding it through a harmonious co-existence.[125]
+History writing was very strongly, and many would say harmfully, influenced by Romanticism.[126] In England, Thomas Carlyle was a highly influential essayist who turned historian; he both invented and exemplified the phrase "hero-worship",[127] lavishing largely uncritical praise on strong leaders such as Oliver Cromwell, Frederick the Great and Napoleon. Romantic nationalism had a largely negative effect on the writing of history in the 19th century, as each nation tended to produce its own version of history, and the critical attitude, even cynicism, of earlier historians was often replaced by a tendency to create romantic stories with clearly distinguished heroes and villains.[128]   Nationalist ideology of the period placed great emphasis on racial coherence, and the antiquity of peoples, and tended to vastly over-emphasize the continuity between past periods and the present, leading to national mysticism.  Much historical effort in the 20th century was devoted to combating the romantic historical myths created in the 19th century.
+To insulate theology from scientism or reductionism in science, 19th-century post-Enlightenment German theologians developed a modernist or so-called liberal conception of Christianity, led by Friedrich Schleiermacher and Albrecht Ritschl. They took the Romantic approach of rooting religion in the inner world of the human spirit, so that it is a person's feeling or sensibility about spiritual matters that comprises religion.[129]
+Romantic chess was the style of chess which emphasized quick, tactical maneuvers characterized by aesthetic beauty rather than long-term strategic planning, which was considered to be of secondary importance.[130] The Romantic era in chess is generally considered to have begun with Joseph MacDonnell and Pierre LaBourdonnais, the two dominant chess players in the 1830s. The 1840s was dominated by Howard Staunton, and other leading players of the era included Adolf Anderssen, Daniel Harrwitz, Henry Bird, Louis Paulsen, and Paul Morphy. The "Immortal Game", played by Adolf Anderssen and Lionel Kieseritzky on 21 June 1851 in London—where Anderssen made bold sacrifices to secure victory, giving up both rooks and a bishop, then his queen, and then checkmating his opponent with his three remaining minor pieces—is considered a supreme example of Romantic chess.[131] The end of the Romantic era in chess is considered to be the 1873 Vienna Tournament where Wilhelm Steinitz popularized positional play and the closed game.
+One of Romanticism's key ideas and most enduring legacies is the assertion of nationalism, which became a central theme of Romantic art and political philosophy. From the earliest parts of the movement, with their focus on development of national languages and folklore, and the importance of local customs and traditions, to the movements that would redraw the map of Europe and lead to calls for self-determination of nationalities, nationalism was one of the key vehicles of Romanticism, its role, expression and meaning. One of the most important functions of medieval references in the 19th century was nationalist. Popular and epic poetry were its workhorses. This is visible in Germany and Ireland, where underlying Germanic or Celtic linguistic substrates dating from before the Romanization-Latinization were sought out.
+Early Romantic nationalism was strongly inspired by Rousseau, and by the ideas of Johann Gottfried von Herder, who in 1784 argued that the geography formed the natural economy of a people, and shaped their customs and society.[citation needed]
+The nature of nationalism changed dramatically, however, after the French Revolution with the rise of Napoleon, and the reactions in other nations. Napoleonic nationalism and republicanism were, at first, inspirational to movements in other nations: self-determination and a consciousness of national unity were held to be two of the reasons why France was able to defeat other countries in battle. But as the French Republic became Napoleon's Empire, Napoleon became not the inspiration for nationalism, but the object of its struggle. In Prussia, the development of spiritual renewal as a means to engage in the struggle against Napoleon was argued by, among others, Johann Gottlieb Fichte, a disciple of Kant. The word Volkstum, or nationality, was coined in German as part of this resistance to the now conquering emperor. Fichte expressed the unity of language and nation in his address "To the German Nation" in 1806:
+Those who speak the same language are joined to each other by a multitude of invisible bonds by nature herself, long before any human art begins; they understand each other and have the power of continuing to make themselves understood more and more clearly; they belong together and are by nature one and an inseparable whole. ...Only when each people, left to itself, develops and forms itself in accordance with its own peculiar quality, and only when in every people each individual develops himself in accordance with that common quality, as well as in accordance with his own peculiar quality—then, and then only, does the manifestation of divinity appear in its true mirror as it ought to be.[132]
+This view of nationalism inspired the collection of folklore by such people as the Brothers Grimm, the revival of old epics as national, and the construction of new epics as if they were old, as in the Kalevala, compiled from Finnish tales and folklore, or Ossian, where the claimed ancient roots were invented. The view that fairy tales, unless contaminated from outside literary sources, were preserved in the same form over thousands of years, was not exclusive to Romantic Nationalists, but fit in well with their views that such tales expressed the primordial nature of a people. For instance, the Brothers Grimm rejected many tales they collected because of their similarity to tales by Charles Perrault, which they thought proved they were not truly German tales;[133] Sleeping Beauty survived in their collection because the tale of Brynhildr convinced them that the figure of the sleeping princess was authentically German. Vuk Karadžić contributed to Serbian folk literature, using peasant culture as the foundation. He regarded the oral literature of the peasants as an integral part of Serbian culture, compiling it to use in his collections of folk songs, tales and proverbs, as well as the first dictionary of vernacular Serbian.[134] Similar projects were undertaken by the Russian Alexander Afanasyev, the Norwegians Peter Christen Asbjørnsen and Jørgen Moe, and the Englishman Joseph Jacobs.[135]
+Romanticism played an essential role in the national awakening of many Central European peoples lacking their own national states, not least in Poland, which had recently failed to restore its independence when Russia's army crushed the Polish Uprising under Nicholas I. Revival and reinterpretation of ancient myths, customs and traditions by Romantic poets and painters helped to distinguish their indigenous cultures from those of the dominant nations and crystallise the mythography of Romantic nationalism. Patriotism, nationalism, revolution and armed struggle for independence also became popular themes in the arts of this period. Arguably, the most distinguished Romantic poet of this part of Europe was Adam Mickiewicz, who developed an idea that Poland was the Messiah of Nations, predestined to suffer just as Jesus had suffered to save all the people. The Polish self-image as a "Christ among nations" or the martyr of Europe can be traced back to its history of Christendom and suffering under invasions. During the periods of foreign occupation, the Catholic Church served as bastion of Poland's national identity and language, and the major promoter of Polish culture. The partitions came to be seen in Poland as a Polish sacrifice for the security for Western civilization. Adam Mickiewicz wrote the patriotic drama Dziady (directed against the Russians), where he depicts Poland as the Christ of Nations. He also wrote "Verily I say unto you, it is not for you to learn civilization from foreigners, but it is you who are to teach them civilization ... You are among the foreigners like the Apostles among the idolaters". In Books of the Polish Nation and Polish Pilgrimage Mickiewicz detailed his vision of Poland as a Messias and a Christ of Nations, that would save mankind. Dziady is known for various interpretation. The most known ones are the moral aspect of part II, individualist and romantic message of part IV, as well as deeply patriotic, messianistic and Christian vision in part III of the poem. Zdzisław Kępiński, however, focuses his interpretation on Slavic pagan and occult elements found in the drama. In his book Mickiewicz hermetyczny he writes about hermetic, theosophic and alchemical philosophy on the book as well as Masonic symbols.
+Joseph Vernet, 1759, Shipwreck; the 18th-century "sublime"
+Joseph Wright, 1774, Cave at evening, Smith College Museum of Art, Northampton, Massachusetts
+Henry Fuseli, 1781, The Nightmare, a classical artist whose themes often anticipate the Romantic
+Philip James de Loutherbourg, Coalbrookdale by Night, 1801, a key location of the English Industrial Revolution
+Théodore Géricault, The Charging Chasseur, c. 1812
+Ingres, The Death of Leonardo da Vinci, 1818, one of his Troubadour style works
+Eugène Delacroix, Collision of Moorish Horsemen, 1843–44
+Eugène Delacroix, The Bride of Abydos, 1857, after the poem by Byron
+Joseph Anton Koch, Waterfalls at Subiaco 1812–1813, a "classical" landscape to art historians
+James Ward, 1814–1815, Gordale Scar
+John Constable, 1821, The Hay Wain, one of Constable's large "six footers"
+J. C. Dahl, 1826, Eruption of Vesuvius, by Friedrich's closest follower
+William Blake, c. 1824–27, The Wood of the Self-Murderers: The Harpies and the Suicides, Tate
+Karl Bryullov, The Last Day of Pompeii, 1833, The State Russian Museum, St. Petersburg, Russia
+Isaac Levitan, Pacific, 1898, State Russian Museum, St.Petersburg
+J. M. W. Turner, The Burning of the Houses of Lords and Commons (1835), Philadelphia Museum of Art
+Hans Gude, Winter Afternoon, 1847, National Gallery of Norway, Oslo
+Ivan Aivazovsky, 1850, The Ninth Wave, Russian Museum, St. Petersburg
+John Martin, 1852, The Destruction of Sodom and Gomorrah, Laing Art Gallery
+Frederic Edwin Church, 1860, Twilight in the Wilderness, Cleveland Museum of Art
+Albert Bierstadt, 1863, The Rocky Mountains, Lander's Peak

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+The Twelve Labours of Heracles or Hercules (Greek: οἱ Ἡρακλέους ἆθλοι, hoi Hērakléous âthloi)[1][2] are a series of episodes concerning a penance carried out by Heracles, the greatest of the Greek heroes, whose name was later romanised as Hercules. They were accomplished at the service of King Eurystheus. The episodes were later connected by a continuous narrative. The establishment of a fixed cycle of twelve labours was attributed by the Greeks to an epic poem, now lost, written by Peisander, dated about 600 BC.[3] After Heracles killed his wife and children, he went to the oracle at Delphi. He prayed to the god Apollo for guidance. Heracles was told to serve the king of Mycenae, Eurystheus, for ten years. During this time, he is sent to perform a series of difficult feats, called labours.[4]
+Driven mad by Hera  (queen of the gods), Heracles slew his sons by his wife Megara.[5] After recovering his sanity, Heracles deeply regretted his actions; he was purified by King Thespius, then traveled to Delphi to inquire how he could atone for his actions. Pythia, the Oracle of Delphi, advised him to go to Tiryns and serve his cousin, King Eurystheus, for ten years, performing whatever labours Eurystheus might set him; in return, he would be rewarded with immortality. Heracles despaired at this, loathing to serve a man whom he knew to be far inferior to himself, yet fearing to oppose his father Zeus. Eventually, he placed himself at Eurystheus's disposal.
+Eurystheus originally ordered Heracles to perform ten labours. Heracles accomplished these tasks, but Eurystheus refused to recognize two: the slaying of the Lernaean Hydra, as Heracles' nephew and charioteer Iolaus had helped him; and the cleansing of the Augeas, because Heracles accepted payment for the labour. Eurystheus set two more tasks (fetching the Golden Apples of Hesperides and capturing Cerberus), which Heracles also performed, bringing the total number of tasks to twelve.
+As they survive, the labours of Heracles are not recounted in any single place, but must be reassembled from many sources. Ruck and Staples[6] assert that there is no one way to interpret the labours, but that six were located in the Peloponnese, culminating with the rededication of Olympia. Six others took the hero farther afield, to places that were, per Ruck, "all previously strongholds of Hera or the 'Goddess' and were Entrances to the Netherworld".[6] In each case, the pattern was the same: Heracles was sent to kill or subdue, or to fetch back for Eurystheus (as Hera's representative) a magical animal or plant.
+A famous depiction of the labours in Greek sculpture is found on the metopes of the Temple of Zeus at Olympia, which date to the 450s BC.[citation needed]
+In his labours, Heracles was sometimes accompanied by a male companion (an eromenos), according to Licymnius[citation needed] and others, such as Iolaus, his nephew. Although he was supposed to perform only ten labours, this assistance led to two labours being disqualified: Eurystheus refused to recognize slaying the Hydra, because Iolaus helped him, and the cleansing of the Augean stables, because Heracles was paid for his services and/or because the rivers did the work. Several of the labours involved the offspring (by various accounts) of Typhon and his mate Echidna, all overcome by Heracles.
+A traditional order of the labours found in the Bibliotheca[7] by Pseudo-Apollodorus is:
+Heracles wandered the area until he came to the town of Cleonae. There he met a boy who said that if Heracles slew the Nemean lion and returned alive within 30 days, the town would sacrifice a lion to Zeus, but if he did not return within 30 days or if he died, the boy would sacrifice himself to Zeus. Another version claims that he met Molorchos, a shepherd who had lost his son to the lion, saying that if he came back within 30 days, a ram would be sacrificed to Zeus. If he did not return within 30 days, it would be sacrificed to the dead Heracles as a mourning offering.
+While searching for the lion, Heracles fletched some arrows to use against it, not knowing that its golden fur was impenetrable. When he found and shot the lion, firing at it with his bow, he discovered the fur's protective property as the arrow bounced harmlessly off the creature's thigh. After some time, Heracles made the lion return to his cave. The cave had two entrances, one of which Heracles blocked; he then entered the other. In those dark and close quarters, Heracles stunned the beast with his club and, using his immense strength, strangled it to death. During the fight the lion bit off one of his fingers.[8] Others say that he shot arrows at it, eventually shooting it in the unarmored mouth. After slaying the lion, he tried to skin it with a knife from his belt, but failed. He then tried sharpening the knife with a stone and even tried with the stone itself. Finally, Athena, noticing the hero's plight, told Heracles to use one of the lion's own claws to skin the pelt. Others say that Heracles' armor was, in fact, the hide of the Lion of Cithaeron.
+When he returned on the 30th day carrying the carcass of the lion on his shoulders, King Eurystheus was amazed and terrified. Eurystheus forbade him ever again to enter the city; from then on he was to display the fruits of his labours outside the city gates. Eurystheus would then tell Heracles his tasks through a herald, not personally. Eurystheus even had a large bronze jar made for him in which to hide from Heracles if need be. Eurystheus then warned him that the tasks would become increasingly difficult.
+Heracles' second labour was to slay the Lernaean Hydra, which Hera had raised just to slay Heracles. Upon reaching the swamp near Lake Lerna, where the Hydra dwelt, Heracles used a cloth to cover his mouth and nose to protect himself from the poisonous fumes. He fired flaming arrows into the Hydra's lair, the spring of Amymone, a deep cave that it only came out of to terrorize neighboring villages.[9] He then confronted the Hydra, wielding a harvesting sickle (according to some early vase-paintings), a sword or his famed club. Ruck and Staples (1994: 170) have pointed out that the chthonic creature's reaction was botanical: upon cutting off each of its heads he found that two grew back, an expression of the hopelessness of such a struggle for any but the hero. Additionally, one of the Hydra's heads - the middle one - was immortal.
+The details of the struggle are explicit in the Bibliotheca (2.5.2): realizing that he could not defeat the Hydra in this way, Heracles called on his nephew Iolaus for help. His nephew then came upon the idea (possibly inspired by Athena) of using a firebrand to scorch the neck stumps after each decapitation. Heracles cut off each head and Iolaus cauterized the open stumps. Seeing that Heracles was winning the struggle, Hera sent a giant crab to distract him. He crushed it under his mighty foot. He cut off the Hydra's one immortal head with a golden sword given to him by Athena. Heracles placed it under a great rock on the sacred way between Lerna and Elaius (Kerenyi 1959:144), and dipped his arrows in the Hydra's poisonous blood, and so his second task was complete. The alternative version of this myth is that after cutting off one head, he then dipped his sword in it and used its venom to burn each head so it could not grow back. Hera, upset that Heracles had slain the beast she raised to kill him, placed it in the dark blue vault of the sky as the constellation Hydra. She then turned the crab into the constellation Cancer.
+Later, Heracles used an arrow dipped in the Hydra's poisonous blood to kill the centaur Nessus; and Nessus's tainted blood was applied to the Tunic of Nessus, by which the centaur had his posthumous revenge. Both Strabo and Pausanias report that the stench of the river Anigrus in Elis, making all the fish of the river inedible, was reputed to be due to the Hydra's venom, washed from the arrows Heracles used on the centaur.[10]
+Eurystheus and Hera were greatly angered that Heracles had survived the Nemean Lion and the Lernaean Hydra. For the third labour, they found a task which they thought would spell doom for the hero. It was not slaying a beast or monster, as it had already been established that Heracles could overcome even the most fearsome opponents. Instead, Eurystheus ordered him to capture the Ceryneian Hind, which was so fast that it could outrun an arrow.
+After beginning the search, Heracles awoke from sleeping and saw the hind by the glint on its antlers. Heracles then chased the hind on foot for a full year through Greece, Thrace, Istria, and the land of the Hyperboreans. In some versions, he captured the hind while it slept, rendering it lame with a trap net. In other versions, he encountered Artemis in her temple; she told him to leave the hind and tell Eurystheus all that had happened, and his third labour would be considered to be completed. Yet another version claims that Heracles trapped the Hind with an arrow between its forelegs.
+Eurystheus had given Heracles this task hoping to incite Artemis' anger at Heracles for his desecration of her sacred animal. As he was returning with the hind, Heracles encountered Artemis and her brother Apollo. He begged the goddess for forgiveness, explaining that he had to catch it as part of his penance, but he promised to return it. Artemis forgave him, foiling Eurystheus' plan to have her punish him.
+Upon bringing the hind to Eurystheus, he was told that it was to become part of the King's menagerie. Heracles knew that he had to return the hind as he had promised, so he agreed to hand it over on the condition that Eurystheus himself come out and take it from him. The King came out, but the moment that Heracles let the hind go, it sprinted back to its mistress and Heracles left, saying that Eurystheus had not been quick enough.
+Eurystheus was disappointed that Heracles had overcome yet another creature and was humiliated by the hind's escape, so he assigned Heracles another dangerous task. By some accounts, the fourth labour was to bring the fearsome Erymanthian Boar back to Eurystheus alive (there is no single definitive telling of the labours). On the way to Mount Erymanthos where the boar lived, Heracles visited Pholus ("caveman"), a kind and hospitable centaur and old friend. Heracles ate with Pholus in his cavern (though the centaur devoured his meat raw) and asked for wine. Pholus had only one jar of wine, a gift from Dionysus to all the centaurs on Mount Erymanthos. Heracles convinced him to open it, and the smell attracted the other centaurs. They did not understand that wine needs to be tempered with water, became drunk, and attacked Heracles. Heracles shot at them with his poisonous arrows, killing many, and the centaurs retreated all the way to Chiron's cave.
+Pholus was curious why the arrows caused so much death. He picked one up but dropped it, and the arrow stabbed his hoof, poisoning him. One version states that a stray arrow hit Chiron as well. He was immortal, but he still felt the pain. Chiron's pain was so great that he volunteered to give up his immortality and take the place of Prometheus, who had been chained to the top of a mountain to have his liver eaten daily by an eagle. Prometheus' torturer, the eagle, continued its torture on Chiron, so Heracles shot it dead with an arrow. It is generally accepted that the tale was meant to show Heracles as being the recipient of Chiron's surrendered immortality. However, this tale contradicts the fact that Chiron later taught Achilles. The tale of the centaurs sometimes appears in other parts of the twelve labours, as does the freeing of Prometheus.
+Heracles had visited Chiron to gain advice on how to catch the boar, and Chiron had told him to drive it into thick snow, which sets this labour in mid-winter. Heracles caught the boar, bound it, and carried it back to Eurystheus, who was frightened of it and ducked down in his half-buried storage pithos, begging Heracles to get rid of the beast.
+The fifth labour was to clean the stables of King Augeas. This assignment was intended to be both humiliating (rather than impressive, as the previous labours had been) and impossible, since the livestock were divinely healthy (and immortal) and therefore produced an enormous quantity of dung. The Augean Stables (/ɔːˈdʒiːən/) had not been cleaned in over 30 years, and over 1,000 cattle lived there. However, Heracles succeeded by re-routing the rivers Alpheus and Peneus to wash out the filth.
+Before starting on the task, Heracles had asked Augeas for one-tenth of the cattle if he finished the task in one day, and Augeas agreed. But afterwards Augeas refused to honour the agreement on the grounds that Heracles had been ordered to carry out the task by Eurystheus anyway. Heracles claimed his reward in court, and was supported by Augeas' son Phyleus. Augeas banished them both before the court had ruled. Heracles returned, slew Augeas, and gave his kingdom to Phyleus. Heracles then founded the Olympic Games.
+The success of this labour was ultimately discounted as the rushing waters had done the work of cleaning the stables and because Heracles was paid for doing the labour.
+Eurystheus said that Heracles still had seven labours to perform.[11]
+The sixth labour was to defeat the Stymphalian birds, man-eating birds with beaks made of bronze and sharp metallic feathers they could launch at their victims. They were sacred to Ares, the god of war. Furthermore, their dung was highly toxic. They had migrated to Lake Stymphalia in Arcadia, where they bred quickly and took over the countryside, destroying local crops, fruit trees, and townspeople. Heracles could not go too far into the swamp, for it would not support his weight. Athena, noticing the hero's plight, gave Heracles a rattle which Hephaestus had made especially for the occasion. Heracles shook the rattle and frightened the birds into the air. Heracles then shot many of them with his arrows. The rest flew far away, never to return. The Argonauts would later encounter them.
+The seventh labour was to capture the Cretan Bull, father of the Minotaur. Heracles sailed to Crete, where King Minos gave Heracles permission to take the bull away and even offered him assistance (which Heracles declined, plausibly because he did not want the labour to be discounted as before).[12] The bull had been wreaking havoc on Crete by uprooting crops and leveling orchard walls. Heracles sneaked up behind the bull and then used his hands to throttle it (stopping before it was killed), and then shipped it back to Tiryns. Eurystheus, who hid in his pithos at first sight of the creature, wanted to sacrifice the bull to Hera, who hated Heracles. She refused the sacrifice because it reflected glory on Heracles. The bull was released and wandered into Marathon, becoming known as the Marathonian Bull.[12] Theseus would later sacrifice the bull to Athena and/or Apollo.
+As the eighth of his Twelve Labours, also categorised as the second of the Non-Peloponneisan labours,[13] Heracles was sent by King Eurystheus to steal the Mares from Diomedes. The mares’ madness was attributed to their unnatural diet which consisted of the flesh[14] of unsuspecting guests or strangers to the island.[15] Some versions of the myth say that the mares also expelled fire when they breathed.[16] The Mares, which were the terror of Thrace, were kept tethered by iron chains to a bronze manger in the now vanished city of Tirida[17] and were named Podargos (the swift), Lampon (the shining), Xanthos (the yellow) and Deinos (or Deinus, the terrible).[18] Although very similar, there are slight variances in the exact details regarding the mares’ capture.
+In one version, Heracles brought a number of volunteers to help him capture the giant horses.[17] After overpowering Diomedes’ men, Heracles broke the chains that tethered the horses and drove the mares down to sea. Unaware that the mares were man-eating and uncontrollable, Heracles left them in the charge of his favored companion, Abderus, while he left to fight Diomedes. Upon his return, Heracles found that the boy was eaten. As revenge, Heracles fed Diomedes to his own horses and then founded Abdera next to the boy's tomb.[15]
+In another version, Heracles, who was visiting the island, stayed awake so that he didn't have his throat cut by Diomedes in the night, and cut the chains binding the horses once everyone was asleep. Having scared the horses onto the high ground of a knoll, Heracles quickly dug a trench through the peninsula, filling it with water and thus flooding the low-lying plain. When Diomedes and his men turned to flee, Heracles killed them with an axe (or a club[17]), and fed Diomedes’ body to the horses to calm them.
+In yet another version, Heracles first captured Diomedes and fed him to the mares before releasing them. Only after realizing that their King was dead did his men, the Bistonians,[15][17] attack Heracles. Upon seeing the mares charging at them, led in a chariot by Abderus, the Bistonians turned and fled.
+All versions have eating human flesh make the horses calmer, giving Heracles the opportunity to bind their mouths shut, and easily take them back to King Eurystheus, who dedicated the horses to Hera.[19] In some versions, they were allowed to roam freely around Argos, having become permanently calm, but in others, Eurystheus ordered the horses taken to Olympus to be sacrificed to Zeus, but Zeus refused them, and sent wolves, lions, and bears to kill them.[20] Roger Lancelyn Green states in his Tales of the Greek Heroes that the mares’ descendants were used in the Trojan War, and survived even to the time of Alexander the Great.[17][21] After the incident, Eurystheus sent Heracles to bring back Hippolyta's Girdle.
+Eurystheus' daughter Admete wanted the Belt of Hippolyta, queen of the Amazons, a gift from her father Ares. To please his daughter, Eurystheus ordered Heracles to retrieve the belt as his ninth labour.
+Taking a band of friends with him, Heracles set sail, stopping at the island of Paros, which was inhabited by some sons of Minos. The sons killed two of Heracles' companions, an act which set Heracles on a rampage. He killed two of the sons of Minos and threatened the other inhabitants until he was offered two men to replace his fallen companions. Heracles agreed and took two of Minos' grandsons, Alcaeus and Sthenelus. They continued their voyage and landed at the court of Lycus, whom Heracles defended in a battle against King Mygdon of Bebryces. After killing King Mygdon, Heracles gave much of the land to his friend Lycus. Lycus called the land Heraclea. The crew then set off for Themiscyra, where Hippolyta lived.
+All would have gone well for Heracles had it not been for Hera. Hippolyta, impressed with Heracles and his exploits, agreed to give him the belt and would have done so had Hera not disguised herself and walked among the Amazons sowing seeds of distrust. She claimed the strangers were plotting to carry off the queen of the Amazons. Alarmed, the women set off on horseback to confront Heracles. When Heracles saw them, he thought Hippolyta had been plotting such treachery all along and had never meant to hand over the belt, so he killed her, took the belt and returned to Eurystheus.
+The tenth labour was to obtain the Cattle of the three-bodied giant Geryon. In the fullest account in the Bibliotheca of Pseudo-Apollodorus,[22] Heracles had to go to the island of Erytheia in the far west (sometimes identified with the Hesperides, or with the island which forms the city of Cádiz) to get the cattle. On the way there, he crossed the Libyan desert[23] and became so frustrated at the heat that he shot an arrow at the Sun. The sun-god Helios "in admiration of his courage" gave Heracles the golden cup Helios used to sail across the sea from west to east each night. Heracles rode the cup to Erytheia; Heracles in the cup was a favorite motif on black-figure pottery.[citation needed] Such a magical conveyance undercuts any literal geography for Erytheia, the "red island" of the sunset.
+When Heracles landed at Erytheia, he was confronted by the two-headed dog Orthrus. With one blow from his olive-wood club, Heracles killed Orthrus. Eurytion the herdsman came to assist Orthrus, but Heracles dealt with him the same way.
+On hearing the commotion, Geryon sprang into action, carrying three shields and three spears, and wearing three helmets. He attacked Heracles at the River Anthemus, but was slain by one of Heracles' poisoned arrows. Heracles shot so forcefully that the arrow pierced Geryon's forehead, "and Geryon bent his neck over to one side, like a poppy that spoils its delicate shapes, shedding its petals all at once."[24]
+Heracles then had to herd the cattle back to Eurystheus. In Roman versions of the narrative, Heracles drove the cattle over the Aventine Hill on the future site of Rome. The giant Cacus, who lived there, stole some of the cattle as Heracles slept, making the cattle walk backwards so that they left no trail, a repetition of the trick of the young Hermes. According to some versions, Heracles drove his remaining cattle past the cave, where Cacus had hidden the stolen animals, and they began calling out to each other. In other versions, Cacus' sister Caca told Heracles where he was. Heracles then killed Cacus, and set up an altar on the spot, later the site of Rome's Forum Boarium (the cattle market).
+To annoy Heracles, Hera sent a gadfly to bite the cattle, irritate them, and scatter them. Within a year, Heracles retrieved them. Hera then sent a flood which raised the level of a river so much that Heracles could not cross with the cattle. He piled stones into the river to make the water shallower. When he finally reached the court of Eurystheus, the cattle were sacrificed to Hera.
+After Heracles completed the first ten labours, Eurystheus gave him two more, claiming that slaying the Hydra did not count (because Iolaus helped Heracles), neither did cleaning the Augean Stables (either because he was paid for the job or because the rivers did the work).
+The first additional labour was to steal three of the golden apples from the garden of the Hesperides. Heracles first caught the Old Man of the Sea, the shapeshifting sea god,[25] to learn where the Garden of the Hesperides was located.[26]
+In some variations, Heracles, either at the start or at the end of this task, meets Antaeus, who was invincible as long as he touched his mother, Gaia, the Earth. Heracles killed Antaeus by holding him aloft and crushing him in a bear hug.[27]
+Herodotus claims that Heracles stopped in Egypt, where King Busiris decided to make him the yearly sacrifice, but Heracles burst out of his chains.
+Heracles finally made his way to the garden of the Hesperides, where he encountered Atlas holding up the heavens on his shoulders. Heracles persuaded Atlas to get the three golden Apples for him by offering to hold up the heavens in his place for a little while. Atlas could get the apples because, in this version, he was the father or otherwise related to the Hesperides. This would have made the labour – like the Hydra and the Augean stables – void because Heracles had received help. When Atlas returned, he decided that he did not want to take the heavens back, and instead offered to deliver the apples himself, but Heracles tricked him by agreeing to remain in place of Atlas on the condition that Atlas relieve him temporarily while Heracles adjusted his cloak. Atlas agreed, but Heracles reneged and walked away with the apples. According to an alternative version, Heracles slew Ladon, the dragon who guarded the apples instead. Eurystheus was furious that Heracles had accomplished something that Eurystheus thought could not possibly be done.
+The twelfth and final labour was the capture of Cerberus, the three-headed, dragon-tailed dog that was the guardian of the gates of the Underworld. To prepare for his descent into the Underworld, Heracles went to Eleusis (or Athens) to be initiated in the Eleusinian Mysteries. He entered the Underworld, and Hermes and Athena were his guides.
+While in the Underworld, Heracles met Theseus and Pirithous. The two companions had been imprisoned by Hades for attempting to kidnap Persephone. One tradition tells of snakes coiling around their legs, then turning into stone; another that Hades feigned hospitality and prepared a feast inviting them to sit. They unknowingly sat in chairs of forgetfulness and were permanently ensnared. When Heracles had pulled Theseus first from his chair, some of his thigh stuck to it (this explains the supposedly lean thighs of Athenians), but the Earth shook at the attempt to liberate Pirithous, whose desire to have the goddess for himself was so insulting he was doomed to stay behind.
+Heracles found Hades and asked permission to bring Cerberus to the surface, which Hades agreed to if Heracles could subdue the beast without using weapons. Heracles overpowered Cerberus with his bare hands and slung the beast over his back. He carried Cerberus out of the Underworld through a cavern entrance in the Peloponnese and brought it to Eurystheus, who again fled into his pithos.  Eurystheus begged Heracles to return Cerberus to the Underworld, offering in return to release him from any further labours when Cerberus disappeared back to his master.
+After completing the Twelve Labours, one tradition says Heracles joined Jason and the Argonauts in their quest for the Golden Fleece. However, Herodorus (c. 400 BC) disputed this and denied Heracles ever sailed with the Argonauts. A separate tradition (e.g. Argonautica) has Heracles accompany the Argonauts, but he did not travel with them as far as Colchis.
+Some ancient Greeks found allegorical meanings of a moral, psychological or philosophical nature in the Labours of Heracles.  This trend became more prominent in the Renaissance.[28]  For example, Heraclitus the Grammarian wrote in his Homeric Problems:
+I turn to Heracles. We must not suppose he attained such power in those days as a result of his physical strength. Rather, he was a man of intellect, an initiate in heavenly wisdom, who, as it were, shed light on philosophy, which had been hidden in deep darkness. The most authoritative of the Stoics agree with this account.... The (Erymanthian) boar which he overcame is the common incontinence of men; the (Nemean) lion is the indiscriminate rush towards improper goals; in the same way, by fettering irrational passions he gave rise to the belief that he had fettered the violent (Cretan) bull. He banished cowardice also from the world, in the shape of the hind of Ceryneia. There was another "labor" too, not properly so called, in which he cleared out the mass of dung (from the Augean stables) — in other words, the foulness that disfigures humanity. The (Stymphalian) birds he scattered are the windy hopes that feed our lives; the many-headed hydra that he burned, as it were, with the fires of exhortation, is pleasure, which begins to grow again as soon as it is cut out.

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+Apiformes (from Latin 'apis')
+Bees are flying insects closely related to wasps and ants, known for their role in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamily Apoidea. They are presently considered a clade, called Anthophila. There are over 16,000 known species of bees in seven recognized biological families.[1][2] Some species — including honey bees, bumblebees, and stingless bees — live socially in colonies while some species — including mason bees, carpenter bees, leafcutter bees, and sweat bees — are solitary.
+Bees are found on every continent except for Antarctica, in every habitat on the planet that contains insect-pollinated flowering plants. The most common bees in the Northern Hemisphere are the Halictidae, or sweat bees, but they are small and often mistaken for wasps or flies. Bees range in size from tiny stingless bee species, whose workers are less than 2 millimetres (0.08 in) long, to Megachile pluto, the largest species of leafcutter bee, whose females can attain a length of 39 millimetres (1.54 in).
+Bees feed on nectar and pollen, the former primarily as an energy source and the latter primarily for protein and other nutrients. Most pollen is used as food for their larvae. Vertebrate predators of bees include birds such as bee-eaters; insect predators include beewolves and dragonflies.
+Bee pollination is important both ecologically and commercially, and the decline in wild bees has increased the value of pollination by commercially managed hives of honey bees. The analysis of 353 wild bee and hoverfly species across Britain from 1980 to 2013 found the insects have been lost from a quarter of the places they inhabited in 1980.[3]
+Human beekeeping or apiculture has been practised for millennia, since at least the times of Ancient Egypt and Ancient Greece. Bees have appeared in mythology and folklore, through all phases of art and literature from ancient times to the present day, although primarily focused in the Northern Hemisphere where beekeeping is far more common.
+The ancestors of bees were wasps in the family Crabronidae, which were predators of other insects. The switch from insect prey to pollen may have resulted from the consumption of prey insects which were flower visitors and were partially covered with pollen when they were fed to the wasp larvae. This same evolutionary scenario may have occurred within the vespoid wasps, where the pollen wasps evolved from predatory ancestors. Until recently, the oldest non-compression bee fossil had been found in New Jersey amber, Cretotrigona prisca of Cretaceous age, a corbiculate bee.[4] A bee fossil from the early Cretaceous (~100 mya), Melittosphex burmensis, is considered "an extinct lineage of pollen-collecting Apoidea sister to the modern bees".[5] Derived features of its morphology (apomorphies) place it clearly within the bees, but it retains two unmodified ancestral traits (plesiomorphies) of the legs (two mid-tibial spurs, and a slender hind basitarsus), showing its transitional status.[5] By the Eocene (~45 mya) there was already considerable diversity among eusocial bee lineages.[6][a]
+The highly eusocial corbiculate Apidae appeared roughly 87 Mya, and the Allodapini (within the Apidae) around 53 Mya.[9]
+The Colletidae appear as fossils only from the late Oligocene (~25 Mya) to early Miocene.[10]
+The Melittidae are known from Palaeomacropis eocenicus in the Early Eocene.[11]
+The Megachilidae are known from trace fossils (characteristic leaf cuttings) from the Middle Eocene.[12]
+The Andrenidae are known from the Eocene-Oligocene boundary, around 34 Mya, of the Florissant shale.[13]
+The Halictidae first appear in the Early Eocene[14] with species[15][16] found in amber. The Stenotritidae are known from fossil brood cells of Pleistocene age.[17]
+The earliest animal-pollinated flowers were shallow, cup-shaped blooms pollinated by insects such as beetles, so the syndrome of insect pollination was well established before the first appearance of bees. The novelty is that bees are specialized as pollination agents, with behavioral and physical modifications that specifically enhance pollination, and are the most efficient pollinating insects. In a process of coevolution, flowers developed floral rewards[18] such as nectar and longer tubes, and bees developed longer tongues to extract the nectar.[19] Bees also developed structures known as scopal hairs and pollen baskets to collect and carry pollen. The location and type differ among and between groups of bees. Most species have scopal hairs on their hind legs or on the underside of their abdomens. Some species in the family Apidae have pollen baskets on their hind legs, while very few lack these and instead collect pollen in their crops.[2] The appearance of these structures drove the adaptive radiation of the angiosperms, and, in turn, bees themselves.[7] Bees coevolved not only with flowers but it is believed that some species coevolved with mites. Some provide tufts of hairs called acarinaria that appear to provide lodgings for mites; in return, it is believed that mites eat fungi that attack pollen, so the relationship in this case may be mutualistc.[20][21]
+This phylogenetic tree is based on Debevic et al, 2012, which used molecular phylogeny to demonstrate that the bees (Anthophila) arose from deep within the Crabronidae, which is therefore paraphyletic. The placement of the Heterogynaidae is uncertain.[22] The small subfamily Mellininae was not included in this analysis.
+Ampulicidae (Cockroach wasps)
+Heterogynaidae (possible placement #1)
+Sphecidae (sensu stricto)
+Crabroninae (part of "Crabronidae")
+Bembicini
+Nyssonini, Astatinae
+Heterogynaidae (possible placement #2)
+Pemphredoninae, Philanthinae
+Anthophila (bees)
+This cladogram of the bee families is based on Hedtke et al., 2013, which places the former families Dasypodaidae and Meganomiidae as subfamilies inside the Melittidae.[23] English names, where available, are given in parentheses.
+Melittidae (inc. Dasypodainae, Meganomiinae) at least 50 Mya
+Apidae (inc. honeybees, cuckoo bees, carpenter bees) ≈87 Mya
+Megachilidae (mason, leafcutter bees) ≈50 Mya
+Andrenidae (mining bees) ≈34 Mya
+Halictidae (sweat bees) ≈50 Mya
+Colletidae (plasterer bees) ≈25 Mya
+Stenotritidae (large Australian bees) ≈2 Mya
+Bees differ from closely related groups such as wasps by having branched or plume-like setae (hairs), combs on the forelimbs for cleaning their antennae, small anatomical differences in limb structure, and the venation of the hind wings; and in females, by having the seventh dorsal abdominal plate divided into two half-plates.[24]
+Bees have the following characteristics:
+The largest species of bee is thought to be Wallace's giant bee Megachile pluto, whose females can attain a length of 39 millimetres (1.54 in).[26] The smallest species may be dwarf stingless bees in the tribe Meliponini whose workers are less than 2 millimetres (0.08 in) in length.[27]
+According to inclusive fitness theory, organisms can gain fitness not just through increasing their own reproductive output, but also that of close relatives. In evolutionary terms, individuals should help relatives when Cost < Relatedness * Benefit. The requirements for eusociality are more easily fulfilled by haplodiploid species such as bees because of their unusual relatedness structure.[28]
+In haplodiploid species, females develop from fertilized eggs and males from unfertilized eggs. Because a male is haploid (has only one copy of each gene), his daughters (which are diploid, with two copies of each gene) share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise to what W. D. Hamilton termed "supersisters", more closely related to their sisters than they would be to their own offspring.[29] Workers often do not reproduce, but they can pass on more of their genes by helping to raise their sisters (as queens) than they would by having their own offspring (each of which would only have 50% of their genes), assuming they would produce similar numbers. This unusual situation has been proposed as an explanation of the multiple (at least 9) evolutions of eusociality within Hymenoptera.[30][31]
+Haplodiploidy is neither necessary nor sufficient for eusociality. Some eusocial species such as termites are not haplodiploid. Conversely, all bees are haplodiploid but not all are eusocial, and among eusocial species many queens mate with multiple males, creating half-sisters that share only 25% of each-other's genes.[32] But, monogamy (queens mating singly) is the ancestral state for all eusocial species so far investigated, so it is likely that haplodiploidy contributed to the evolution of eusociality in bees.[30]
+Bees may be solitary or may live in various types of communities. Eusociality appears to have originated from at least three independent origins in halictid bees.[33] The most advanced of these are species with eusocial colonies; these are characterised by cooperative brood care and a division of labour into reproductive and non-reproductive adults, plus overlapping generations.[34] This division of labour creates specialized groups within eusocial societies which are called castes. In some species, groups of cohabiting females may be sisters, and if there is a division of labour within the group, they are considered semisocial. The group is called eusocial if, in addition, the group consists of a mother (the queen) and her daughters (workers). When the castes are purely behavioural alternatives, with no morphological differentiation other than size, the system is considered primitively eusocial, as in many paper wasps; when the castes are morphologically discrete, the system is considered highly eusocial.[19]
+True honey bees (genus Apis, of which seven species are currently recognized) are highly eusocial, and are among the best known insects. Their colonies are established by swarms, consisting of a queen and several hundred workers. There are 29 subspecies of one of these species, Apis mellifera, native to Europe, the Middle East, and Africa. Africanized bees are a hybrid strain of A. mellifera that escaped from experiments involving crossing European and African subspecies; they are extremely defensive.[35]
+Stingless bees are also highly eusocial. They practise mass provisioning, with complex nest architecture and perennial colonies also established via swarming.[36]
+Many bumblebees are eusocial, similar to the eusocial Vespidae such as hornets in that the queen initiates a nest on her own rather than by swarming. Bumblebee colonies typically have from 50 to 200 bees at peak population, which occurs in mid to late summer. Nest architecture is simple, limited by the size of the pre-existing nest cavity, and colonies rarely last more than a year.[37]  In 2011, the International Union for Conservation of Nature set up the Bumblebee Specialist Group to review the threat status of all bumblebee species worldwide using the IUCN Red List criteria.[38]
+There are many more species of primitively eusocial than highly eusocial bees, but they have been studied less often. Most are in the family Halictidae, or "sweat bees". Colonies are typically small, with a dozen or fewer workers, on average. Queens and workers differ only in size, if at all. Most species have a single season colony cycle, even in the tropics, and only mated females hibernate. A few species have long active seasons and attain colony sizes in the hundreds, such as Halictus hesperus.[39] Some species are eusocial in parts of their range and solitary in others,[40] or have a mix of eusocial and solitary nests in the same population.[41] The orchid bees (Apidae) include some primitively eusocial species with similar biology. Some allodapine bees (Apidae) form primitively eusocial colonies, with progressive provisioning: a larva's food is supplied gradually as it develops, as is the case in honey bees and some bumblebees.[42]
+Most other bees, including familiar insects such as carpenter bees, leafcutter bees and mason bees are solitary in the sense that every female is fertile, and typically inhabits a nest she constructs herself. There is no division of labor so these nests lack queens and worker bees for these species. Solitary bees typically produce neither honey nor beeswax.
+Bees collect pollen to feed their young, and have the necessary adaptations to do this. However, certain wasp species such as pollen wasps have similar behaviours, and a few species of bee scavenge from carcases to feed their offspring.[24] Solitary bees are important pollinators; they gather pollen to provision their nests with food for their brood. Often it is mixed with nectar to form a paste-like consistency. Some solitary bees have advanced types of pollen-carrying structures on their bodies. Very few species of solitary bee are being cultured for commercial pollination. Most of these species belong to a distinct set of genera which are commonly known by their nesting behavior or preferences, namely: carpenter bees, sweat bees, mason bees, plasterer bees, squash bees, dwarf carpenter bees, leafcutter bees, alkali bees and digger bees.[43]
+Most solitary bees nest in the ground in a variety of soil textures and conditions while others create nests in hollow reeds or twigs, holes in wood. The female typically creates a compartment (a "cell") with an egg and some provisions for the resulting larva, then seals it off. A nest may consist of numerous cells. When the nest is in wood, usually the last (those closer to the entrance) contain eggs that will become males. The adult does not provide care for the brood once the egg is laid, and usually dies after making one or more nests. The males typically emerge first and are ready for mating when the females emerge. Solitary bees are either stingless or very unlikely to sting (only in self-defense, if ever).[44][45]
+While solitary, females each make individual nests. Some species, such as the European mason bee Hoplitis anthocopoides,[46] and the Dawson's Burrowing bee, Amegilla dawsoni,[47] are gregarious, preferring to make nests near others of the same species, and giving the appearance of being social. Large groups of solitary bee nests are called aggregations, to distinguish them from colonies. In some species, multiple females share a common nest, but each makes and provisions her own cells independently. This type of group is called "communal" and is not uncommon. The primary advantage appears to be that a nest entrance is easier to defend from predators and parasites when there are multiple females using that same entrance on a regular basis.[46]
+The life cycle of a bee, be it a solitary or social species, involves the laying of an egg, the development through several moults of a legless larva, a pupation stage during which the insect undergoes complete metamorphosis, followed by the emergence of a winged adult. Most solitary bees and bumble bees in temperate climates overwinter as adults or pupae and emerge in spring when increasing numbers of flowering plants come into bloom. The males usually emerge first and search for females with which to mate. The sex of a bee is determined by whether or not the egg is fertilised; after mating, a female stores the sperm, and determines which sex is required at the time each individual egg is laid, fertilised eggs producing female offspring and unfertilised eggs, males. Tropical bees may have several generations in a year and no diapause stage.[48][49][50][51]
+The egg is generally oblong, slightly curved and tapering at one end. Solitary bees, lay each egg in a separate cell with a supply of mixed pollen and nectar next to it. This may be rolled into a pellet or placed in a pile and is known as mass provisioning. Social bee species provision progressively, that is, they feed the larva regularly while it grows. The nest varies from a hole in the ground or in wood, in solitary bees, to a substantial structure with wax combs in bumblebees and honey bees.[52]
+In most species, larvae are whitish grubs, roughly oval and bluntly-pointed at both ends. They have 15 segments and spiracles in each segment for breathing. They have no legs but move within the cell, helped by tubercles on their sides. They have short horns on the head, jaws for chewing food and an appendage on either side of the mouth tipped with a bristle. There is a gland under the mouth that secretes a viscous liquid which solidifies into the silk they use to produce a cocoon. The cocoon is semi-transparent and the pupa can be seen through it. Over the course of a few days, the larva undergoes metamorphosis into a winged adult. When ready to emerge, the adult splits its skin dorsally and climbs out of the exuviae and breaks out of the cell.[52]
+Nest of common carder bumblebee, wax canopy removed to show winged workers and pupae in irregularly placed wax cells
+Carpenter bee nests in a cedar wood beam (sawn open)
+Honeybees on brood comb with eggs and larvae in cells
+Antoine Magnan's 1934 book Le vol des insectes, says that he and André Sainte-Laguë had applied the equations of air resistance to insects and found that their flight could not be explained by fixed-wing calculations, but that "One shouldn't be surprised that the results of the calculations don't square with reality".[53] This has led to a common misconception that bees "violate aerodynamic theory". In fact it merely confirms that bees do not engage in fixed-wing flight, and that their flight is explained by other mechanics, such as those used by helicopters.[54] In 1996 it was shown that vortices created by many insects' wings helped to provide lift.[55] High-speed cinematography[56] and robotic mock-up of a bee wing[57] showed that lift was generated by "the unconventional combination of short, choppy wing strokes, a rapid rotation of the wing as it flops over and reverses direction, and a very fast wing-beat frequency". Wing-beat frequency normally increases as size decreases, but as the bee's wing beat covers such a small arc, it flaps approximately 230 times per second, faster than a fruitfly (200 times per second) which is 80 times smaller.[58]
+The ethologist Karl von Frisch studied navigation in the honey bee. He showed that honey bees communicate by the waggle dance, in which a worker indicates the location of a food source to other workers in the hive. He demonstrated that bees can recognize a desired compass direction in three different ways: by the sun, by the polarization pattern of the blue sky, and by the earth's magnetic field. He showed that the sun is the preferred or main compass; the other mechanisms are used under cloudy skies or inside a dark beehive.[59] Bees navigate using spatial memory with a "rich, map-like organization".[60]
+The gut of bees is relatively simple, but multiple metabolic strategies exist in the gut microbiota.[61] Pollinating bees consume nectar and pollen, which require different digestion strategies by somewhat specialized bacteria. While nectar is a liquid of mostly monosaccharide sugars and so easily absorbed, pollen contains complex polysaccharides: branching pectin and hemicellulose.[62] Approximately five groups of bacteria are involved in digestion. Three groups specialize in simple sugars (Snodgrassella and two groups of Lactobacillus), and two other groups in complex sugars (Gilliamella and Bifidobacterium). Digestion of pectin and hemicellulose is dominated by bacterial clades Gilliamella and Bifidobacterium respectively. Bacteria that cannot digest polysaccharides obtain enzymes from their neighbors, and bacteria that lack certain amino acids do the same, creating multiple ecological niches.[63]
+Although most bee species are nectarivorous and palynivorous, some are not. Particularly unusual are vulture bees in the genus Trigona, which consume carrion and wasp brood, turning meat into a honey-like substance.[64]
+Most bees are polylectic (generalist) meaning they collect pollen from a range of flowering plants, but some are oligoleges (specialists), in that they only gather pollen from one or a few species or genera of closely related plants.[65] Specialist pollinators also include bee species which gather floral oils instead of pollen, and male orchid bees, which gather aromatic compounds from orchids (one of the few cases where male bees are effective pollinators). Bees are able to sense the presence of desirable flowers through ultraviolet patterning on flowers, floral odors,[66] and even electromagnetic fields.[67] Once landed, a bee then uses nectar quality[66] and pollen taste[68] to determine whether to continue visiting similar flowers.
+In rare cases, a plant species may only be effectively pollinated by a single bee species, and some plants are endangered at least in part because their pollinator is also threatened. But, there is a pronounced tendency for oligolectic bees to be associated with common, widespread plants visited by multiple pollinator species. For example, the creosote bush in the arid parts of the United States southwest is associated with some 40 oligoleges.[69]
+Many bees are aposematically coloured, typically orange and black, warning of their ability to defend themselves with a powerful sting. As such they are models for Batesian mimicry by non-stinging insects such as bee-flies, robber flies and hoverflies,[70] all of which gain a measure of protection by superficially looking and behaving like bees.[70]
+Bees are themselves Müllerian mimics of other aposematic insects with the same colour scheme, including wasps, lycid and other beetles, and many butterflies and moths (Lepidoptera) which are themselves distasteful, often through acquiring bitter and poisonous chemicals from their plant food. All the Müllerian mimics, including bees, benefit from the reduced risk of predation that results from their easily recognised warning coloration.[71]
+Bees are also mimicked by plants such as the bee orchid which imitates both the appearance and the scent of a female bee; male bees attempt to mate (pseudocopulation) with the furry lip of the flower, thus pollinating it.[72]
+Brood parasites occur in several bee families including the apid subfamily Nomadinae.[73] Females of these species lack pollen collecting structures (the scopa) and do not construct their own nests. They typically enter the nests of pollen collecting species, and lay their eggs in cells provisioned by the host bee. When the "cuckoo" bee larva hatches, it consumes the host larva's pollen ball, and often the host egg also.[74] In particular, the Arctic bee species, Bombus hyperboreus is an aggressive species that attacks and enslaves other bees of the same subgenus. However, unlike many other bee brood parasites, they have pollen baskets and often collect pollen.[75]
+In Southern Africa, hives of African honeybees (A. mellifera scutellata) are being destroyed by parasitic workers of the Cape honeybee, A. m. capensis. These lay diploid eggs ("thelytoky"), escaping normal worker policing, leading to the colony's destruction; the parasites can then move to other hives.[76]
+The cuckoo bees in the Bombus subgenus Psithyrus are closely related to, and resemble, their hosts in looks and size. This common pattern gave rise to the ecological principle "Emery's rule". Others parasitize bees in different families, like Townsendiella, a nomadine apid, two species of which are cleptoparasites of the dasypodaid genus Hesperapis,[77] while the other species in the same genus attacks halictid bees.[78]
+Four bee families (Andrenidae, Colletidae, Halictidae, and Apidae) contain some species that are crepuscular. Most are tropical or subtropical, but some live in arid regions at higher latitudes. These bees have greatly enlarged ocelli, which are extremely sensitive to light and dark, though incapable of forming images. Some have refracting superposition compound eyes: these combine the output of many elements of their compound eyes to provide enough light for each retinal photoreceptor. Their ability to fly by night enables them to avoid many predators, and to exploit flowers that produce nectar only or also at night.[79]
+Vertebrate predators of bees include bee-eaters, shrikes and flycatchers, which make short sallies to catch insects in flight.[80] Swifts and swallows[80] fly almost continually, catching insects as they go. The honey buzzard attacks bees' nests and eats the larvae.[81] The greater honeyguide interacts with humans by guiding them to the nests of wild bees. The humans break open the nests and take the honey and the bird feeds on the larvae and the wax.[82] Among mammals, predators such as the badger dig up bumblebee nests and eat both the larvae and any stored food.[83]
+Specialist ambush predators of visitors to flowers include crab spiders, which wait on flowering plants for pollinating insects; predatory bugs, and praying mantises,[80] some of which (the flower mantises of the tropics) wait motionless, aggressive mimics camouflaged as flowers.[84] Beewolves are large wasps that habitually attack bees;[80] the ethologist Niko Tinbergen estimated that a single colony of the beewolf Philanthus triangulum might kill several thousand honeybees in a day: all the prey he observed were honeybees.[85] Other predatory insects that sometimes catch bees include robber flies and dragonflies.[80] Honey bees are affected by parasites including acarine and Varroa mites.[86] However, some bees are believed to have a mutualistic relationship with mites.[21]
+Homer's Hymn to Hermes describes three bee-maidens with the power of divination and thus speaking truth, and identifies the food of the gods as honey. Sources associated the bee maidens with Apollo and, until the 1980s, scholars followed Gottfried Hermann (1806) in incorrectly identifying the bee-maidens with the Thriae.[87] Honey, according to a Greek myth, was discovered by a nymph called Melissa ("Bee"); and honey was offered to the Greek gods from Mycenean times. Bees were also associated with the Delphic oracle and the prophetess was sometimes called a bee.[88]
+The image of a community of honey bees has been used from ancient to modern times, in Aristotle and Plato; in Virgil and Seneca; in Erasmus and Shakespeare; Tolstoy, and by political and social theorists such as Bernard Mandeville and Karl Marx as a model for human society.[89] In English folklore, bees would be told of important events in the household, in a custom known as "Telling the bees".[90]
+Some of the oldest examples of bees in art are rock paintings in Spain which have been dated to 15,000 BC.[91]
+W. B. Yeats's poem The Lake Isle of Innisfree (1888) contains the couplet "Nine bean rows will I have there, a hive for the honey bee, / And live alone in the bee loud glade." At the time he was living in Bedford Park in the West of London.[92] Beatrix Potter's illustrated book The Tale of Mrs Tittlemouse (1910) features Babbity Bumble and her brood (pictured). Kit Williams' treasure hunt book The Bee on the Comb (1984) uses bees and beekeeping as part of its story and puzzle. Sue Monk Kidd's The Secret Life of Bees (2004), and the 2009 film starring Dakota Fanning, tells the story of a girl who escapes her abusive home and finds her way to live with a family of beekeepers, the Boatwrights.
+The humorous 2007 animated film Bee Movie used Jerry Seinfeld's first script and was his first work for children; he starred as a bee named Barry B. Benson, alongside Renée Zellweger. Critics found its premise awkward and its delivery tame.[93] Dave Goulson's A Sting in the Tale (2014) describes his efforts to save bumblebees in Britain, as well as much about their biology. The playwright Laline Paull's fantasy The Bees (2015) tells the tale of a hive bee named Flora 717 from hatching onwards.[94]
+Humans have kept honey bee colonies, commonly in hives, for millennia. Beekeepers collect honey, beeswax, propolis, pollen, and royal jelly from hives; bees are also kept to pollinate crops and to produce bees for sale to other beekeepers.
+Depictions of humans collecting honey from wild bees date to 15,000 years ago; efforts to domesticate them are shown in Egyptian art around 4,500 years ago.[95] Simple hives and smoke were used;[96][97] jars of honey were found in the tombs of pharaohs such as Tutankhamun. From the 18th century, European understanding of the colonies and biology of bees allowed the construction of the moveable comb hive so that honey could be harvested without destroying the colony.[98][99] Among Classical Era authors, beekeeping with the use of smoke is described in Aristotle's History of Animals Book 9.[100] The account mentions that bees die after stinging; that workers remove corpses from the hive, and guard it; castes including workers and non-working drones, but "kings" rather than queens; predators including toads and bee-eaters; and the waggle dance, with the "irresistible suggestion" of άpοσειονται ("aroseiontai", it waggles) and παρακολουθούσιν ("parakolouthousin", they watch).[101][b]
+Beekeeping is described in detail by Virgil in his Eclogues; it is also mentioned in his Aeneid, and in Pliny's Natural History.[101]
+Bees play an important role in pollinating flowering plants, and are the major type of pollinator in many ecosystems that contain flowering plants. It is estimated that one third of the human food supply depends on pollination by insects, birds and bats, most of which is accomplished by bees, whether wild or domesticated.[102][103] Over the last half century, there has been a general decline in the species richness of wild bees and other pollinators, probably attributable to stress from increased parasites and disease, the use of pesticides, and a general decrease in the number of wild flowers. Climate change probably exacerbates the problem.[104]
+Contract pollination has overtaken the role of honey production for beekeepers in many countries. After the introduction of Varroa mites, feral honey bees declined dramatically in the US, though their numbers have since recovered.[105][106] The number of colonies kept by beekeepers declined slightly, through urbanization, systematic pesticide use, tracheal and Varroa mites, and the closure of beekeeping businesses. In 2006 and 2007 the rate of attrition increased, and was described as colony collapse disorder.[107] In 2010 invertebrate iridescent virus and the fungus Nosema ceranae were shown to be in every killed colony, and deadly in combination.[108][109][110][111] Winter losses increased to about 1/3.[112][113] Varroa mites were thought to be responsible for about half the losses.[114]
+Apart from colony collapse disorder, losses outside the US have been attributed to causes including pesticide seed dressings, using neonicotinoids such as Clothianidin, Imidacloprid and Thiamethoxam.[115][116] From 2013 the European Union restricted some pesticides to stop bee populations from declining further.[117] In 2014 the Intergovernmental Panel on Climate Change report warned that bees faced increased risk of extinction because of global warming.[118] In 2018 the European Union decided to ban field use of all three major neonicotinoids; they remain permitted in veterinary, greenhouse, and vehicle transport usage.[119]
+Farmers have focused on alternative solutions to mitigate these problems. By raising native plants, they provide food for native bee pollinators like Lasioglossum vierecki[120] and L. leucozonium,[121] leading to less reliance on honey bee populations.
+Honey is a natural product produced by bees and stored for their own use, but its sweetness has always appealed to humans. Before domestication of bees was even attempted, humans were raiding their nests for their honey. Smoke was often used to subdue the bees and such activities are depicted in rock paintings in Spain dated to 15,000 BC.[91]
+Honey bees are used commercially to produce honey.[122] They also produce some substances used as dietary supplements with possible health benefits, pollen,[123] propolis,[124] and royal jelly,[125] though all of these can also cause allergic reactions.
+Bees are partly considered edible insects. Indigenous people in many countries eat insects, including the larvae and pupae of bees, mostly stingless species. They also gather larvae, pupae and surrounding cells, known as bee brood, for consumption.[126] In the Indonesian dish botok tawon from Central and East Java, bee larvae are eaten as a companion to rice, after being mixed with shredded coconut,  wrapped in banana leaves, and steamed.[127][128]
+Bee brood (pupae and larvae) although low in calcium, has been found to be high in protein  and carbohydrate, and a useful source of phosphorus, magnesium, potassium, and trace minerals iron, zinc, copper, and selenium. In addition, while bee brood was high in fat, it contained no fat soluble vitamins (such as A, D, and E) but it was a good source of most of the water-soluble B-vitamins including choline as well as vitamin C. The fat was composed mostly of saturated and monounsaturated fatty acids with 2.0% being polyunsaturated fatty acids.[129][130]
+Apitherapy is a branch of alternative medicine that uses honey bee products, including raw honey, royal jelly, pollen, propolis, beeswax and apitoxin (Bee venom).[131] The claim that apitherapy treats cancer, which some proponents of apitherapy make, remains unsupported by evidence-based medicine.[132][133]
+The painful stings of bees are mostly associated with the poison gland and the Dufour's gland which are abdominal exocrine glands containing various chemicals. In Lasioglossum leucozonium, the Dufour's Gland mostly contains octadecanolide as well as some eicosanolide. There is also evidence of n-triscosane, n-heptacosane,[134] and 22-docosanolide.[135] However, the secretions of these glands could also be used for nest construction.[134]

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+Blood is a body fluid in humans and other animals that delivers necessary substances such as nutrients and oxygen to the cells and transports metabolic waste products away from those same cells.[1]
+In vertebrates, it is composed of blood cells suspended in blood plasma. Plasma, which constitutes 55% of blood fluid, is mostly water (92% by volume),[2] and contains proteins, glucose, mineral ions, hormones, carbon dioxide (plasma being the main medium for excretory product transportation), and blood cells themselves. Albumin is the main protein in plasma, and it functions to regulate the colloidal osmotic pressure of blood. The blood cells are mainly red blood cells (also called RBCs or erythrocytes), white blood cells (also called WBCs or leukocytes) and platelets (also called thrombocytes). The most abundant cells in vertebrate blood are red blood cells. These contain hemoglobin, an iron-containing protein, which facilitates oxygen transport by reversibly binding to this respiratory gas and greatly increasing its solubility in blood. In contrast, carbon dioxide is mostly transported extracellularly as bicarbonate ion transported in plasma.
+Vertebrate blood is bright red when its hemoglobin is oxygenated and dark red when it is deoxygenated. Some animals, such as crustaceans and mollusks, use hemocyanin to carry oxygen, instead of hemoglobin. Insects and some mollusks use a fluid called hemolymph instead of blood, the difference being that hemolymph is not contained in a closed circulatory system. In most insects, this "blood" does not contain oxygen-carrying molecules such as hemoglobin because their bodies are small enough for their tracheal system to suffice for supplying oxygen.
+Jawed vertebrates have an adaptive immune system, based largely on white blood cells. White blood cells help to resist infections and parasites. Platelets are important in the clotting of blood. Arthropods, using hemolymph, have hemocytes as part of their immune system.
+Blood is circulated around the body through blood vessels by the pumping action of the heart. In animals with lungs, arterial blood carries oxygen from inhaled air to the tissues of the body, and venous blood carries carbon dioxide, a waste product of metabolism produced by cells, from the tissues to the lungs to be exhaled.
+Medical terms related to blood often begin with hemo- or hemato- (also spelled haemo- and haemato-) from the Greek word αἷμα (haima) for "blood". In terms of anatomy and histology, blood is considered a specialized form of connective tissue, given its origin in the bones and the presence of potential molecular fibers in the form of fibrinogen.
+Blood performs many important functions within the body, including:
+Blood accounts for 7% of the human body weight,[3][4] with an average density around 1060 kg/m3, very close to pure water's density of 1000 kg/m3.[5] The average adult has a blood volume of roughly 5 litres (11 US pt) or 1.3 gallons,[4] which is composed of plasma and several kinds of cells. These blood cells (which are also called corpuscles or "formed elements") consist of erythrocytes (red blood cells, RBCs), leukocytes (white blood cells), and thrombocytes (platelets). By volume, the red blood cells constitute about 45% of whole blood, the plasma about 54.3%, and white cells about 0.7%.
+Whole blood (plasma and cells) exhibits non-Newtonian fluid dynamics.[specify]
+Human blood fractioned by centrifugation: Plasma (upper, yellow layer), buffy coat (middle, thin white layer) and erythrocyte layer (bottom, red layer) can be seen.
+Blood circulation: Red = oxygenated, blue = deoxygenated
+Illustration depicting formed elements of blood
+Two tubes of EDTA-anticoagulated blood. Left tube: after standing, the RBCs have settled at the bottom of the tube. Right tube: Freshly drawn blood
+One microliter of blood contains:
+45 ± 7 (38–52%) for males
+42 ± 5 (37–47%) for females
+Oxygenated: 98–99%
+Deoxygenated: 75%
+About 55% of blood is blood plasma, a fluid that is the blood's liquid medium, which by itself is straw-yellow in color. The blood plasma volume totals of 2.7–3.0 liters (2.8–3.2 quarts) in an average human. It is essentially an aqueous solution containing 92% water, 8% blood plasma proteins, and trace amounts of other materials. Plasma circulates dissolved nutrients, such as glucose, amino acids, and fatty acids (dissolved in the blood or bound to plasma proteins), and removes waste products, such as carbon dioxide, urea, and lactic acid.
+Other important components include:
+The term serum refers to plasma from which the clotting proteins have been removed. Most of the proteins remaining are albumin and immunoglobulins.
+Blood pH is regulated to stay within the narrow range of 7.35 to 7.45, making it slightly basic.[9][10] Blood that has a pH below 7.35 is too acidic, whereas blood pH above 7.45 is too basic. Blood pH, partial pressure of oxygen (pO2), partial pressure of carbon dioxide (pCO2), and bicarbonate (HCO3−) are carefully regulated by a number of homeostatic mechanisms, which exert their influence principally through the respiratory system and the urinary system to control the acid-base balance and respiration. An arterial blood gas test measures these. Plasma also circulates hormones transmitting their messages to various tissues. The list of normal reference ranges for various blood electrolytes is extensive.
+Human blood is typical of that of mammals, although the precise details concerning cell numbers, size, protein structure, and so on, vary somewhat between species. In non-mammalian vertebrates, however, there are some key differences:[11]
+Blood is circulated around the body through blood vessels by the pumping action of the heart. In humans, blood is pumped from the strong left ventricle of the heart through arteries to peripheral tissues and returns to the right atrium of the heart through veins. It then enters the right ventricle and is pumped through the pulmonary artery to the lungs and returns to the left atrium through the pulmonary veins. Blood then enters the left ventricle to be circulated again. Arterial blood carries oxygen from inhaled air to all of the cells of the body, and venous blood carries carbon dioxide, a waste product of metabolism by cells, to the lungs to be exhaled. However, one exception includes pulmonary arteries, which contain the most deoxygenated blood in the body, while the pulmonary veins contain oxygenated blood.
+Additional return flow may be generated by the movement of skeletal muscles, which can compress veins and push blood through the valves in veins toward the right atrium.
+The blood circulation was famously described by William Harvey in 1628.[12]
+In vertebrates, the various cells of blood are made in the bone marrow in a process called hematopoiesis, which includes erythropoiesis, the production of red blood cells; and myelopoiesis, the production of white blood cells and platelets. During childhood, almost every human bone produces red blood cells; as adults, red blood cell production is limited to the larger bones: the bodies of the vertebrae, the breastbone (sternum), the ribcage, the pelvic bones, and the bones of the upper arms and legs. In addition, during childhood, the thymus gland, found in the mediastinum, is an important source of T lymphocytes.[13]
+The proteinaceous component of blood (including clotting proteins) is produced predominantly by the liver, while hormones are produced by the endocrine glands and the watery fraction is regulated by the hypothalamus and maintained by the kidney.
+Healthy erythrocytes have a plasma life of about 120 days before they are degraded by the spleen, and the Kupffer cells in the liver. The liver also clears some proteins, lipids, and amino acids. The kidney actively secretes waste products into the urine.
+About 98.5% [14] of the oxygen in a sample of arterial blood in a healthy human breathing air at sea-level pressure is chemically combined with the hemoglobin. About 1.5% is physically dissolved in the other blood liquids and not connected to hemoglobin. The hemoglobin molecule is the primary transporter of oxygen in mammals and many other species (for exceptions, see below). Hemoglobin has an oxygen binding capacity between 1.36 and 1.40 ml O2 per gram hemoglobin,[15] which increases the total blood oxygen capacity seventyfold,[16] compared to if oxygen solely were carried by its solubility of 0.03 ml O2 per liter blood per mm Hg partial pressure of oxygen (about 100 mm Hg in arteries).[16]
+With the exception of pulmonary and umbilical arteries and their corresponding veins, arteries carry oxygenated blood away from the heart and deliver it to the body via arterioles and capillaries, where the oxygen is consumed; afterwards, venules and veins carry deoxygenated blood back to the heart.
+Under normal conditions in adult humans at rest, hemoglobin in blood leaving the lungs is about 98–99% saturated with oxygen, achieving an oxygen delivery between 950 and 1150 ml/min[17] to the body. In a healthy adult at rest, oxygen consumption is approximately 200–250 ml/min,[17] and deoxygenated blood returning to the lungs is still roughly 75%[18][19] (70 to 78%)[17] saturated. Increased oxygen consumption during sustained exercise reduces the oxygen saturation of venous blood, which can reach less than 15% in a trained athlete; although breathing rate and blood flow increase to compensate, oxygen saturation in arterial blood can drop to 95% or less under these conditions.[20] Oxygen saturation this low is considered dangerous in an individual at rest (for instance, during surgery under anesthesia). Sustained hypoxia (oxygenation less than 90%), is dangerous to health, and severe hypoxia (saturations less than 30%) may be rapidly fatal.[21]
+A fetus, receiving oxygen via the placenta, is exposed to much lower oxygen pressures (about 21% of the level found in an adult's lungs), so fetuses produce another form of hemoglobin with a much higher affinity for oxygen (hemoglobin F) to function under these conditions.[22]
+CO2 is carried in blood in three different ways. (The exact percentages vary depending whether it is arterial or venous blood). Most of it (about 70%) is converted to bicarbonate ions HCO−3 by the enzyme carbonic anhydrase in the red blood cells by the reaction CO2 + H2O → H2CO3 → H+ + HCO−3; about 7% is dissolved in the plasma; and about 23% is bound to hemoglobin as carbamino compounds.[23][24]
+Hemoglobin, the main oxygen-carrying molecule in red blood cells, carries both oxygen and carbon dioxide. However, the CO2 bound to hemoglobin does not bind to the same site as oxygen. Instead, it combines with the N-terminal groups on the four globin chains. However, because of allosteric effects on the hemoglobin molecule, the binding of CO2 decreases the amount of oxygen that is bound for a given partial pressure of oxygen. The decreased binding to carbon dioxide in the blood due to increased oxygen levels is known as the Haldane effect, and is important in the transport of carbon dioxide from the tissues to the lungs. A rise in the partial pressure of CO2 or a lower pH will cause offloading of oxygen from hemoglobin, which is known as the Bohr effect.
+Some oxyhemoglobin loses oxygen and becomes deoxyhemoglobin. Deoxyhemoglobin binds most of the hydrogen ions as it has a much greater affinity for more hydrogen than does oxyhemoglobin.
+In mammals, blood is in equilibrium with lymph, which is continuously formed in tissues from blood by capillary ultrafiltration. Lymph is collected by a system of small lymphatic vessels and directed to the thoracic duct, which drains into the left subclavian vein, where lymph rejoins the systemic blood circulation.
+Blood circulation transports heat throughout the body, and adjustments to this flow are an important part of thermoregulation. Increasing blood flow to the surface (e.g., during warm weather or strenuous exercise) causes warmer skin, resulting in faster heat loss. In contrast, when the external temperature is low, blood flow to the extremities and surface of the skin is reduced and to prevent heat loss and is circulated to the important organs of the body, preferentially.
+Rate of blood flow varies greatly between different organs. Liver has the most abundant blood supply with an approximate flow of 1350 ml/min. Kidney and brain are the second and the third most supplied organs, with 1100 ml/min and ~700 ml/min, respectively.[25]
+Relative rates of blood flow per 100 g of tissue are different, with kidney, adrenal gland and thyroid being the first, second and third most supplied tissues, respectively.[25]
+The restriction of blood flow can also be used in specialized tissues to cause engorgement, resulting in an erection of that tissue; examples are the erectile tissue in the penis and clitoris.
+Another example of a hydraulic function is the jumping spider, in which blood forced into the legs under pressure causes them to straighten for a powerful jump, without the need for bulky muscular legs.[26]
+In insects, the blood (more properly called hemolymph) is not involved in the transport of oxygen. (Openings called tracheae allow oxygen from the air to diffuse directly to the tissues.) Insect blood moves nutrients to the tissues and removes waste products in an open system.
+Other invertebrates use respiratory proteins to increase the oxygen-carrying capacity. Hemoglobin is the most common respiratory protein found in nature. Hemocyanin (blue) contains copper and is found in crustaceans and mollusks. It is thought that tunicates (sea squirts) might use vanabins (proteins containing vanadium) for respiratory pigment (bright-green, blue, or orange).
+In many invertebrates, these oxygen-carrying proteins are freely soluble in the blood; in vertebrates they are contained in specialized red blood cells, allowing for a higher concentration of respiratory pigments without increasing viscosity or damaging blood filtering organs like the kidneys.
+Giant tube worms have unusual hemoglobins that allow them to live in extraordinary environments. These hemoglobins also carry sulfides normally fatal in other animals.
+The coloring matter of blood (hemochrome) is largely due to the protein in the blood responsible for oxygen transport. Different groups of organisms use different proteins.
+Hemoglobin is the principal determinant of the color of blood in vertebrates. Each molecule has four heme groups, and their interaction with various molecules alters the exact color. In vertebrates and other hemoglobin-using creatures, arterial blood and capillary blood are bright red, as oxygen imparts a strong red color to the heme group. Deoxygenated blood is a darker shade of red; this is present in veins, and can be seen during blood donation and when venous blood samples are taken. This is because the spectrum of light absorbed by hemoglobin differs between the oxygenated and deoxygenated states.[27]
+Blood in carbon monoxide poisoning is bright red, because carbon monoxide causes the formation of carboxyhemoglobin. In cyanide poisoning, the body cannot utilize oxygen, so the venous blood remains oxygenated, increasing the redness. There are some conditions affecting the heme groups present in hemoglobin that can make the skin appear blue – a symptom called cyanosis. If the heme is oxidized, methemoglobin, which is more brownish and cannot transport oxygen, is formed. In the rare condition sulfhemoglobinemia, arterial hemoglobin is partially oxygenated, and appears dark red with a bluish hue.
+Veins close to the surface of the skin appear blue for a variety of reasons. However, the factors that contribute to this alteration of color perception are related to the light-scattering properties of the skin and the processing of visual input by the visual cortex, rather than the actual color of the venous blood.[28]
+Skinks in the genus Prasinohaema have green blood due to a buildup of the waste product biliverdin.[29]
+The blood of most mollusks – including cephalopods and gastropods – as well as some arthropods, such as horseshoe crabs, is blue, as it contains the copper-containing protein hemocyanin at concentrations of about 50 grams per liter.[30] Hemocyanin is colorless when deoxygenated and dark blue when oxygenated. The blood in the circulation of these creatures, which generally live in cold environments with low oxygen tensions, is grey-white to pale yellow,[30] and it turns dark blue when exposed to the oxygen in the air, as seen when they bleed.[30] This is due to change in color of hemocyanin when it is oxidized.[30] Hemocyanin carries oxygen in extracellular fluid, which is in contrast to the intracellular oxygen transport in mammals by hemoglobin in RBCs.[30]
+The blood of most annelid worms and some marine polychaetes use chlorocruorin to transport oxygen. It is green in color in dilute solutions.[31]
+Hemerythrin is used for oxygen transport in the marine invertebrates sipunculids, priapulids, brachiopods, and the annelid worm, magelona. Hemerythrin is violet-pink when oxygenated.[31]
+The blood of some species of ascidians and tunicates, also known as sea squirts, contains proteins called vanadins. These proteins are based on vanadium, and give the creatures a concentration of vanadium in their bodies 100 times higher than the surrounding sea water. Unlike hemocyanin and hemoglobin, hemovanadin is not an oxygen carrier. When exposed to oxygen, however, vanadins turn a mustard yellow.
+Substances other than oxygen can bind to hemoglobin; in some cases this can cause irreversible damage to the body. Carbon monoxide, for example, is extremely dangerous when carried to the blood via the lungs by inhalation, because carbon monoxide irreversibly binds to hemoglobin to form carboxyhemoglobin, so that less hemoglobin is free to bind oxygen, and fewer oxygen molecules can be transported throughout the blood. This can cause suffocation insidiously. A fire burning in an enclosed room with poor ventilation presents a very dangerous hazard, since it can create a build-up of carbon monoxide in the air. Some carbon monoxide binds to hemoglobin when smoking tobacco.[34]
+Blood for transfusion is obtained from human donors by blood donation and stored in a blood bank. There are many different blood types in humans, the ABO blood group system, and the Rhesus blood group system being the most important. Transfusion of blood of an incompatible blood group may cause severe, often fatal, complications, so crossmatching is done to ensure that a compatible blood product is transfused.
+Other blood products administered intravenously are platelets, blood plasma, cryoprecipitate, and specific coagulation factor concentrates.
+Many forms of medication (from antibiotics to chemotherapy) are administered intravenously, as they are not readily or adequately absorbed by the digestive tract.
+After severe acute blood loss, liquid preparations, generically known as plasma expanders, can be given intravenously, either solutions of salts (NaCl, KCl, CaCl2 etc.) at physiological concentrations, or colloidal solutions, such as dextrans, human serum albumin, or fresh frozen plasma. In these emergency situations, a plasma expander is a more effective life-saving procedure than a blood transfusion, because the metabolism of transfused red blood cells does not restart immediately after a transfusion.
+In modern evidence-based medicine, bloodletting is used in management of a few rare diseases, including hemochromatosis and polycythemia. However, bloodletting and leeching were common unvalidated interventions used until the 19th century, as many diseases were incorrectly thought to be due to an excess of blood, according to Hippocratic medicine.
+English blood (Old English blod) derives from Germanic and has cognates with a similar range of meanings in all other Germanic languages (e.g. German Blut, Swedish blod, Gothic blōþ). There is no accepted Indo-European etymology.[35]
+Robin Fåhræus [pl; sv] (a Swedish physician who devised the erythrocyte sedimentation rate) suggested that the Ancient Greek system of humorism, wherein the body was thought to contain four distinct bodily fluids (associated with different temperaments), were based upon the observation of blood clotting in a transparent container. When blood is drawn in a glass container and left undisturbed for about an hour, four different layers can be seen. A dark clot forms at the bottom (the "black bile"). Above the clot is a layer of red blood cells (the "blood"). Above this is a whitish layer of white blood cells (the "phlegm"). The top layer is clear yellow serum (the "yellow bile").[36]
+The ABO blood group system was discovered in the year 1900 by Karl Landsteiner. Jan Janský is credited with the first classification of blood into the four types (A, B, AB, and O) in 1907, which remains in use today. In 1907 the first blood transfusion was performed that used the ABO system to predict compatibility.[37] The first non-direct transfusion was performed on March 27, 1914. The Rhesus factor was discovered in 1937.
+Due to its importance to life, blood is associated with a large number of beliefs. One of the most basic is the use of blood as a symbol for family relationships through birth/parentage; to be "related by blood" is to be related by ancestry or descendence, rather than marriage. This bears closely to bloodlines, and sayings such as "blood is thicker than water" and "bad blood", as well as "Blood brother".
+Blood is given particular emphasis in the Jewish and Christian religions, because Leviticus 17:11 says "the life of a creature is in the blood." This phrase is part of the Levitical law forbidding the drinking of blood or eating meat with the blood still intact instead of being poured off.
+Mythic references to blood can sometimes be connected to the life-giving nature of blood, seen in such events as childbirth, as contrasted with the blood of injury or death.
+In many indigenous Australian Aboriginal peoples' traditions, ochre (particularly red) and blood, both high in iron content and considered Maban, are applied to the bodies of dancers for ritual. As Lawlor states:
+In many Aboriginal rituals and ceremonies, red ochre is rubbed all over the naked bodies of the dancers. In secret, sacred male ceremonies, blood extracted from the veins of the participant's arms is exchanged and rubbed on their bodies. Red ochre is used in similar ways in less-secret ceremonies. Blood is also used to fasten the feathers of birds onto people's bodies. Bird feathers contain a protein that is highly magnetically sensitive.[38]
+Lawlor comments that blood employed in this fashion is held by these peoples to attune the dancers to the invisible energetic realm of the Dreamtime. Lawlor then connects these invisible energetic realms and magnetic fields, because iron is magnetic.
+Among the Germanic tribes, blood was used during their sacrifices; the Blóts. The blood was considered to have the power of its originator, and, after the butchering, the blood was sprinkled on the walls, on the statues of the gods, and on the participants themselves. This act of sprinkling blood was called blóedsian in Old English, and the terminology was borrowed by the Roman Catholic Church becoming to bless and blessing. The Hittite word for blood, ishar was a cognate to words for "oath" and "bond", see Ishara.
+The Ancient Greeks believed that the blood of the gods, ichor, was a substance that was poisonous to mortals.
+As a relic of Germanic Law, the cruentation, an ordeal where the corpse of the victim was supposed to start bleeding in the presence of the murderer, was used until the early 17th century.
+In Genesis 9:4, God prohibited Noah and his sons from eating blood (see Noahide Law). This command continued to be observed by the Eastern Orthodox.
+It is also found in the Bible that when the Angel of Death came around to the Hebrew house that the first-born child would not die if the angel saw lamb's blood wiped across the doorway.
+At the Council of Jerusalem, the apostles prohibited certain Christians from consuming blood – this is documented in Acts 15:20 and 29. This chapter specifies a reason (especially in verses 19–21): It was to avoid offending Jews who had become Christians, because the Mosaic Law Code prohibited the practice.
+Christ's blood is the means for the atonement of sins. Also, ″... the blood of Jesus Christ his [God] Son cleanseth us from all sin." (1 John 1:7), “... Unto him [God] that loved us, and washed us from our sins in his own blood." (Revelation 1:5), and "And they overcame him (Satan) by the blood
+of the Lamb [Jesus the Christ], and by the word of their testimony ...” (Revelation 12:11).
+Some Christian churches, including Roman Catholicism, Eastern Orthodoxy, Oriental Orthodoxy, and the Assyrian Church of the East teach that, when consecrated, the Eucharistic wine actually becomes the blood of Jesus for worshippers to drink. Thus in the consecrated wine, Jesus becomes spiritually and physically present. This teaching is rooted in the Last Supper, as written in the four gospels of the Bible, in which Jesus stated to his disciples that the bread that they ate was his body, and the wine was his blood. "This cup is the new testament in my blood, which is shed for you." (Luke 22:20).
+Most forms of Protestantism, especially those of a Methodist or Presbyterian lineage, teach that the wine is no more than a symbol of the blood of Christ, who is spiritually but not physically present. Lutheran theology teaches that the body and blood is present together "in, with, and under" the bread and wine of the Eucharistic feast.
+In Judaism, animal blood may not be consumed even in the smallest quantity (Leviticus 3:17 and elsewhere); this is reflected in Jewish dietary laws (Kashrut). Blood is purged from meat by rinsing and soaking in water (to loosen clots), salting and then rinsing with water again several times.[39] Eggs must also be checked and any blood spots removed before consumption.[40] Although blood from fish is biblically kosher, it is rabbinically forbidden to consume fish blood to avoid the appearance of breaking the Biblical prohibition.[41]
+Another ritual involving blood involves the covering of the blood of fowl and game after slaughtering (Leviticus 17:13); the reason given by the Torah is: "Because the life of the animal is [in] its blood" (ibid 17:14). In relation to human beings, Kabbalah expounds on this verse that the animal soul of a person is in the blood, and that physical desires stem from it.
+Likewise, the mystical reason for salting temple sacrifices and slaughtered meat is to remove the blood of animal-like passions from the person. By removing the animal's blood, the animal energies and life-force contained in the blood are removed, making the meat fit for human consumption.[42]
+Consumption of food containing blood is forbidden by Islamic dietary laws. This is derived from the statement in the Qur'an, sura Al-Ma'ida (5:3): "Forbidden to you (for food) are: dead meat, blood, the flesh of swine, and that on which has been invoked the name of other than Allah."
+Blood is considered unclean, hence there are specific methods to obtain physical and ritual status of cleanliness once bleeding has occurred. Specific rules and prohibitions apply to menstruation, postnatal bleeding and irregular vaginal bleeding. When an animal has been slaughtered, the animal's neck is cut in a way to ensure that the spine is not severed, hence the brain may send commands to the heart to pump blood to it for oxygen. In this way, blood is removed from the body, and the meat is generally now safe to cook and eat. In modern times, blood transfusions are generally not considered against the rules.
+Based on their interpretation of scriptures such as Acts 15:28, 29 ("Keep abstaining...from blood."), many Jehovah's Witnesses neither consume blood nor accept transfusions of whole blood or its major components: red blood cells, white blood cells, platelets (thrombocytes), and plasma. Members may personally decide whether they will accept medical procedures that involve their own blood or substances that are further fractionated from the four major components.[43]
+In south East Asian popular culture, it is often said that if a man's nose produces a small flow of blood, he is experiencing sexual desire. This often appears in Chinese-language and Hong Kong films as well as in Japanese and Korean culture parodied in anime, manga, and drama. Characters, mostly males, will often be shown with a nosebleed if they have just seen someone nude or in little clothing, or if they have had an erotic thought or fantasy; this is based on the idea that a male's blood pressure will spike dramatically when aroused.[44][unreliable source?]
+Vampires are mythical creatures that drink blood directly for sustenance, usually with a preference for human blood. Cultures all over the world have myths of this kind; for example the 'Nosferatu' legend, a human who achieves damnation and immortality by drinking the blood of others, originates from Eastern European folklore. Ticks, leeches, female mosquitoes, vampire bats, and an assortment of other natural creatures do consume the blood of other animals, but only bats are associated with vampires. This has no relation to vampire bats, which are new world creatures discovered well after the origins of the European myths.
+Blood residue can help forensic investigators identify weapons, reconstruct a criminal action, and link suspects to the crime. Through bloodstain pattern analysis, forensic information can also be gained from the spatial distribution of bloodstains.
+Blood residue analysis is also a technique used in archeology.
+Blood is one of the body fluids that has been used in art.[45] In particular, the performances of Viennese Actionist Hermann Nitsch, Istvan Kantor, Franko B, Lennie Lee, Ron Athey, Yang Zhichao, Lucas Abela and Kira O'Reilly, along with the photography of Andres Serrano, have incorporated blood as a prominent visual element. Marc Quinn has made sculptures using frozen blood, including a cast of his own head made using his own blood.
+The term blood is used in genealogical circles to refer to one's ancestry, origins, and ethnic background as in the word bloodline. Other terms where blood is used in a family history sense are blue-blood, royal blood, mixed-blood and blood relative.

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+Mesothelae
+Opisthothelae
+ See Spider taxonomy.
+Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom,[2] and spinnerets that extrude silk.[3] They are the largest order of arachnids and rank seventh in total species diversity among all orders of organisms.[4] Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every habitat with the exceptions of air and sea colonization. As of July 2019[update], at least 48,200 spider species, and 120 families have been recorded by taxonomists.[1] However, there has been dissension within the scientific community as to how all these families should be classified, as evidenced by the over 20 different classifications that have been proposed since 1900.[5]
+Anatomically, spiders (as with all arachnids) differ from other arthropods in that the usual body segments are fused into two tagmata, the prosoma, or cephalothorax, and opisthosoma, or abdomen, and joined by a small, cylindrical pedicel (however, as there is currently neither paleontological nor embryological evidence that spiders ever had a separate thorax-like division, there exists an argument against the validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly, arguments can be formed against use of the term abdomen, as the opisthosoma of all spiders contains a heart and respiratory organs, organs atypical of an abdomen[6]). Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.
+Their abdomens bear appendages that have been modified into spinnerets that extrude silk from up to six types of glands. Spider webs vary widely in size, shape and the amount of sticky thread used. It now appears that the spiral orb web may be one of the earliest forms, and spiders that produce tangled cobwebs are more abundant and diverse than orb-web spiders. Spider-like arachnids with silk-producing spigots appeared in the Devonian period about 386 million years ago, but these animals apparently lacked spinnerets. True spiders have been found in Carboniferous rocks from 318 to 299 million years ago, and are very similar to the most primitive surviving suborder, the Mesothelae. The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, before 200 million years ago.
+The species Bagheera kiplingi was described as herbivorous in 2008,[7] but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. It is estimated that the world's 25 million tons of spiders kill 400–800 million tons of prey per year.[8] Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus Portia show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, so they liquefy their food by flooding it with digestive enzymes. They also grind food with the bases of their pedipalps, as arachnids do not have the mandibles that crustaceans and insects have.
+To avoid being eaten by the females, which are typically much larger, male spiders identify themselves to potential mates by a variety of complex courtship rituals. Males of most species survive a few matings, limited mainly by their short life spans. Females weave silk egg-cases, each of which may contain hundreds of eggs. Females of many species care for their young, for example by carrying them around or by sharing food with them. A minority of species are social, building communal webs that may house anywhere from a few to 50,000 individuals. Social behavior ranges from precarious toleration, as in the widow spiders, to co-operative hunting and food-sharing. Although most spiders live for at most two years, tarantulas and other mygalomorph spiders can live up to 25 years in captivity.
+While the venom of a few species is dangerous to humans, scientists are now researching the use of spider venom in medicine and as non-polluting pesticides. Spider silk provides a combination of lightness, strength and elasticity that is superior to that of synthetic materials, and spider silk genes have been inserted into mammals and plants to see if these can be used as silk factories. As a result of their wide range of behaviors, spiders have become common symbols in art and mythology symbolizing various combinations of patience, cruelty and creative powers. An abnormal fear of spiders is called arachnophobia.
+1: pedipalp
+ 2: trichobothria
+ 3: carapace of prosoma (cephalothorax)
+ 4: opisthosoma (abdomen)
+ 5: eyes – AL (anterior lateral)
+    AM (anterior median)
+    PL (posterior lateral)
+    PM (posterior median)
+Leg segments:
+ 6: costa
+ 7: trochanter
+ 8: patella
+ 9: tibia
+10: metatarsus
+11: tarsus
+13: claw
+14: chelicera
+15: sternum of prosoma
+16: pedicel (also called pedicle)
+17: book lung sac
+18: book lung stigma
+19: epigastric fold
+20: epigyne
+21: anterior spinneret
+22: posterior spinneret
+Spiders are chelicerates and therefore arthropods.[9] As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo.[10] Being chelicerates, their bodies consist of two tagmata, sets of segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete fusion of the segments that in an insect would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma.[9] In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel.[11] The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws".[10][12] The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.[9]
+Spiders and scorpions are members of one chelicerate group, the arachnids.[12] Scorpions' chelicerae have three sections and are used in feeding.[13] Spiders' chelicerae have two sections and terminate in fangs that are generally venomous, and fold away behind the upper sections while not in use. The upper sections generally have thick "beards" that filter solid lumps out of their food, as spiders can take only liquid food.[11] Scorpions' pedipalps generally form large claws for capturing prey,[13] while those of spiders are fairly small appendages whose bases also act as an extension of the mouth; in addition, those of male spiders have enlarged last sections used for sperm transfer.[11]
+In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex carapace, while the underside is covered by two rather flat plates. The abdomen is soft and egg-shaped. It shows no sign of segmentation, except that the primitive Mesothelae, whose living members are the Liphistiidae, have segmented plates on the upper surface.[11]
+Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas round the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end.[14] However, in spiders, it occupies only the upper part of the abdomen, and blood is discharged into the hemocoel by one artery that opens at the rear end of the abdomen and by branching arteries that pass through the pedicle and open into several parts of the cephalothorax. Hence spiders have open circulatory systems.[11] The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.[12]
+Spiders have developed several different respiratory anatomies, based on book lungs, a tracheal system, or both. Mygalomorph and Mesothelae spiders have two pairs of book lungs filled with haemolymph, where openings on the ventral surface of the abdomen allow air to enter and diffuse oxygen. This is also the case for some basal araneomorph spiders, like the family Hypochilidae, but the remaining members of this group have just the anterior pair of book lungs intact while the posterior pair of breathing organs are partly or fully modified into tracheae, through which oxygen is diffused into the haemolymph or directly to the tissue and organs.[11] The tracheal system has most likely evolved in small ancestors to help resist desiccation.[12] The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets.[11] Spiders that have tracheae generally have higher metabolic rates and better water conservation.[15] Spiders are ectotherms, so environmental temperatures affect their activity.[16]
+Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae.[11] The families Uloboridae and Holarchaeidae, and some Liphistiidae spiders, have lost their venom glands, and kill their prey with silk instead.[17] Like most arachnids, including scorpions,[12] spiders have a narrow gut that can only cope with liquid food and two sets of filters to keep solids out.[11] They use one of two different systems of external digestion. Some pump digestive enzymes from the midgut into the prey and then suck the liquified tissues of the prey into the gut, eventually leaving behind the empty husk of the prey. Others grind the prey to pulp using the chelicerae and the bases of the pedipalps, while flooding it with enzymes; in these species, the chelicerae and the bases of the pedipalps form a preoral cavity that holds the food they are processing.[11]
+The stomach in the cephalothorax acts as a pump that sends the food deeper into the digestive system. The midgut bears many digestive ceca, compartments with no other exit, that extract nutrients from the food; most are in the abdomen, which is dominated by the digestive system, but a few are found in the cephalothorax.[11]
+Most spiders convert nitrogenous waste products into uric acid, which can be excreted as a dry material. Malphigian tubules ("little tubes") extract these wastes from the blood in the hemocoel and dump them into the cloacal chamber, from which they are expelled through the anus.[11] Production of uric acid and its removal via Malphigian tubules are a water-conserving feature that has evolved independently in several arthropod lineages that can live far away from water,[18] for example the tubules of insects and arachnids develop from completely different parts of the embryo.[12] However, a few primitive spiders, the suborder Mesothelae and infraorder Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"),[11] which use large amounts of water to excrete nitrogenous waste products as ammonia.[18]
+The basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia.[19] Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen;[11][12][19] in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.[15]
+Despite the relatively small central nervous system, some spiders (like Portia) exhibit complex behaviour, including the ability to use a trial-and-error approach.[20][21]
+Spiders have primarily four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another.[11] The principal pair at the front are of the type called pigment-cup ocelli ("little eyes"), which in most arthropods are only capable of detecting the direction from which light is coming, using the shadow cast by the walls of the cup. However, in spiders these eyes are capable of forming images.[22][23] The other pairs, called secondary eyes, are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the principal eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torchlight reflected from the tapeta. On the other hand, jumping spiders' secondary eyes have no tapeta.[11]
+Other differences between the principal and secondary eyes are that the latter have rhabdomeres that point away from incoming light, just like in vertebrates, while the arrangement is the opposite in the former. The principal eyes are also the only ones with eye muscles, allowing them to move the retina. Having no muscles, the secondary eyes are immobile.[24]
+Some jumping spiders' visual acuity exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects; in fact the human eye is only about five times sharper than a jumping spider's. They achieve this by a telephotographic series of lenses, a four-layer retina and the ability to swivel their eyes and integrate images from different stages in the scan. The downside is that the scanning and integrating processes are relatively slow.[20]
+There are spiders with a reduced number of eyes. Of these, those with six eyes (such as Periegops suterii) are the most numerous and are missing a pair of eyes on the anterior median line;[25] other species have four eyes and some just two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.
+As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae.[22] An adult Araneus may have up to 1,000 such chemosensitive setae, most on the tarsi of the first pair of legs. Males have more chemosensitive bristles on their pedipalps than females. They have been shown to be responsive to sex pheromones produced by females, both contact and air-borne.[26] The jumping spider Evarcha culicivora uses the scent of blood from mammals and other vertebrates, which is obtained by capturing blood-filled mosquitoes, to attract the opposite sex. Because they are able to tell the sexes apart, it is assumed the blood scent is mixed with pheromones.[27] Spiders also have in the joints of their limbs slit sensillae that detect force and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.[11]
+Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' proprioceptors, sensors that report the force exerted by muscles and the degree of bending in the body and joints, are well-understood. On the other hand, little is known about what other internal sensors spiders or other arthropods may have.[22]
+Each of the eight legs of a spider consists of seven distinct parts. The part closest to and attaching the leg to the cephalothorax is the coxa; the next segment is the short trochanter that works as a hinge for the following long segment, the femur; next is the spider's knee, the patella, which acts as the hinge for the tibia; the metatarsus is next, and it connects the tibia to the tarsus (which may be thought of as a foot of sorts); the tarsus ends in a claw made up of either two or three points, depending on the family to which the spider belongs. Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors.[28] The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter).[29] As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up.[11] Spiders can generate pressures up to eight times their resting level to extend their legs,[30] and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs.[11] Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.[29]
+Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine bristles between the paired claws at the tips of their legs. These tufts, known as scopulae, consist of bristles whose ends are split into as many as 1,000 branches, and enable spiders with scopulae to walk up vertical glass and upside down on ceilings. It appears that scopulae get their grip from contact with extremely thin layers of water on surfaces.[11] Spiders, like most other arachnids, keep at least four legs on the surface while walking or running.[31]
+The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk.[11]
+Silk is mainly composed of a protein very similar to that used in insect silk. It is initially a liquid, and hardens not by exposure to air but as a result of being drawn out, which changes the internal structure of the protein.[32] It is similar in tensile strength to nylon and biological materials such as chitin, collagen and cellulose, but is much more elastic. In other words, it can stretch much further before breaking or losing shape.[11]
+Some spiders have a cribellum, a modified spinneret with up to 40,000 spigots, each of which produces a single very fine fiber. The fibers are pulled out by the calamistrum, a comblike set of bristles on the jointed tip of the cribellum, and combined into a composite woolly thread that is very effective in snagging the bristles of insects. The earliest spiders had cribella, which produced the first silk capable of capturing insects, before spiders developed silk coated with sticky droplets. However, most modern groups of spiders have lost the cribellum.[11]
+Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a "safety rope"; for nest-building; and as "parachutes" by the young of some species.[11]
+Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods,[33] male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs onto which they ejaculate and then transfer the sperm to special syringe-styled structures, palpal bulbs or palpal organs, borne on the tips of the pedipalps of mature males. When a male detects signs of a female nearby he checks whether she is of the same species and whether she is ready to mate; for example in species that produce webs or "safety ropes", the male can identify the species and sex of these objects by "smell".[11]
+Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female via one or two openings on the underside of her abdomen.[11]
+Female spiders' reproductive tracts are arranged in one of two ways. The ancestral arrangement ("haplogyne" or "non-entelegyne") consists of a single genital opening, leading to two seminal receptacles (spermathecae) in which females store sperm. In the more advanced arrangement ("entelegyne" ), there are two further openings leading directly to the spermathecae, creating a "flow through" system rather than a "first-in first-out" one. Eggs are as a general rule only fertilized during oviposition when the stored sperm is released from its chamber, rather than in the ovarian cavity.[34] A few exceptions exist, such as Parasteatoda tepidariorum. In these species the female appears to be able to activate the dormant sperm before oviposition, allowing them to migrate to the ovarian cavity where fertilization occurs.[35][36][37] The only known example of direct fertilization between male and female is an Israeli spider, Harpactea sadistica, which has evolved traumatic insemination. In this species the male will penetrate its pedipalps through the female's body wall and inject his sperm directly into her ovaries, where the embryos inside the fertilized eggs will start to develop before being laid.[38]
+Males of the genus Tidarren amputate one of their palps before maturation and enter adult life with one palp only. The palps are 20% of the male's body mass in this species, and detaching one of the two improves mobility. In the Yemeni species Tidarren argo, the remaining palp is then torn off by the female. The separated palp remains attached to the female's epigynum for about four hours and apparently continues to function independently. In the meantime, the female feeds on the palpless male.[39] In over 60% of cases, the female of the Australian redback spider kills and eats the male after it inserts its second palp into the female's genital opening; in fact, the males co-operate by trying to impale themselves on the females' fangs. Observation shows that most male redbacks never get an opportunity to mate, and the "lucky" ones increase the likely number of offspring by ensuring that the females are well-fed.[40] However, males of most species survive a few matings, limited mainly by their short life spans. Some even live for a while in their mates' webs.[41]
+The tiny male of the Golden orb weaver (Trichonephila clavipes) (near the top of the leaf) is protected from the female by producing the right vibrations in the web, and may be too small to be worth eating.
+Orange spider egg sac hanging from ceiling
+Gasteracantha mammosa spiderlings next to their eggs capsule
+Wolf spider carrying its young on its abdomen
+Females lay up to 3,000 eggs in one or more silk egg sacs,[11] which maintain a fairly constant humidity level.[41] In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.[11]
+Baby spiders pass all their larval stages inside the egg and hatch as spiderlings, very small and sexually immature but similar in shape to adults. Some spiders care for their young, for example a wolf spider's brood clings to rough bristles on the mother's back,[11] and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.[41]
+Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch.[42] In some species males mate with newly-molted females, which are too weak to be dangerous to the males.[41] Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years,[11][43] and an Australian female trapdoor spider was documented to have lived in the wild for 43 years, dying of a parasitic wasp attack.[44]
+Spiders occur in a large range of sizes. The smallest, Patu digua from Colombia, are less than 0.37 mm (0.015 in) in body length. The largest and heaviest spiders occur among tarantulas, which can have body lengths up to 90 mm (3.5 in) and leg spans up to 250 mm (9.8 in).[45]
+Only three classes of pigment (ommochromes, bilins and guanine) have been identified in spiders, although other pigments have been detected but not yet characterized. Melanins, carotenoids and pterins, very common in other animals, are apparently absent. In some species, the exocuticle of the legs and prosoma is modified by a tanning process, resulting in a brown coloration.[46]
+Bilins are found, for example, in Micrommata virescens, resulting in its green color. Guanine is responsible for the white markings of the European garden spider Araneus diadematus. It is in many species accumulated in specialized cells called guanocytes. In genera such as Tetragnatha, Leucauge, Argyrodes or Theridiosoma, guanine creates their silvery appearance. While guanine is originally an end-product of protein metabolism, its excretion can be blocked in spiders, leading to an increase in its storage.[46] Structural colors occur in some species, which are the result of the diffraction, scattering or interference of light, for example by modified setae or scales. The white prosoma of Argiope results from bristles reflecting the light, Lycosa and Josa both have areas of modified cuticle that act as light reflectors.[46]
+While in many spiders color is fixed throughout their lifespan, in some groups, color may be variable in response to environmental and internal conditions.[46] Choice of prey may be able to alter the color of spiders. For example, the abdomen of Theridion grallator will become orange if the spider ingests certain species of Diptera and adult Lepidoptera, but if it consumes Homoptera or larval Lepidoptera, then the abdomen becomes green.[47] Environmentally induced color changes may be morphological (occurring over several days) or physiological (occurring near instantly). Morphological changes require pigment synthesis and degradation. In contrast to this, physiological changes occur by changing the position of pigment-containing cells.[46] An example of morphological color changes is background matching. Misumena vatia for instance can change its body color to match the substrate it lives on which makes it more difficult to be detected by prey.[48] An example of physiological color change is observed in Cyrtophora cicatrosa, which can change its body color from white to brown near instantly.[46]
+Although spiders are generally regarded as predatory, the jumping spider Bagheera kiplingi gets over 90% of its food from fairly solid plant material produced by acacias as part of a mutually beneficial relationship with a species of ant.[49]
+Juveniles of some spiders in the families Anyphaenidae, Corinnidae, Clubionidae, Thomisidae and Salticidae feed on plant nectar. Laboratory studies show that they do so deliberately and over extended periods, and periodically clean themselves while feeding. These spiders also prefer sugar solutions to plain water, which indicates that they are seeking nutrients. Since many spiders are nocturnal, the extent of nectar consumption by spiders may have been underestimated. Nectar contains amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.[50]
+Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.[50]
+The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.[11]
+Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it.[51] A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.[11]
+Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus Hyptiotes and the family Theridiosomatidae stretch their webs and then release them when prey strike them, but do not actively move their webs. Those of the family Deinopidae weave even smaller webs, hold them outstretched between their first two pairs of legs, and lunge and push the webs as much as twice their own body length to trap prey, and this move may increase the webs' area by a factor of up to ten. Experiments have shown that Deinopis spinosus has two different techniques for trapping prey: backwards strikes to catch flying insects, whose vibrations it detects; and forward strikes to catch ground-walking prey that it sees. These two techniques have also been observed in other deinopids. Walking insects form most of the prey of most deinopids, but one population of Deinopis subrufa appears to live mainly on tipulid flies that they catch with the backwards strike.[52]
+Mature female bolas spiders of the genus Mastophora build "webs" that consist of only a single "trapeze line", which they patrol. They also construct a bolas made of a single thread, tipped with a large ball of very wet sticky silk. They emit chemicals that resemble the pheromones of moths, and then swing the bolas at the moths. Although they miss on about 50% of strikes, they catch about the same weight of insects per night as web-weaving spiders of similar size. The spiders eat the bolas if they have not made a kill in about 30 minutes, rest for a while, and then make new bolas.[53][54] Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.[55]
+The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush predators that lurk in burrows, often closed by trapdoors and often surrounded by networks of silk threads that alert these spiders to the presence of prey.[15] Other ambush predators do without such aids, including many crab spiders,[11] and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking.[46] Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.[11]
+Some jumping spiders of the genus Portia hunt other spiders in ways that seem intelligent,[20] outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species.[56] However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.[20]
+Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective bristles to resemble the shiny bodies of ants. In some spider species, males and females mimic different ant species, as female spiders are usually much larger than males. Ant-mimicking spiders also modify their behavior to resemble that of the target species of ant; for example, many adopt a zig-zag pattern of movement, ant-mimicking jumping spiders avoid jumping, and spiders of the genus Synemosyna walk on the outer edges of leaves in the same way as Pseudomyrmex. Ant mimicry in many spiders and other arthropods may be for protection from predators that hunt by sight, including birds, lizards and spiders. However, several ant-mimicking spiders prey either on ants or on the ants' "livestock", such as aphids. When at rest, the ant-mimicking crab spider Amyciaea does not closely resemble Oecophylla, but while hunting it imitates the behavior of a dying ant to attract worker ants. After a kill, some ant-mimicking spiders hold their victims between themselves and large groups of ants to avoid being attacked.[57]
+There is strong evidence that spiders' coloration is camouflage that helps them to evade their major predators, birds and parasitic wasps, both of which have good color vision. Many spider species are colored so as to merge with their most common backgrounds, and some have disruptive coloration, stripes and blotches that break up their outlines. In a few species, such as the Hawaiian happy-face spider, Theridion grallator, several coloration schemes are present in a ratio that appears to remain constant, and this may make it more difficult for predators to recognize the species. Most spiders are insufficiently dangerous or unpleasant-tasting for warning coloration to offer much benefit. However, a few species with powerful venom, large jaws or irritant bristles have patches of warning colors, and some actively display these colors when threatened.[46][58]
+Many of the family Theraphosidae, which includes tarantulas and baboon spiders, have urticating hairs on their abdomens and use their legs to flick them at attackers. These bristles are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom.[59] A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles.[60] The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.[61]
+A few spider species that build webs live together in large colonies and show social behavior, although not as complex as in social insects. Anelosimus eximius (in the family Theridiidae) can form colonies of up to 50,000 individuals.[62] The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social.[63] Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food.[64] Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and Mallos gregalis (family Dictynidae).[65] Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this.[66] The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings.[49] Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.[67]
+There is no consistent relationship between the classification of spiders and the types of web they build: species in the same genus may build very similar or significantly different webs. Nor is there much correspondence between spiders' classification and the chemical composition of their silks. Convergent evolution in web construction, in other words use of similar techniques by remotely related species, is rampant. Orb web designs and the spinning behaviors that produce them are the best understood. The basic radial-then-spiral sequence visible in orb webs and the sense of direction required to build them may have been inherited from the common ancestors of most spider groups.[68] However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a subgroup that evolved from orb-web spiders, and non-orb spiders have over 40% more species and are four times as abundant as orb-web spiders. Their greater success may be because sphecid wasps, which are often the dominant predators of spiders, much prefer to attack spiders that have flat webs.[69]
+About half the potential prey that hit orb webs escape. A web has to perform three functions: intercepting the prey (intersection), absorbing its momentum without breaking (stopping), and trapping the prey by entangling it or sticking to it (retention). No single design is best for all prey. For example: wider spacing of lines will increase the web's area and hence its ability to intercept prey, but reduce its stopping power and retention; closer spacing, larger sticky droplets and thicker lines would improve retention, but would make it easier for potential prey to see and avoid the web, at least during the day. However, there are no consistent differences between orb webs built for use during the day and those built for use at night. In fact, there is no simple relationship between orb web design features and the prey they capture, as each orb-weaving species takes a wide range of prey.[68]
+The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.[68]
+Horizontal orb webs are fairly common, despite being less effective at intercepting and retaining prey and more vulnerable to damage by rain and falling debris. Various researchers have suggested that horizontal webs offer compensating advantages, such as reduced vulnerability to wind damage; reduced visibility to prey flying upwards, because of the backlighting from the sky; enabling oscillations to catch insects in slow horizontal flight. However, there is no single explanation for the common use of horizontal orb webs.[68]
+Spiders often attach highly visible silk bands, called decorations or stabilimenta, to their webs. Field research suggests that webs with more decorative bands captured more prey per hour.[70] However, a laboratory study showed that spiders reduce the building of these decorations if they sense the presence of predators.[71]
+There are several unusual variants of orb web, many of them convergently evolved, including: attachment of lines to the surface of water, possibly to trap insects in or on the surface; webs with twigs through their centers, possibly to hide the spiders from predators; "ladderlike" webs that appear most effective in catching moths. However, the significance of many variations is unclear.[68]
+In 1973, Skylab 3 took two orb-web spiders into space to test their web-spinning capabilities in zero gravity. At first, both produced rather sloppy webs, but they adapted quickly.[72]
+Members of the family Theridiidae weave irregular, tangled, three-dimensional webs, popularly known as cobwebs. There seems to be an evolutionary trend towards a reduction in the amount of sticky silk used, leading to its total absence in some species. The construction of cobwebs is less stereotyped than that of orb-webs, and may take several days.[69]
+The Linyphiidae generally make horizontal but uneven sheets, with tangles of stopping threads above. Insects that hit the stopping threads fall onto the sheet or are shaken onto it by the spider, and are held by sticky threads on the sheet until the spider can attack from below.[73]
+Although the fossil record of spiders is considered poor,[74] almost 1000 species have been described from fossils.[75] Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber.[75] The oldest known amber that contains fossil arthropods dates from 130 million years ago in the Early Cretaceous period. In addition to preserving spiders' anatomy in very fine detail, pieces of amber show spiders mating, killing prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached;[76] the oldest fossil web found so far is 100 million years old.[77] Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.[76]
+The oldest known exclusively terrestrial arachnid is the trigonotarbid Palaeotarbus jerami, from about 420 million years ago in the Silurian period, and had a triangular cephalothorax and segmented abdomen, as well as eight legs and a pair of pedipalps.[78] Attercopus fimbriunguis, from 386 million years ago in the Devonian period, bears the earliest known silk-producing spigots, and was therefore hailed as a spider at the time of its discovery.[79] However, these spigots may have been mounted on the underside of the abdomen rather than on spinnerets, which are modified appendages and whose mobility is important in the building of webs. Hence Attercopus and the similar Permian arachnid Permarachne may not have been true spiders, and probably used silk for lining nests or producing egg cases rather than for building webs.[3] The largest known fossil spider as of 2011 is the araneid Nephila jurassica, from about 165 million years ago, recorded from Daohuogo, Inner Mongolia in China.[80] Its body length is almost 25 mm, (i.e., almost one inch).
+Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae.[79]
+The mesothelid Paleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets.[81] Although the Permian period 299 to 251 million years ago saw rapid diversification of flying insects, there are very few fossil spiders from this period.[79]
+The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appear in the Triassic well before 200 million years ago. Some Triassic mygalomorphs appear to be members of the family Hexathelidae, whose modern members include the notorious Sydney funnel-web spider, and their spinnerets appear adapted for building funnel-shaped webs to catch jumping insects. Araneomorphae account for the great majority of modern spiders, including those that weave the familiar orb-shaped webs. The Jurassic and Cretaceous periods provide a large number of fossil spiders, including representatives of many modern families.[79]
+Xiphosura (horseshoe crabs)
+Eurypterida†
+Chasmataspidida†
+Scorpiones
+Opiliones (harvestmen)
+Pseudoscorpiones
+Solifugae (sun spiders)
+Palpigradi (microwhip scorpions)
+Trigonotarbida†
+Araneae (spiders)
+Haptopoda†
+Amblypygi (whip spiders)
+Thelyphonida (whip scorpions)
+Schizomida
+Ricinulei (hooded tickspiders)
+Anactinotrichida
+Acariformes (mites)
+It is now agreed that spiders (Araneae) are monophyletic (i.e., members of a group of organisms that form a clade, consisting of a last common ancestor and all of its descendants).[83] There has been debate about what their closest evolutionary relatives are, and how all of these evolved from the ancestral chelicerates, which were marine animals. The cladogram on the right is based on J. W. Shultz' analysis (2007). Other views include proposals that: scorpions are more closely related to the extinct marine scorpioid eurypterids than to spiders; spiders and Amblypygi are a monophyletic group. The appearance of several multi-way branchings in the tree on the right shows that there are still uncertainties about relationships between the groups involved.[83]
+Arachnids lack some features of other chelicerates, including backward-pointing mouths and gnathobases ("jaw bases") at the bases of their legs;[83] both of these features are part of the ancestral arthropod feeding system.[84] Instead, they have mouths that point forwards and downwards, and all have some means of breathing air.[83] Spiders (Araneae) are distinguished from other arachnid groups by several characteristics, including spinnerets and, in males, pedipalps that are specially adapted for sperm transfer.[85]
+Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two infraorders, Mygalomorphae and Araneomorphae. Over 48,000 living species of spiders (order Araneae) have been identified and as of 2019 grouped into 120 families and about 4,100 genera by arachnologists.[1]
+The only living members of the primitive Mesothelae are the family Liphistiidae, found only in Southeast Asia, China, and Japan.[85] Most of the Liphistiidae construct silk-lined burrows with thin trapdoors, although some species of the genus Liphistius build camouflaged silk tubes with a second trapdoor as an emergency exit. Members of the genus Liphistius run silk "tripwires" outwards from their tunnels to help them detect approaching prey, while those of the genus Heptathela do not and instead rely on their built-in vibration sensors.[88] Spiders of the genus Heptathela have no venom glands, although they do have venom gland outlets on the fang tip.[89]
+The extinct families Arthrolycosidae, found in Carboniferous and Permian rocks, and Arthromygalidae, so far found only in Carboniferous rocks, have been classified as members of the Mesothelae.[90]
+The Mygalomorphae, which first appeared in the Triassic period,[79] are generally heavily built and ″hairy″, with large, robust chelicerae and fangs (technically, spiders do not have true hairs, but rather setae).[91][85] Well-known examples include tarantulas, ctenizid trapdoor spiders and the Australasian funnel-web spiders.[11] Most spend the majority of their time in burrows, and some run silk tripwires out from these, but a few build webs to capture prey. However, mygalomorphs cannot produce the pirifom silk that the Araneomorphae use as an instant adhesive to glue silk to surfaces or to other strands of silk, and this makes web construction more difficult for mygalomorphs. Since mygalomorphs rarely "balloon" by using air currents for transport, their populations often form clumps.[85] In addition to arthropods, some mygalomorphs are known to prey on frogs, small mammals, lizards, snakes, snails, and small birds.[92][93]
+In addition to accounting for over 90% of spider species, the Araneomorphae, also known as the "true spiders", include orb-web spiders, the cursorial wolf spiders, and jumping spiders,[85] as well as the only known herbivorous spider, Bagheera kiplingi.[49] They are distinguished by having fangs that oppose each other and cross in a pinching action, in contrast to the Mygalomorphae, which have fangs that are nearly parallel in alignment.[94]
+Although spiders are widely feared, only a few species are dangerous to people.[96] Spiders will only bite humans in self-defense, and few produce worse effects than a mosquito bite or bee sting.[97] Most of those with medically serious bites, such as recluse spiders (genus Loxosceles) and widow spiders (genus Latrodectus), would rather flee and bite only when trapped, although this can easily arise by accident.[98][99] The defensive tactics of Australian funnel-web spiders (family Atracidae) include fang display. Their venom, although they rarely inject much, has resulted in 13 attributed human deaths over 50 years.[100] They have been deemed to be the world's most dangerous spiders on clinical and venom toxicity grounds,[96] though this claim has also been attributed to the Brazilian wandering spider (genus Phoneutria).[101]
+There were about 100 reliably reported deaths from spider bites in the 20th century,[102] compared to about 1,500 from jellyfish stings.[103] Many alleged cases of spider bites may represent incorrect diagnoses,[104] which would make it more difficult to check the effectiveness of treatments for genuine bites.[105] A review published in 2016 agreed with this conclusion, showing that 78% of 134 published medical case studies of supposed spider bites did not meet the necessary criteria for a spider bite to be verified. In the case of the two genera with the highest reported number of bites, Loxosceles and Latrodectus, spider bites were not verified in over 90% of the reports. Even when verification had occurred, details of the treatment and its effects were often lacking.[106]
+Spider venoms may be a less polluting alternative to conventional pesticides, as they are deadly to insects but the great majority are harmless to vertebrates. Australian funnel web spiders are a promising source, as most of the world's insect pests have had no opportunity to develop any immunity to their venom, and funnel web spiders thrive in captivity and are easy to "milk". It may be possible to target specific pests by engineering genes for the production of spider toxins into viruses that infect species such as cotton bollworms.[107]
+The Ch'ol Maya use a beverage created from the tarantula species Brachypelma vagans for the treatment of a condition they term 'tarantula wind', the symptoms of which include chest pain, asthma and coughing.[108]
+Possible medical uses for spider venoms are being investigated, for the treatment of cardiac arrhythmia,[109] Alzheimer's disease,[110] strokes,[111] and erectile dysfunction.[112]
+The peptide GsMtx-4, found in the venom of Brachypelma vagans, is being researched to determine whether or not it could effectively be used for the treatment of cardiac arrhythmia, muscular dystrophy or glioma.[108]
+Because spider silk is both light and very strong, attempts are being made to produce it in goats' milk and in the leaves of plants, by means of genetic engineering.[113][114]
+Spiders can also be used as food. Cooked tarantulas are considered a delicacy in Cambodia,[115] and by the Piaroa Indians of southern Venezuela – provided the highly irritant bristles, the spiders' main defense system, are removed first.[116]
+Arachnophobia is a specific phobia—it is the abnormal fear of spiders or anything reminiscent of spiders, such as webs or spiderlike shapes. It is one of the most common specific phobias,[117][118] and some statistics show that 50% of women and 10% of men show symptoms.[119] It may be an exaggerated form of an instinctive response that helped early humans to survive,[120] or a cultural phenomenon that is most common in predominantly European societies.[121]
+Spiders have been the focus of stories and mythologies of various cultures for centuries.[122] Uttu, the ancient Sumerian goddess of weaving, was envisioned as a spider spinning her web.[123][124] According to her main myth, she resisted her father Enki's sexual advances by ensconcing herself in her web,[124] but let him in after he promised her fresh produce as a marriage gift,[124] thereby allowing him to intoxicate her with beer and rape her.[124] Enki's wife Ninhursag heard Uttu's screams and rescued her,[124] removing Enki's semen from her vagina and planting it in the ground to produce eight previously-nonexistent plants.[124] In a story told by the Roman poet Ovid in his Metamorphoses, Arachne was a Lydian girl who challenged the goddess Athena to a weaving contest.[125][126] Arachne won, but Athena destroyed her tapestry out of jealousy,[126][127] causing Arachne to hang herself.[126][127] In an act of mercy, Athena brought Arachne back to life as the first spider.[126][127] Stories about the trickster-spider Anansi are prominent in the folktales of West Africa and the Caribbean.[128]
+In some cultures, spiders have symbolized patience due to their hunting technique of setting webs and waiting for prey, as well as mischief and malice due to their venomous bites.[129] The Italian tarantella is a dance to rid the young woman of the lustful effects of a spider bite. Web-spinning also caused the association of the spider with creation myths, as they seem to have the ability to produce their own worlds.[130] Dreamcatchers are depictions of spiderwebs. The Moche people of ancient Peru worshipped nature.[131] They placed emphasis on animals and often depicted spiders in their art.[132]

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+The wild boar (Sus scrofa), also known as the "wild swine",[3] "common wild pig",[4] or simply "wild pig",[5] is a suid native to much of the Palearctic, as well as being introduced in the Nearctic, Neotropic, Oceania, the Caribbean islands, and Southeast Asia. Human intervention has spread it further, making the species one of the widest-ranging mammals in the world, as well as the most widespread suiform.[4] It has been assessed as least concern on the IUCN Red List due to its wide range, high numbers, and adaptability to a diversity of habitats.[1] It has become an invasive species in part of its introduced range. Wild boars probably originated in Southeast Asia during the Early Pleistocene[6] and outcompeted other suid species as they spread throughout the Old World.[7]
+As of 1990, up to 16 subspecies are recognized, which are divided into four regional groupings based on skull height and lacrimal bone length.[2] The species lives in matriarchal societies consisting of interrelated females and their young (both male and female). Fully grown males are usually solitary outside the breeding season.[8] The grey wolf is the wild boar's main predator in most of its natural range except in the Far East and the Lesser Sunda Islands, where it is replaced by the tiger and Komodo dragon respectively.[9][10] The wild boar has a long history of association with humans, having been the ancestor of most domestic pig breeds and a big-game animal for millennia. Boars have also re-hybridized in recent decades with feral pigs; these boar–pig hybrids have become a serious pest wild animal in the Americas and Australia.
+As true wild boars became extinct in Great Britain before the development of Modern English, the same terms are often used for both true wild boar and pigs, especially large or semi-wild ones. The English 'boar' stems from the Old English bar, which is thought to be derived from the West Germanic *bairaz, of unknown origin.[11] Boar is sometimes used specifically to refer to males, and may also be used to refer to male domesticated pigs, especially breeding males that have not been castrated.
+'Sow', the traditional name for a female, again comes from Old English and Germanic; it stems from Proto-Indo-European, and is related to the Latin: sus and Greek hus, and more closely to the New High German Sau. The young may be called 'piglets'.
+The animals' specific name scrofa is Latin for 'sow'.[12]
+In hunting terminology, boars are given different designations according to their age:[13]
+MtDNA studies indicate that the wild boar originated from islands in Southeast Asia such as Indonesia and the Philippines, and subsequently spread onto mainland Eurasia and North Africa.[6] The earliest fossil finds of the species come from both Europe and Asia, and date back to the Early Pleistocene.[14] By the late Villafranchian, S. scrofa largely displaced the related S. strozzii, a large, possibly swamp-adapted suid ancestral to the modern S. verrucosus throughout the Eurasian mainland, restricting it to insular Asia.[7] Its closest wild relative is the bearded pig of Malacca and surrounding islands.[3]
+As of 2005[update],[2] 16 subspecies are recognised, which are divided into four regional groupings:
+sahariensis (Heim de Balzac, 1937)
+nipponicus (Heude, 1899)
+mediterraneus (Ulmansky, 1911)
+reiseri (Bolkay, 1925)
+sardous (Ströbel, 1882)
+With the exception of domestic pigs in Timor and Papua New Guinea (which appear to be of Sulawesi warty pig stock), the wild boar is the ancestor of most pig breeds.[16][24] Archaeological evidence suggests that pigs were domesticated from wild boar as early as 13,000–12,700 BCE in the Near East in the Tigris Basin,[25] being managed in the wild in a way similar to the way they are managed by some modern New Guineans.[26] Remains of pigs have been dated to earlier than 11,400 BCE in Cyprus. Those animals must have been introduced from the mainland, which suggests domestication in the adjacent mainland by then.[27] There was also a separate domestication in China, which took place about 8,000 years ago.[28][29]
+DNA evidence from sub-fossil remains of teeth and jawbones of Neolithic pigs shows that the first domestic pigs in Europe had been brought from the Near East. This stimulated the domestication of local European wild boars, resulting in a third domestication event with the Near Eastern genes dying out in European pig stock. Modern domesticated pigs have involved complex exchanges, with European domesticated lines being exported in turn to the ancient Near East.[30][31] Historical records indicate that Asian pigs were introduced into Europe during the 18th and early 19th centuries.[28] Domestic pigs tend to have much more developed hindquarters than their wild boar ancestors, to the point where 70% of their body weight is concentrated in the posterior, which is the opposite of wild boar, where most of the muscles are concentrated on the head and shoulders.[32]
+The wild boar is a bulky, massively built suid with short and relatively thin legs. The trunk is short and robust, while the hindquarters are comparatively underdeveloped. The region behind the shoulder blades rises into a hump and the neck is short and thick to the point of being nearly immobile. The animal's head is very large, taking up to one-third of the body's entire length.[3] The structure of the head is well suited for digging. The head acts as a plough, while the powerful neck muscles allow the animal to upturn considerable amounts of soil:[33] it is capable of digging 8–10 cm (3.1–3.9 in) into frozen ground and can upturn rocks weighing 40–50 kg (88–110 lb).[9] The eyes are small and deep-set and the ears long and broad. The species has well developed canine teeth, which protrude from the mouths of adult males. The middle hooves are larger and more elongated than the lateral ones and are capable of quick movements.[3] The animal can run at a maximum speed of 40 km/h (25 mph) and jump at a height of 140–150 cm (55–59 in).[9]
+Sexual dimorphism is very pronounced in the species, with males being typically 5–10% larger and 20–30% heavier than females. Males also sport a mane running down the back, which is particularly apparent during autumn and winter.[34] The canine teeth are also much more prominent in males and grow throughout life. The upper canines are relatively short and grow sideways early in life, though they gradually curve upwards. The lower canines are much sharper and longer, with the exposed parts measuring 10–12 cm (3.9–4.7 in) in length. In the breeding period, males develop a coating of subcutaneous tissue, which may be 2–3 cm (0.79–1.18 in) thick, extending from the shoulder blades to the rump, thus protecting vital organs during fights. Males sport a roughly egg-sized sack near the opening of the penis, which collects urine and emits a sharp odour. The function of this sack is not fully understood.[3]
+Adult size and weight is largely determined by environmental factors; boars living in arid areas with little productivity tend to attain smaller sizes than their counterparts inhabiting areas with abundant food and water. In most of Europe, males average 75–100 kg (165–220 lb) in weight, 75–80 cm (30–31 in) in shoulder height and 150 cm (59 in) in body length, whereas females average 60–80 kg (130–180 lb) in weight, 70 cm (28 in) in shoulder height and 140 cm (55 in) in body length. In Europe's Mediterranean regions, males may reach average weights as low as 50 kg (110 lb) and females 45 kg (99 lb), with shoulder heights of 63–65 cm (25–26 in). In the more productive areas of Eastern Europe, males average 110–130 kg (240–290 lb) in weight, 95 cm (37 in) in shoulder height and 160 cm (63 in) in body length, while females weigh 95 kg (209 lb), reach 85–90 cm (33–35 in) in shoulder height and 145 cm (57 in) in body length. In Western and Central Europe, the largest males weigh 200 kg (440 lb) and females 120 kg (260 lb). In Northeastern Asia, large males can reach brown bear-like sizes, weighing 270 kg (600 lb) and measuring 110–118 cm (43–46 in) in shoulder height. Some adult males in Ussuriland and Manchuria have been recorded to weigh 300–350 kg (660–770 lb) and measure 125 cm (49 in) in shoulder height. Adults of this size are generally immune from wolf predation.[35] Such giants are rare in modern times, due to past overhunting preventing animals from attaining their full growth.[3]
+The winter coat consists of long, coarse bristles underlaid with short brown downy fur. The length of these bristles varies along the body, with the shortest being around the face and limbs and the longest running along the back. These back bristles form the aforementioned mane prominent in males and stand erect when the animal is agitated. Colour is highly variable; specimens around Lake Balkhash are very lightly coloured, and can even be white, while some boars from Belarus and Ussuriland can be black. Some subspecies sport a light-coloured patch running backward from the corners of the mouth. Coat colour also varies with age, with piglets having light brown or rusty-brown fur with pale bands extending from the flanks and back.[3]
+The wild boar produces a number of different sounds which are divided into three categories:
+Its sense of smell is very well developed to the point that the animal is used for drug detection in Germany.[37] Its hearing is also acute, though its eyesight is comparatively weak,[3] lacking color vision[37] and being unable to recognise a standing human 10–15 metres (33–49 ft) away.[9]
+Pigs are one of four known mammalian species which possess mutations in the nicotinic acetylcholine receptor that protect against snake venom. Mongooses, honey badgers, hedgehogs, and pigs all have modifications to the receptor pocket which prevents the snake venom α-neurotoxin from binding. These represent four separate, independent mutations.[38]
+Boars are typically social animals, living in female-dominated sounders consisting of barren sows and mothers with young led by an old matriarch. Male boars leave their sounder at the age of 8–15 months, while females either remain with their mothers or establish new territories nearby. Subadult males may live in loosely knit groups, while adult and elderly males tend to be solitary outside the breeding season.[8][a]
+The breeding period in most areas lasts from November to January, though most mating only lasts a month and a half. Prior to mating, the males develop their subcutaneous armour in preparation for confronting rivals. The testicles double in size and the glands secrete a foamy yellowish liquid. Once ready to reproduce, males travel long distances in search of a sounder of sows, eating little on the way. Once a sounder has been located, the male drives off all young animals and persistently chases the sows. At this point, the male fiercely fights potential rivals.[3] A single male can mate with 5–10 sows.[9] By the end of the rut, males are often badly mauled and have lost 20% of their body weight,[3] with bite-induced injuries to the penis being common.[40] The gestation period varies according to the age of the expecting mother. For first-time breeders, it lasts 114–130 days, while it lasts 133–140 days in older sows. Farrowing occurs between March and May, with litter sizes depending on the age and nutrition of the mother. The average litter consists of 4–6 piglets, with the maximum being 10–12.[3][b] The piglets are whelped in a nest constructed from twigs, grasses and leaves. Should the mother die prematurely, the piglets are adopted by the other sows in the sounder.[42]
+Newborn piglets weigh around 600–1,000 grams, lacking underfur and bearing a single milk incisor and canine on each half of the jaw.[3] There is intense competition between the piglets over the most milk-rich nipples, as the best-fed young grow faster and have stronger constitutions.[42] The piglets do not leave the lair for their first week of life. Should the mother be absent, the piglets lie closely pressed to each other. By two weeks of age, the piglets begin accompanying their mother on her journeys. Should danger be detected, the piglets take cover or stand immobile, relying on their camouflage to keep them hidden. The neonatal coat fades after three months, with adult colouration being attained at eight months. Although the lactation period lasts 2.5–3.5 months, the piglets begin displaying adult feeding behaviours at the age of 2–3 weeks. The permanent dentition is fully formed by 1–2 years. With the exception of the canines in males, the teeth stop growing during the middle of the fourth year. The canines in old males continue to grow throughout their lives, curving strongly as they age. Sows attain sexual maturity at the age of one year, with males attaining it a year later. However, estrus usually first occurs after two years in sows, while males begin participating in the rut after 4–5 years, as they are not permitted to mate by the older males.[3] The maximum lifespan in the wild is 10–14 years, though few specimens survive past 4–5 years.[43] Boars in captivity have lived for 20 years.[9]
+The wild boar inhabits a diverse array of habitats from boreal taigas to deserts.[3] In mountainous regions, it can even occupy alpine zones, occurring up to 1,900 m (6,200 ft) in the Carpathians, 2,600 m (8,500 ft) in the Caucasus and up to 3,600–4,000 m (11,800–13,100 ft) in the mountains in Central Asia and Kazakhstan.[3] In order to survive in a given area, wild boars require a habitat fulfilling three conditions: heavily brushed areas providing shelter from predators, water for drinking and bathing purposes and an absence of regular snowfall.[44]
+The main habitats favored by boars in Europe are deciduous and mixed forests, with the most favorable areas consisting of forest composed of oak and beech enclosing marshes and meadows. In the Białowieża Forest, the animal's primary habitat consists of well-developed broad-leaved and mixed forests, along with marshy mixed forests, with coniferous forests and undergrowths being of secondary importance. Forests made up entirely of oak groves and beeches are used only during the fruit-bearing season. This is in contrast to the Caucasian and Transcaucasian mountain areas, where boars will occupy such fruit-bearing forests year-round. In the mountainous areas of the Russian Far East, the species inhabits nutpine groves, hilly mixed forests where Mongolian oak and Korean pine are present, swampy mixed taiga and coastal oak forests. In Transbaikalia, boars are restricted to river valleys with nut pine and shrubs. Boars are regularly encountered in pistachio groves in winter in some areas of Tajikistan and Turkmenistan, while in spring they migrate to open deserts; boar have also colonized deserts in several areas they have been introduced to.[3][44][45]
+On the islands of Komodo and Rinca, the boar mostly inhabits savanna or open monsoon forests, avoiding heavily forested areas unless pursued by humans.[10] Wild boar are known to be competent swimmers, capable of covering long distances. In 2013, one boar was reported to have completed the 11-kilometre (7 mi) swim from France to Alderney in the Channel Islands. Due to concerns about disease, it was shot and incinerated.[46]
+Wild boar rest in shelters, which contain insulating material like spruce branches and dry hay. These resting places are occupied by whole families (though males lie separately) and are often located in the vicinity of streams, in swamp forests and in tall grass or shrub thickets. Boars never defecate in their shelters and will cover themselves with soil and pine needles when irritated by insects.[9]
+The wild boar is a highly versatile omnivore, whose diversity in choice of food is comparable to that of humans.[33] Their foods can be divided into four categories:
+A 50 kg (110 lb) boar needs around 4,000–4,500 calories of food per day, though this required amount increases during winter and pregnancy,[33] with the majority of its diet consisting of food items dug from the ground, like underground plant material and burrowing animals.[3] Acorns and beechnuts are invariably its most important food items in temperate zones,[47] as they are rich in the carbohydrates necessary for the buildup of fat reserves needed to survive lean periods.[33] In Western Europe, underground plant material favoured by boars includes bracken, willow herb, bulbs, meadow herb roots and bulbs and the bulbs of cultivated crops. Such food is favoured in early spring and summer, but may also be eaten in autumn and winter during beechnut and acorn crop failures. Should regular wild foods become scarce, boars will eat tree bark and fungi, as well as visit cultivated potato and artichoke fields.[3] Boar soil disturbance and foraging have been shown to facilitate invasive plants.[48][49] Boars of the vittatus subspecies in Ujung Kulon National Park in Java differ from most other populations by their primarily frugivorous diet, which consists of 50 different fruit species, especially figs, thus making them important seed dispersers.[4] The wild boar can consume numerous genera of poisonous plants without ill effect, including Aconitum, Anemone, Calla, Caltha, Ferula and Pteridium.[9]
+Boars may occasionally prey on small vertebrates like newborn deer fawns, leporids and galliform chicks.[33] Boars inhabiting the Volga Delta and near some lakes and rivers of Kazakhstan have been recorded to feed extensively on fish like carp and Caspian roach. Boars in the former area will also feed on cormorant and heron chicks, bivalved molluscs, trapped muskrats and mice.[3] There is at least one record of a boar killing and eating a bonnet macaque in southern India's Bandipur National Park, though this may have been a case of intraguild predation, brought on by interspecific competition for human handouts.[50]
+Piglets are vulnerable to attack from medium-sized felids like Eurasian lynx, jungle cats and snow leopards and other carnivorans like brown bears and yellow-throated martens.[3]
+The grey wolf is the main predator of wild boar throughout most of its range. A single wolf can kill around 50 to 80 boars of differing ages in one year.[3] In Italy[51] and Belarus' Belovezhskaya Pushcha National Park, boars are the wolf's primary prey, despite an abundance of alternative, less powerful ungulates.[51] Wolves are particularly threatening during the winter, when deep snow impedes the boars' movements. In the Baltic regions, heavy snowfall can allow wolves to eliminate boars from an area almost completely. Wolves primarily target piglets and subadults and only rarely attack adult sows. Adult males are usually avoided entirely.[3] Dholes may also prey on boars, to the point of keeping their numbers down in northwestern Bhutan, despite there being many more cattle in the area.[52]
+Leopards are predators of wild boar in the Caucasus (particularly Transcaucasia), the Russian Far East, India, China[53] and Iran. In most areas, boars constitute only a small part of the leopard's diet. However, in Iran's Sarigol National Park, boars are the second most frequently targeted prey species after mouflon, though adult individuals are generally avoided, as they are above the leopard's preferred weight range of 10–40 kg (22–88 lb).[54] This dependence on wild boar is largely due in part to the local leopard subspecies' large size.[55]
+Boars of all ages were once the primary prey of tigers in Transcaucasia, Kazakhstan, Middle Asia and the Far East up until the late 19th century. In modern times, tiger numbers are too low to have a limiting effect on boar populations. A single tiger can systematically destroy an entire sounder by preying on its members one by one, before moving on to another sounder. Tigers have been noted to chase boars for longer distances than with other prey. In two rare cases, boars were reported to gore a small tiger and a tigress to death in self-defense.[56] In the Amur region, wild boars are one of the two most important prey species for tigers alongside the Manchurian wapiti, with the two species collectively comprising roughly 80% of the felid's prey.[57] In Sikhote Alin, a tiger can kill 30–34 boars a year.[9] Studies of tigers in India indicate that boars are usually secondary in preference to various cervids and bovids,[58] though when boars are targeted, healthy adults are caught more frequently than young and sick specimens.[59]
+On the islands of Komodo, Rinca and Flores, the boar's main predator is the Komodo dragon.[10]
+The species originally occurred in North Africa and much of Eurasia; from the British Isles to Korea and the Sunda Islands. The northern limit of its range extended from southern Scandinavia to southern Siberia and Japan. Within this range, it was only absent in extremely dry deserts and alpine zones. It was once found in North Africa along the Nile valley up to Khartum and north of the Sahara. The species occurs on a few Ionian and Aegean Islands, sometimes swimming between islands.[60] The reconstructed northern boundary of the animal's Asian range ran from Lake Ladoga (at 60°N) through the area of Novgorod and Moscow into the southern Urals, where it reached 52°N. From there, the boundary passed Ishim and farther east the Irtysh at 56°N. In the eastern Baraba steppe (near Novosibirsk) the boundary turned steep south, encircled the Altai Mountains and went again eastward including the Tannu-Ola Mountains and Lake Baikal. From here, the boundary went slightly north of the Amur River eastward to its lower reaches at the Sea of Okhotsk. On Sakhalin, there are only fossil reports of wild boar. The southern boundaries in Europe and Asia were almost invariably identical to the seashores of these continents. It is absent in the dry regions of Mongolia from 44–46°N southward, in China westward of Sichuan and in India north of the Himalayas. It is absent in the higher elevations of the Pamir and the Tien Shan, though they do occur in the Tarim basin and on the lower slopes of the Tien Shan.[3]
+In recent centuries, the range of wild boar has changed dramatically, largely due to hunting by humans and more recently because of captive wild boar escaping into the wild. Prior to the 20th century, boar populations had declined in numerous areas, with British populations probably becoming extinct during the 13th century.[61] In the warm period after the ice age, wild boar lived in the southern parts of Sweden and Norway and north of Lake Ladoga in Karelia.[62] It was previously thought that the species did not live in Finland during prehistory because no prehistoric wild boar bones had been found within the borders of the country.[63][64] It was not until 2013, when a wild boar bone was found in Askola, that the species was found to have lived in Finland more than 8,000 years ago. It is believed, however, that man prevented its establishment by hunting.[65][66] In Denmark, the last boar was shot at the beginning of the 19th century, and by 1900 they were absent in Tunisia and Sudan and large areas of Germany, Austria and Italy. In Russia, they were extirpated in wide areas by the 1930s.[3] The last boar in Egypt reportedly died on 20 December 1912 in the Giza Zoo, with wild populations having disappeared by 1894–1902. Prince Kamal el Dine Hussein attempted to repopulate Wadi El Natrun with boars of Hungarian stock, but they were quickly exterminated by poachers.[67]
+A revival of boar populations began in the middle of the 20th century. By 1950, wild boar had once again reached their original northern boundary in many parts of their Asiatic range. By 1960, they reached Leningrad and Moscow and by 1975, they were to be found in Archangelsk and Astrakhan. In the 1970s they again occurred in Denmark and Sweden, where captive animals escaped and now survive in the wild. In England, wild boar populations re-established themselves in the 1990s, after escaping from specialist farms that had imported European stock.[61]
+Wild boars were apparently already becoming rare by the 11th century since a 1087 forestry law enacted by William the Conqueror punishes through blinding the unlawful killing of a boar. Charles I attempted to reintroduce the species into the New Forest, though this population was exterminated during the Civil War.
+Between their medieval extinction and the 1980s, when wild boar farming began, only a handful of captive wild boar, imported from the continent, were present in Britain. Occasional escapes of wild boar from wildlife parks have occurred as early as the 1970s, but since the early 1990s significant populations have re-established themselves after escapes from farms, the number of which has increased as the demand for meat from the species has grown. A 1998 MAFF (now DEFRA) study on wild boar living wild in Britain confirmed the presence of two populations of wild boar living in Britain; one in Kent/East Sussex and another in Dorset.[61] Another DEFRA report, in February 2008,[68] confirmed the existence of these two sites as 'established breeding areas' and identified a third in Gloucestershire/Herefordshire; in the Forest of Dean/Ross on Wye area. A 'new breeding population' was also identified in Devon. There is another significant population in Dumfries and Galloway. Populations estimates were as follows:
+Population estimates for the Forest of Dean are disputed as, at the time that the DEFRA population estimate was 100, a photo of a boar sounder in the forest near Staunton with over 33 animals visible was published and at about the same time over 30 boar were seen in a field near the original escape location of Weston under Penyard many kilometres or miles away. In early 2010 the Forestry Commission embarked on a cull,[70] with the aim of reducing the boar population from an estimated 150 animals to 100. By August it was stated that efforts were being made to reduce the population from 200 to 90, but that only 25 had been killed.[71] The failure to meet cull targets was confirmed in February 2011.[72]
+Wild boars have crossed the River Wye into Monmouthshire, Wales. Iolo Williams, the BBC Wales wildlife expert, attempted to film Welsh boar in late 2012.[73] Many other sightings, across the UK, have also been reported.[74] The effects of wild boar on the U.K.'s woodlands were discussed with Ralph Harmer of the Forestry Commission on the BBC Radio's Farming Today radio programme in 2011. The programme prompted activist writer George Monbiot to propose a thorough population study, followed by the introduction of permit-controlled culling.[75]
+Wild boars are an invasive species in the Americas and cause problems including out-competing native species for food, destroying the nests of ground-nesting species, killing fawns and young domestic livestock, destroying agricultural crops, eating tree seeds and seedlings, destroying native vegetation and wetlands through wallowing, damaging water quality, coming into violent conflict with humans and pets and carrying pig and human diseases including brucellosis, trichinosis and pseudorabies. In some jurisdictions, it is illegal to import, breed, release, possess, sell, distribute, trade, transport, hunt, or trap Eurasian boars. Hunting and trapping is done systematically, to increase the chance of eradication and to remove the incentive to illegally release boars, which have mostly been spread deliberately by sport hunters.[76]
+While domestic pigs, both captive and feral (popularly termed "razorbacks"), have been in North America since the earliest days of European colonization, pure wild boars were not introduced into the New World until the 19th century. The suids were released into the wild by wealthy landowners as big game animals. The initial introductions took place in fenced enclosures, though several escapes occurred, with the escapees sometimes intermixing with already established feral pig populations.
+The first of these introductions occurred in New Hampshire in 1890. Thirteen wild boars from Germany were purchased by Austin Corbin from Carl Hagenbeck and released into a 9,500-hectare (23,000-acre) game preserve in Sullivan County. Several of these boars escaped, though they were quickly hunted down by locals. Two further introductions were made from the original stocking, with several escapes taking place due to breaches in the game preserve's fencing. These escapees have ranged widely, with some specimens having been observed crossing into Vermont.[77]
+In 1902, 15–20 wild boar from Germany were released into a 3,200-hectare (7,900-acre) estate in Hamilton County, New York. Several specimens escaped six years later, dispersing into the William C. Whitney Wilderness Area, with their descendants surviving for at least 20 years.[77]
+The most extensive boar introduction in the US took place in western North Carolina in 1912, when 13 boars of undetermined European origin were released into two fenced enclosures in a game preserve in Hooper Bald, Graham County. Most of the specimens remained in the preserve for the next decade, until a large-scale hunt caused the remaining animals to break through their confines and escape. Some of the boars migrated to Tennessee, where they intermixed with both free-ranging and feral pigs in the area. In 1924, a dozen Hooper Bald wild pigs were shipped to California and released in a property between Carmel Valley and the Los Padres National Forest. These hybrid boar were later used as breeding stock on various private and public lands throughout the state, as well as in other states like Florida, Georgia, South Carolina, West Virginia and Mississippi.[77]
+Several wild boars from Leon Springs and the San Antonio, Saint Louis and San Diego Zoos were released in the Powder Horn Ranch in Calhoun County, Texas, in 1939. These specimens escaped and established themselves in surrounding ranchlands and coastal areas, with some crossing the Espiritu Santo Bay and colonizing Matagorda Island. Descendants of the Powder Horn Ranch boars were later released onto San José Island and the coast of Chalmette, Louisiana.[77]
+Wild boar of unknown origin were stocked in a ranch in the Edwards Plateau in the 1940s, only to escape during a storm and hybridize with local feral pig populations, later spreading into neighboring counties.[77]
+Starting in the mid-1980s, several boars purchased from the San Diego Zoo and Tierpark Berlin were released into the United States. A decade later, more specimens from farms in Canada and Białowieża Forest were let loose. In recent years, wild pig populations have been reported in 44 states within the US, most of which are likely wild boar–feral hog hybrids. Pure wild boar populations may still be present, but are extremely localized.[77]
+Wild boars are known to host at least 20 different parasitic worm species, with maximum infections occurring in summer. Young animals are vulnerable to helminths like Metastrongylus, which are consumed by boars through earthworms and cause death by parasitising the lungs. Wild boar also carry parasites known to infect humans, including Gastrodiscoides, Trichinella spiralis, Taenia solium, Balantidium coli and Toxoplasma gondii.[78] Wild boar in southern regions are frequently infested with ticks (Dermacentor, Rhipicephalus, and Hyalomma) and hog lice. The species also suffers from blood-sucking flies, which it escapes by bathing frequently or hiding in dense shrubs.[3]
+Swine plague spreads very quickly in wild boar, with epizootics being recorded in Germany, Poland, Hungary, Belarus, the Caucasus, the Far East, Kazakhstan and other regions. Foot-and-mouth disease can also take on epidemic proportions in boar populations. The species occasionally, but rarely contracts Pasteurellosis, hemorrhagic sepsis, tularemia, and anthrax. Wild boar may on occasion contract swine erysipelas through rodents or hog lice and ticks.[3]
+The wild boar features prominently in the cultures of Indo-European people, many of which saw the animal as embodying warrior virtues. Cultures throughout Europe and Asia Minor saw the killing of a boar as proof of one's valor and strength. Neolithic hunter gatherers depicted reliefs of ferocious wild boars on their temple pillars at Göbekli Tepe some 11,600 years ago.[80][81] Virtually all heroes in Greek mythology fight or kill a boar at one point. The demigod Herakles' third labour involves the capture of the Erymanthian Boar, Theseus slays the wild sow Phaea, and a disguised Odysseus is recognised by his handmaiden Eurycleia by the scars inflicted on him by a boar during a hunt in his youth.[82] To the mythical Hyperboreans, the boar represented spiritual authority.[79] Several Greek myths use the boar as a symbol of darkness, death and winter. One example is the story of the youthful Adonis, who is killed by a boar and is permitted by Zeus to depart from Hades only during the spring and summer period. This theme also occurs in Irish and Egyptian mythology, where the animal is explicitly linked to the month of October, therefore autumn. This association likely arose from aspects of the boar's actual nature. Its dark colour was linked to the night, while its solitary habits, proclivity to consume crops and nocturnal nature were associated with evil.[83] The foundation myth of Ephesus has the city being built over the site where Prince Androklos of Athens killed a boar.[84] Boars were frequently depicted on Greek funerary monuments alongside lions, representing gallant losers who have finally met their match, as opposed to victorious hunters as lions are. The theme of the doomed, yet valorous boar warrior also occurred in Hittite culture, where it was traditional to sacrifice a boar alongside a dog and a prisoner of war after a military defeat.[82]
+The boar as a warrior also appears in Scandinavian, Germanic and Anglo-Saxon culture, with its image having been frequently engraved on helmets, shields and swords. According to Tacitus, the Baltic Aesti featured boars on their helmets and may have also worn boar masks (see for example the Guilden Morden boar). The boar and pig were held in particularly high esteem by the Celts, who considered them to be their most important sacred animal. Some Celtic deities linked to boars include Moccus and Veteris. It has been suggested that some early myths surrounding the Welsh hero Culhwch involved the character being the son of a boar god.[82] Nevertheless, the importance of the boar as a culinary item among Celtic tribes may have been exaggerated in popular culture by the Asterix series, as wild boar bones are rare among Celtic archaeological sites and the few that do occur show no signs of butchery, having probably been used in sacrificial rituals.[85]
+The boar also appears in Vedic mythology and Hindu mythology. A story present in the Brahmanas has the god Indra slaying an avaricious boar, who has stolen the treasure of the asuras, then giving its carcass to the god Vishnu, who offered it as a sacrifice to the gods. In the story's retelling in the Charaka Samhita, the boar is described as a form of Prajapati and is credited with having raised the Earth from the primeval waters. In the Ramayana and the Puranas, the same boar is portrayed as Varaha, an avatar of Vishnu.[86]
+In Japanese culture, the boar is widely seen as a fearsome and reckless animal, to the point that several words and expressions in Japanese referring to recklessness include references to boars. The boar is the last animal of the Oriental zodiac, with people born during the year of the Pig being said to embody the boar-like traits of determination and impetuosity. Among Japanese hunters, the boar's courage and defiance is a source of admiration and it is not uncommon for hunters and mountain people to name their sons after the animal inoshishi (猪). Boars are also seen as symbols of fertility and prosperity; in some regions, it is thought that boars are drawn to fields owned by families including pregnant women, and hunters with pregnant wives are thought to have greater chances of success when boar hunting. The animal's link to prosperity was illustrated by its inclusion on the ¥10 note during the Meiji period and it was once believed that a man could become wealthy by keeping a clump of boar hair in his wallet.[87]
+In the folklore of the Mongol Altai Uriankhai tribe, the wild boar was associated with the watery underworld, as it was thought that the spirits of the dead entered the animal's head, to be ultimately transported to the water.[88] Prior to the conversion to Islam, the Kyrgyz people believed that they were descended from boars and thus did not eat pork. In Buryat mythology, the forefathers of the Buryats descended from heaven and were nourished by a boar.[89] In China, the boar is the emblem of the Miao people.[79]
+The boar (sanglier) is frequently displayed in English, Scottish and Welsh heraldry. As with the lion, the boar is often shown as armed and langued. As with the bear, Scottish and Welsh heraldry displays the boar's head with the neck cropped, unlike the English version, which retains the neck.[90] The white boar served as the badge of King Richard III of England, who distributed it among his northern retainers during his tenure as Duke of Gloucester.[91]
+Humans have been hunting boar for millennia, the earliest artistic depictions of such activities dating back to the Upper Paleolithic.[82] The animal was seen as a source of food among the Ancient Greeks, as well as a sporting challenge and source of epic narratives. The Romans inherited this tradition, with one of its first practitioners being Scipio Aemilianus. Boar hunting became particularly popular among the young nobility during the 3rd century BC as preparation for manhood and battle. A typical Roman boar hunting tactic involved surrounding a given area with large nets, then flushing the boar with dogs and immobilizing it with smaller nets. The animal would then be dispatched with a venabulum, a short spear with a crossguard at the base of the blade. More than their Greek predecessors, the Romans extensively took inspiration from boar hunting in their art and sculpture. With the ascension of Constantine the Great, boar hunting took on Christian allegorical themes, with the animal being portrayed as a "black beast" analogous to the dragon of Saint George. Boar hunting continued after the fall of the Western Roman Empire, though the Germanic tribes considered the red deer to be a more noble and worthy quarry. The post-Roman nobility hunted boar as their predecessors did, but primarily as training for battle rather than sport. It was not uncommon for medieval hunters to deliberately hunt boars during the breeding season when the animals were more aggressive. During the Renaissance, when deforestation and the introduction of firearms reduced boar numbers, boar hunting became the sole prerogative of the nobility, one of many charges brought up against the rich during the German Peasants' War and the French Revolution.[92] During the mid-20th century, 7,000–8,000 boars were caught in the Caucasus, 6,000–7,000 in Kazakhstan and about 5,000 in Central Asia during the Soviet period, primarily through the use of dogs and beats.[3] In Nepal, farmers and poachers eliminate boars by baiting balls of wheat flour containing explosives with kerosene oil, with the animals' chewing motions triggering the devices.[93]
+Wild boar can thrive in captivity, though piglets grow slowly and poorly without their mothers. Products derived from wild boar include meat, hide and bristles.[3] Apicius devotes a whole chapter to the cooking of boar meat, providing 10 recipes involving roasting, boiling and what sauces to use. The Romans usually served boar meat with garum.[94] Boar's head was the centrepiece of most medieval Christmas celebrations among the nobility.[95] Although growing in popularity as a captive-bred source of food, the wild boar takes longer to mature than most domestic pigs and it is usually smaller and produces less meat. Nevertheless, wild boar meat is leaner and healthier than pork,[96] being of higher nutritional value and having a much higher concentration of essential amino acids.[97] Most meat-dressing organizations agree that a boar carcass should yield 50 kg (110 lb) of meat on average. Large specimens can yield 15–20 kg (33–44 lb) of fat, with some giants yielding 30 kg (66 lb) or more. A boar hide can measure 300 dm2 (4,700 sq in) and can yield 350–1,000 grams (12–35 oz) of bristle and 400 grams (14 oz) of underwool.[3]
+Roman relief of a dog confronting a boar, Cologne
+Southern Indian depiction of boar hunt, c. 1540
+Pig-sticking in British India
+Boar shot in Volgograd Oblast, Russia
+The Boar Hunt – Hans Wertinger, c. 1530, the Danube Valley
+Boars can be damaging to agriculture in situations where their natural habitat is sparse. Populations living on the outskirts of towns or farms can dig up potatoes and damage melons, watermelons and maize. However, they generally only encroach upon farms when natural food is scarce. In the Belovezh forest for example, 34–47% of the local boar population will enter fields in years of moderate availability of natural foods. While the role of boars in damaging crops is often exaggerated,[3] cases are known of boar depredations causing famines, as was the case in Hachinohe, Japan in 1749, where 3,000 people died of what became known as the "wild boar famine". Still, within Japanese culture, the boar's status as vermin is expressed through its title as "king of pests" and the popular saying (addressed to young men in rural areas) "When you get married, choose a place with no wild boar."[87][98]
+In Central Europe, farmers typically repel boars through distraction or fright, while in Kazakhstan it is usual to employ guard dogs in plantations. Although large boar populations can play an important role in limiting forest growth, they are also useful in keeping pest populations such as June bugs under control.[3] The growth of urban areas and the corresponding decline in natural boar habitats has led to some sounders entering human habitations in search of food. As in natural conditions, sounders in peri-urban areas are matriarchal, though males tend to be much less represented and adults of both sexes can be up to 35% heavier than their forest-dwelling counterparts. As of 2010, at least 44 cities in 15 countries have experienced problems of some kind relating to the presence of habituated wild boar.[99]
+Actual attacks on humans are rare, but can be serious, resulting in penetrating injuries to the lower part of the body. They generally occur during the boars' rutting season from November to January, in agricultural areas bordering forests or on paths leading through forests. The animal typically attacks by charging and pointing its tusks towards the intended victim, with most injuries occurring on the thigh region. Once the initial attack is over, the boar steps back, takes position and attacks again if the victim is still moving, only ending once the victim is completely incapacitated.[100][101]
+Boar attacks on humans have been documented since the Stone Age, with one of the oldest depictions being a cave painting in Bhimbetaka, India. The Romans and Ancient Greeks wrote of these attacks (Odysseus was wounded by a boar and Adonis was killed by one). A 2012 study compiling recorded attacks from 1825–2012 found accounts of 665 human victims of both wild boars and feral pigs, with the majority (19%) of attacks in the animal's native range occurring in India. Most of the attacks occurred in rural areas during the winter months in non-hunting contexts and were committed by solitary males.[102]

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+Animals (also called Metazoa) are multicellular eukaryotic organisms that form the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and grow from a hollow sphere of cells, the blastula, during embryonic development. Over 1.5 million living animal species have been described—of which around 1 million are insects—but it has been estimated there are over 7 million animal species in total. Animals range in length from 8.5 micrometres (0.00033 in) to 33.6 metres (110 ft). They have complex interactions with each other and their environments, forming intricate food webs. The kingdom Animalia includes humans but in colloquial use the term animal often refers only to non-human animals. The scientific study of animals is known as zoology.
+Most living animal species are in Bilateria, a clade whose members have a bilaterally symmetric body plan. The Bilateria include the protostomes—in which many groups of invertebrates are found, such as nematodes, arthropods, and molluscs—and the deuterostomes, containing both the echinoderms as well as the chordates, the latter containing the vertebrates. Life forms interpreted as early animals were present in the Ediacaran biota of the late Precambrian. Many modern animal phyla became clearly established in the fossil record as marine species during the Cambrian explosion, which began around 542 million years ago. 6,331 groups of genes common to all living animals have been identified; these may have arisen from a single common ancestor that lived 650 million years ago.
+Historically, Aristotle divided animals into those with blood and those without. Carl Linnaeus created the first hierarchical biological classification for animals in 1758 with his Systema Naturae, which Jean-Baptiste Lamarck expanded into 14 phyla by 1809. In 1874, Ernst Haeckel divided the animal kingdom into the multicellular Metazoa (synonymous for Animalia) and the Protozoa, single-celled organisms no longer considered animals. In modern times, the biological classification of animals relies on advanced techniques, such as molecular phylogenetics, which are effective at demonstrating the evolutionary relationships between animal taxa.
+Humans make use of many other animal species, such as for food (including meat, milk, and eggs), for materials (such as leather and wool), and also as pets, and for transports, as working animals. Dogs have been used in hunting, while many terrestrial and aquatic animals were hunted for sports. Non-human animals have appeared in art from the earliest times and are featured in mythology and religion.
+The word "animal" comes from the Latin animalis, meaning having breath, having soul or living being.[1] The biological definition includes all members of the kingdom Animalia.[2] In colloquial usage, as a consequence of anthropocentrism, the term animal is sometimes used nonscientifically to refer only to non-human animals.[3][4][5][6]
+Animals have several characteristics that set them apart from other living things. Animals are eukaryotic and multicellular,[7][8] unlike bacteria, which are prokaryotic, and unlike protists, which are eukaryotic but unicellular. Unlike plants and algae, which produce their own nutrients[9] animals are heterotrophic,[8][10] feeding on organic material and digesting it internally.[11] With very few exceptions, animals respire aerobically.[12] All animals are motile[13] (able to spontaneously move their bodies) during at least part of their life cycle, but some animals, such as sponges, corals, mussels, and barnacles, later become sessile. The blastula is a stage in embryonic development that is unique to most animals,[14] allowing cells to be differentiated into specialised tissues and organs.
+All animals are composed of cells, surrounded by a characteristic extracellular matrix composed of collagen and elastic glycoproteins.[15] During development, the animal extracellular matrix forms a relatively flexible framework upon which cells can move about and be reorganised, making the formation of complex structures possible. This may be calcified, forming structures such as shells, bones, and spicules.[16] In contrast, the cells of other multicellular organisms (primarily algae, plants, and fungi) are held in place by cell walls, and so develop by progressive growth.[17] Animal cells uniquely possess the cell junctions called tight junctions, gap junctions, and desmosomes.[18]
+With few exceptions—in particular, the sponges and placozoans—animal bodies are differentiated into tissues.[19] These include muscles, which enable locomotion, and nerve tissues, which transmit signals and coordinate the body. Typically, there is also an internal digestive chamber with either one opening (in Ctenophora, Cnidaria, and flatworms) or two openings (in most bilaterians).[20]
+Nearly all animals make use of some form of sexual reproduction.[21] They produce haploid gametes by meiosis; the smaller, motile gametes are spermatozoa and the larger, non-motile gametes are ova.[22] These fuse to form zygotes,[23] which develop via mitosis into a hollow sphere, called a blastula. In sponges, blastula larvae swim to a new location, attach to the seabed, and develop into a new sponge.[24] In most other groups, the blastula  undergoes more complicated rearrangement.[25] It first invaginates to form a gastrula with a digestive chamber and two separate germ layers, an external ectoderm and an internal endoderm.[26] In most cases, a third germ layer, the mesoderm, also develops between them.[27] These germ layers then differentiate to form tissues and organs.[28]
+Repeated instances of mating with a close relative during sexual reproduction generally leads to inbreeding depression within a population due to the increased prevalence of harmful recessive traits.[29][30] Animals have evolved numerous mechanisms for avoiding close inbreeding.[31]
+Some animals are capable of asexual reproduction, which often results in a genetic clone of the parent. This may take place through fragmentation; budding, such as in Hydra and other cnidarians; or parthenogenesis, where fertile eggs are produced without mating, such as in aphids.[32][33]
+Animals are categorised into ecological groups depending on how they obtain or consume organic material, including carnivores, herbivores, omnivores, detritivores,[34] and parasites.[35] Interactions between animals form complex food webs. In carnivorous or omnivorous species, predation is a consumer-resource interaction where a predator feeds on another organism (called its prey).[36] Selective pressures imposed on one another lead to an evolutionary arms race between predator and prey, resulting in various anti-predator adaptations.[37][38] Almost all multicellular predators are animals.[39] Some consumers use multiple methods; for example, in parasitoid wasps, the larvae feed on the hosts' living tissues, killing them in the process,[40] but the adults primarily consume nectar from flowers.[41] Other animals may have very specific feeding behaviours, such as hawksbill sea turtles primarily eating sponges.[42]
+Most animals rely on the biomass and energy produced by plants through photosynthesis. Herbivores eat plant material directly, while carnivores, and other animals on higher trophic levels typically acquire it indirectly by eating other animals. Animals oxidize carbohydrates, lipids, proteins, and other biomolecules to unlock the chemical energy of molecular oxygen,[43] which allows the animal to grow and to sustain biological processes such as locomotion.[44][45][46] Animals living close to hydrothermal vents and cold seeps on the dark sea floor consume organic matter of archaea and bacteria produced in these locations through chemosynthesis (by oxidizing inorganic compounds, such as hydrogen sulfide).[47]
+Animals originally evolved in the sea. Lineages of arthropods colonised land around the same time as land plants, probably between 510–471 million years ago during the Late Cambrian or Early Ordovician.[48] Vertebrates such as the lobe-finned fish Tiktaalik started to move on to land in the late Devonian, about 375 million years ago.[49][50] Animals occupy virtually all of earth's habitats and microhabitats, including salt water, hydrothermal vents, fresh water, hot springs, swamps, forests, pastures, deserts, air, and the interiors of animals, plants, fungi and rocks.[51] Animals are however not particularly heat tolerant; very few of them can survive at constant temperatures above 50 °C (122 °F).[52] Only very few species of animals (mostly nematodes) inhabit the most extreme cold deserts of continental Antarctica.[53]
+The blue whale (Balaenoptera musculus) is the largest animal that has ever lived, weighing up to at least 190 tonnes and measuring up to 33.6 metres (110 ft) long.[54][55][56] The largest extant terrestrial animal is the African bush elephant (Loxodonta africana), weighing up to 12.25 tonnes[54] and measuring up to 10.67 metres (35.0 ft) long.[54] The largest terrestrial animals that ever lived were titanosaur sauropod dinosaurs such as Argentinosaurus, which may have weighed as much as 73 tonnes.[57] Several animals are microscopic; some Myxozoa (obligate parasites within the Cnidaria) never grow larger than 20 µm,[58] and one of the smallest species (Myxobolus shekel) is no more than 8.5 µm when fully grown.[59]
+The following table lists estimated numbers of described extant species for the animal groups with the largest numbers of species,[60] along with their principal habitats (terrestrial, fresh water,[61] and marine),[62] and free-living or parasitic ways of life.[63] Species estimates shown here are based on numbers described scientifically; much larger estimates have been calculated based on various means of prediction, and these can vary wildly. For instance, around 25,000–27,000 species of nematodes have been described, while published estimates of the total number of nematode species include 10,000–20,000; 500,000; 10 million; and 100 million.[64] Using patterns within the taxonomic hierarchy, the total number of animal species—including those not yet described—was calculated to be about 7.77 million in 2011.[65][66][a]
+3,000–6,500[74]
+4,000–25,000[74]
+The first fossils that might represent animals appear in the 665-million-year-old rocks of the Trezona Formation of South Australia. These fossils are interpreted as most probably being early sponges.[78]
+The oldest animals are found in the Ediacaran biota, towards the end of the Precambrian, around 610 million years ago. It had long been doubtful whether these included animals,[79][80][81] but the discovery of the animal lipid cholesterol in fossils of Dickinsonia establishes that these were indeed animals.[77] Animals are thought to have originated under low-oxygen conditions, suggesting that they were capable of living entirely by anaerobic respiration, but as they became specialized for aerobic metabolism they became fully dependent on oxygen in their environments.[82]
+Many animal phyla first appear in the fossil record during the Cambrian explosion, starting about 542 million years ago, in beds such as the Burgess shale. Extant phyla in these rocks include molluscs, brachiopods, onychophorans, tardigrades, arthropods, echinoderms and hemichordates, along with numerous now-extinct forms such as the predatory Anomalocaris. The apparent suddenness of the event may however be an artefact of the fossil record, rather than showing that all these animals appeared simultaneously.[83][84][85][86]
+Some palaeontologists have suggested that animals appeared much earlier than the Cambrian explosion, possibly as early as 1 billion years ago.[87] Trace fossils such as tracks and burrows found in the Tonian period may indicate the presence of triploblastic worm-like animals, roughly as large (about 5 mm wide) and complex as earthworms.[88] However, similar tracks are produced today by the giant single-celled protist Gromia sphaerica, so the Tonian trace fossils may not indicate early animal evolution.[89][90] Around the same time, the layered mats of microorganisms called stromatolites decreased in diversity, perhaps due to grazing by newly-evolved animals.[91]
+Animals are monophyletic, meaning they are derived from a common ancestor. Animals are sister to the Choanoflagellata, with which they form the Choanozoa.[92] The most basal animals, the Porifera, Ctenophora, Cnidaria, and Placozoa, have body plans that lack bilateral symmetry. Their relationships are still disputed; the sister group to all other animals could be the Porifera or the Ctenophora, both of which lack hox genes, important in body plan development.[93]
+These genes are found in the Placozoa[94][95] and the higher animals, the Bilateria.[96][97] 6,331 groups of genes common to all living animals have been identified; these may have arisen from a single common ancestor that lived 650 million years ago in the Precambrian. 25 of these are novel core gene groups, found only in animals; of those, 8 are for essential components of the Wnt and TGF-beta signalling pathways which may have enabled animals to become multicellular by providing a pattern for the body's system of axes (in three dimensions), and another 7 are for transcription factors including homeodomain proteins involved in the control of development.[98][99]
+The phylogenetic tree (of major lineages only) indicates approximately how many millions of years ago (mya) the lineages split.[100][101][102][103][104]
+Choanoflagellata
+Porifera
+Ctenophora
+Placozoa
+Cnidaria
+Xenacoelomorpha
+Chordata
+Ambulacraria
+Scalidophora
+Arthropoda and allies
+Nematoda and allies
+Rotifera and allies
+Chaetognatha
+Platyhelminthes and allies
+Mollusca and allies
+Annelida and allies
+Several animal phyla lack bilateral symmetry. Among these, the sponges (Porifera) probably diverged first, representing the oldest animal phylum.[105] Sponges lack the complex organization found in most other animal phyla;[106] their cells are differentiated, but in most cases not organised into distinct tissues.[107] They typically feed by drawing in water through pores.[108]
+The Ctenophora (comb jellies) and Cnidaria (which includes jellyfish, sea anemones, and corals) are radially symmetric and have digestive chambers with a single opening, which serves as both mouth and anus.[109] Animals in both phyla have distinct tissues, but these are not organised into organs.[110] They are diploblastic, having only two main germ layers, ectoderm and endoderm.[111] The tiny placozoans are similar, but they do not have a permanent digestive chamber.[112][113]
+The remaining animals, the great majority—comprising some 29 phyla and over a million species—form a clade, the Bilateria. The body is triploblastic, with three well-developed germ layers, and their tissues form distinct organs. The digestive chamber has two openings, a mouth and an anus, and there is an internal body cavity, a coelom or pseudocoelom. Animals with this bilaterally symmetric body plan and a tendency to move in one direction have a head end (anterior) and a tail end (posterior) as well as a back (dorsal) and a belly (ventral); therefore they also have a left side and a right side.[114][115]
+Having a front end means that this part of the body encounters stimuli, such as food, favouring cephalisation, the development of a head with sense organs and a mouth. Many bilaterians have a combination of circular muscles that constrict the body, making it longer, and an opposing set of longitudinal muscles, that shorten the body;[115] these enable soft-bodied animals with a hydrostatic skeleton to move by peristalsis.[116] They also have a gut that extends through the basically cylindrical body from mouth to anus. Many bilaterian phyla have primary larvae which swim with cilia and have an apical organ containing sensory cells. However, there are exceptions to each of these characteristics; for example, adult echinoderms are radially symmetric (unlike their larvae), while some parasitic worms have extremely simplified body structures.[114][115]
+Genetic studies have considerably changed zoologists' understanding of the relationships within the Bilateria. Most appear to belong to two major lineages, the  protostomes and the deuterostomes.[117] The basalmost bilaterians are the Xenacoelomorpha.[118][119][120]
+Protostomes and deuterostomes differ in several ways. Early in development, deuterostome embryos undergo radial cleavage during cell division, while many protostomes (the Spiralia) undergo spiral cleavage.[121]
+Animals from both groups possess a complete digestive tract, but in protostomes the first opening of the embryonic gut develops into the mouth, and the anus forms secondarily. In deuterostomes, the anus forms first while the mouth develops secondarily.[122][123] Most protostomes have schizocoelous development, where cells simply fill in the interior of the gastrula to form the mesoderm. In deuterostomes, the mesoderm forms by enterocoelic pouching, through invagination of the endoderm.[124]
+The main deuterostome phyla are the Echinodermata and the Chordata.[125] Echinoderms are exclusively marine and include starfish, sea urchins, and sea cucumbers.[126] The chordates are dominated by the vertebrates (animals with backbones),[127] which consist of fishes, amphibians, reptiles, birds, and mammals.[128] The deuterostomes also include the Hemichordata (acorn worms).[129][130]
+The Ecdysozoa are protostomes, named after their shared trait of ecdysis, growth by moulting.[131] They include the largest animal phylum, the Arthropoda, which contains insects, spiders, crabs, and their kin. All of these have a body divided into repeating segments, typically with paired appendages. Two smaller phyla, the Onychophora and Tardigrada, are close relatives of the arthropods and share these traits. The ecdysozoans also include the Nematoda or roundworms, perhaps the second largest animal phylum. Roundworms are typically microscopic, and occur in nearly every environment where there is water;[132] some are important parasites.[133] Smaller phyla related to them are the Nematomorpha or horsehair worms, and the Kinorhyncha, Priapulida, and Loricifera. These groups have a reduced coelom, called a pseudocoelom.[134]
+The Spiralia are a large group of protostomes that develop by spiral cleavage in the early embryo.[135] The Spiralia's phylogeny has been disputed, but it contains a large clade, the superphylum Lophotrochozoa, and smaller groups of phyla such as the Rouphozoa which includes the gastrotrichs and the flatworms. All of these are grouped as the Platytrochozoa, which has a sister group, the Gnathifera, which includes the rotifers.[136][137]
+The Lophotrochozoa includes the molluscs, annelids, brachiopods, nemerteans, bryozoa and entoprocts.[136][138][139] The molluscs, the second-largest animal phylum by number of described species, includes snails, clams, and squids, while the annelids are the segmented worms, such as earthworms, lugworms, and leeches. These two groups have long been considered close relatives because they share trochophore larvae.[140][141]
+In the classical era, Aristotle divided animals,[d] based on his own observations, into those with blood (roughly, the vertebrates) and those without. The animals were then arranged on a scale from man (with blood, 2 legs, rational soul) down through the live-bearing tetrapods (with blood, 4 legs, sensitive soul) and other groups such as crustaceans (no blood, many legs, sensitive soul) down to spontaneously-generating creatures like sponges (no blood, no legs, vegetable soul). Aristotle was uncertain whether sponges were animals, which in his system ought to have sensation, appetite, and locomotion, or plants, which did not: he knew that sponges could sense touch, and would contract if about to be pulled off their rocks, but that they were rooted like plants and never moved about.[143]
+In 1758, Carl Linnaeus created the first hierarchical classification in his Systema Naturae.[144] In his original scheme, the animals were one of three kingdoms, divided into the classes of Vermes, Insecta, Pisces, Amphibia, Aves, and Mammalia. Since then the last four have all been subsumed into a single phylum, the Chordata, while his Insecta (which included the crustaceans and arachnids) and Vermes have been renamed or broken up. The process was begun in 1793 by Jean-Baptiste de Lamarck, who called the Vermes une espèce de chaos (a chaotic mess)[e] and split the group into three new phyla, worms, echinoderms, and polyps (which contained corals and jellyfish). By 1809, in his Philosophie Zoologique, Lamarck had created 9 phyla apart from vertebrates (where he still had 4 phyla: mammals, birds, reptiles, and fish) and molluscs, namely cirripedes, annelids, crustaceans, arachnids, insects, worms, radiates, polyps, and infusorians.[142]
+In his 1817 Le Règne Animal, Georges Cuvier used comparative anatomy to group the animals into four embranchements ("branches" with different body plans, roughly corresponding to phyla), namely vertebrates, molluscs, articulated animals (arthropods and annelids), and zoophytes (radiata) (echinoderms, cnidaria and other forms).[146] This division into four was followed by the embryologist Karl Ernst von Baer in 1828, the zoologist Louis Agassiz in 1857, and the comparative anatomist Richard Owen in 1860.[147]
+In 1874, Ernst Haeckel divided the animal kingdom into two subkingdoms: Metazoa (multicellular animals, with five phyla: coelenterates, echinoderms, articulates, molluscs, and vertebrates) and Protozoa (single-celled animals), including a sixth animal phylum, sponges.[148][147] The protozoa were later moved to the former kingdom Protista, leaving only the Metazoa as a synonym of Animalia.[149]
+The human population exploits a large number of other animal species for food, both of domesticated livestock species in animal husbandry and, mainly at sea, by hunting wild species.[150][151] Marine fish of many species are caught commercially for food. A smaller number of species are farmed commercially.[150][152][153]
+Invertebrates including cephalopods, crustaceans, and bivalve or gastropod molluscs are hunted or farmed for food.[154]
+Chickens, cattle, sheep, pigs and other animals are raised as livestock for meat across the world.[151][155][156]
+Animal fibres such as wool are used to make textiles, while animal sinews have been used as lashings and bindings, and leather is widely used to make shoes and other items. Animals have been hunted and farmed for their fur to make items such as coats and hats.[157][158] Dyestuffs including carmine (cochineal),[159][160] shellac,[161][162] and kermes[163][164] have been made from the bodies of insects. Working animals including cattle and horses have been used for work and transport from the first days of agriculture.[165]
+Animals such as the fruit fly Drosophila melanogaster serve a major role in science as experimental models.[166][167][168][169] Animals have been used to create vaccines since their discovery in the 18th century.[170] Some medicines such as the cancer drug Yondelis are based on toxins or other molecules of animal origin.[171]
+People have used hunting dogs to help chase down and retrieve animals,[172] and birds of prey to catch birds and mammals,[173] while tethered cormorants have been used to catch fish.[174] Poison dart frogs have been used to poison the tips of blowpipe darts.[175][176]
+A wide variety of animals are kept as pets, from invertebrates such as tarantulas and octopuses, insects including praying mantises,[177] reptiles such as snakes and chameleons,[178] and birds including canaries, parakeets, and parrots[179] all finding a place. However, the most kept pet species are mammals, namely dogs, cats, and rabbits.[180][181][182] There is a tension between the role of animals as companions to humans, and their existence as individuals with rights of their own.[183]
+A wide variety of terrestrial and aquatic animals are hunted for sport.[184]
+Animals have been the subjects of art from the earliest times, both historical, as in Ancient Egypt, and prehistoric, as in the cave paintings at Lascaux. Major animal paintings include Albrecht Dürer's 1515 The Rhinoceros, and George Stubbs's c. 1762 horse portrait Whistlejacket.[185]
+Insects, birds and mammals play roles in literature and film,[186] such as in giant bug movies.[187][188][189] Animals including insects[190] and mammals[191] feature in mythology and religion. In both Japan and Europe, a butterfly was seen as the personification of a person's soul,[190][192][193] while the scarab beetle was sacred in ancient Egypt.[194] Among the mammals, cattle,[195] deer,[191] horses,[196] lions,[197] bats,[198] bears,[199] and wolves[200] are the subjects of myths and worship. The signs of the Western and Chinese zodiacs are based on animals.[201][202]

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+An Arab (/ˈær.əb/;[59] Arabic: عَرَبٌ‎, ISO 233: ‘arab; Arabic pronunciation: [ˈʕarab] (listen)) is a person inhabiting the Arab world. Arabs primarily live among the Arab states in Western Asia, Northern Africa, the Horn of Africa, and Western Indian Ocean islands (including the Comoros). They also live, in significant numbers, in the Americas, Western Europe, Indonesia, Israel, Turkey, and Iran.[60] The Arab diaspora is established around the world.[61]
+The first mention of Arabs appeared in the mid-9th century BCE as a tribal people in eastern and southern Syria, and the north of the Arabian Peninsula.[62] The Arabs appear to have been under the vassalage of the Neo-Assyrian Empire (911–612 BCE), as well as the succeeding Neo-Babylonian (626–539 BCE), Achaemenid (539–332 BCE), Seleucid, and Parthian empires.[63] The Nabataeans, an Arab people, formed their Kingdom near Petra in the 3rd century BC. Arab tribes, most notably the Ghassanids and Lakhmids, begin to appear in the southern Syrian Desert from the mid 3rd century CE onward, during the mid to later stages of the Roman and Sasanian empires.[64]
+Before the expansion of the Rashidun Caliphate (632–661 C.E.), "Arab" referred to any of the largely nomadic and settled Semitic people from the Arabian Peninsula, Syrian Desert, and North and Lower Mesopotamia.[65] Today, "Arab" refers to a large number of people whose native regions form the Arab world due to the spread of Arabs and the Arabic language throughout the region during the early Muslim conquests of the 7th and 8th centuries and the subsequent Arabisation of indigenous populations.[66][67] The Arabs forged the Rashidun (632–661), Umayyad (661–750), Abbasid (750–1517) and the Fatimid (901–1071) caliphates, whose borders reached southern France in the west, China in the east, Anatolia in the north, and the Sudan in the south. This was one of the largest land empires in history.[68] In the early 20th century, the First World War signalled the end of the Ottoman Empire, which had ruled much of the Arab world since conquering the Mamluk Sultanate in 1517.[69] The end culminated in the 1922 defeat and dissolution of the empire and the partition of its territories, forming the modern Arab states.[70] Following the adoption of the Alexandria Protocol in 1944, the Arab League was founded on 22 March 1945.[71] The Charter of the Arab League endorsed the principle of an Arab homeland whilst respecting the individual sovereignty of its member states.[72]
+Today, Arabs primarily inhabit the 22 Arab states within the Arab League: Algeria, Bahrain, the Comoros, Djibouti, Egypt, Iraq, Jordan, Kuwait, Lebanon, Libya, Mauritania, Morocco, Oman, Palestine, Qatar, Saudi Arabia, Somalia, Sudan, Syria, Tunisia, United Arab Emirates, and Yemen. The Arab world stretches around 13 million km2, from the Atlantic Ocean in the west to the Arabian Sea in the east, and from the Mediterranean Sea in the north to the Horn of Africa and the Indian Ocean in the southeast. Beyond the boundaries of the League of Arab States, Arabs can also be found in the global diaspora.[60] The ties that bind Arabs are ethnic, linguistic, cultural, historical, identical, nationalist, geographical and political.[73] The Arabs have their own customs, language, architecture, art, literature, music, dance, media, cuisine, dress, society, sports and mythology.[74]
+Arabs are a diverse group in terms of religious affiliations and practices. In the pre-Islamic era, most Arabs followed polytheistic religions. Some tribes had adopted Christianity or Judaism, and a few individuals, the hanifs, apparently observed another form of monotheism.[75] Today, about 93% of Arabs are adherents of Islam,[76] and there are sizable Christian minorities.[77] Arab Muslims primarily belong to the Sunni, Shiite, Ibadi, and Alawite denominations. Arab Christians generally follow one of the Eastern Christian Churches, such as the Oriental Orthodox or Eastern Catholic churches.[78] There also exist small amounts of Arab Jews still living in Arab countries, and a much larger population of Jews descended from Arab Jewish communities living in Israel and various Western countries, who may or may not consider themselves Arab today. Many Christians in Arab countries may also not consider themselves Arab, especially Copts and Assyrians. Other smaller minority religions are also followed, such as the Bahá'í Faith and Druze.
+Arabs have greatly influenced and contributed to diverse fields, notably the arts and architecture, language, philosophy, mythology, ethics, literature, politics, business, music, dance, cinema, medicine, science and technology in the ancient and modern history.[79]
+The earliest documented use of the word Arab in reference to a people appears in the Kurkh Monoliths, an Akkadian-language record of the Assyrian conquest of Aram (9th century BCE). The Monoliths used the term to refer to Bedouins of the Arabian Peninsula under King Gindibu, who fought as part of a coalition opposed to Assyria.[80] Listed among the booty captured by the army of the Assyrian king Shalmaneser III in the Battle of Qarqar (853 BCE) are 1000 camels of "Gi-in-di-bu'u the ar-ba-a-a" or "[the man] Gindibu belonging to the Arabs" (ar-ba-a-a being an adjectival nisba of the noun ʿarab).[80]
+The related word ʾaʿrāb is used to refer to Bedouins today, in contrast to ʿarab which refers to Arabs in general.[81] Both terms are mentioned around 40 times in pre-Islamic Sabaean inscriptions. The term ʿarab ('Arab') occurs also in the titles of the Himyarite kings from the time of 'Abu Karab Asad until MadiKarib Ya'fur. According to Sabaean grammar, the term ʾaʿrāb is derived from the term ʿarab. The term is also mentioned in Quranic verses, referring to people who were living in Madina and it might be a south Arabian loanword into Quranic language.[82]
+The oldest surviving indication of an Arab national identity is an inscription made in an archaic form of Arabic in 328 CE using the Nabataean alphabet, which refers to Imru' al-Qays ibn 'Amr as 'King of all the Arabs'.[83][84] Herodotus refers to the Arabs in the Sinai, southern Palestine, and the frankincense region (Southern Arabia). Other Ancient-Greek historians like Agatharchides, Diodorus Siculus, and Strabo mention Arabs living in Mesopotamia (along the Euphrates), in Egypt (the Sinai and the Red Sea), southern Jordan (the Nabataeans), the Syrian steppe, and in eastern Arabia (the people of Gerrha). Inscriptions dating to the 6th century BCE in Yemen include the term 'Arab'.[85]
+The most popular Arab account holds that the word Arab came from an eponymous father named Ya'rub, who was supposedly the first to speak Arabic. Abu Muhammad al-Hasan al-Hamdani had another view; he states that Arabs were called gharab ('westerners') by Mesopotamians because Bedouins originally resided to the west of Mesopotamia; the term was then corrupted into arab.
+Yet another view is held by al-Masudi that the word Arab was initially applied to the Ishmaelites of the Arabah valley. In Biblical etymology, Arab (Hebrew: arvi) comes both from the desert origin of the Bedouins it originally described (arava means 'wilderness').
+The root ʿ-r-b has several additional meanings in Semitic languages—including 'west, sunset', 'desert', 'mingle', 'mixed', 'merchant', and 'raven'—and are "comprehensible" with all of these having varying degrees of relevance to the emergence of the name. It is also possible that some forms were metathetical from ʿ-B-R, 'moving around' (Arabic: ʿ-B-R, 'traverse'), and hence, it is alleged, 'nomadic'.[86]
+Pre-Islamic Arabia refers to the Arabian Peninsula prior to the rise of Islam in the 630s. The study of Pre-Islamic Arabia is important to Islamic studies as it provides the context for the development of Islam. Some of the settled communities in the Arabian Peninsula developed into distinctive civilizations. Sources for these civilizations are not extensive, and are limited to archaeological evidence, accounts written outside of Arabia, and Arab oral traditions later recorded by Islamic scholars. Among the most prominent civilizations was Dilmun, which arose around the 4th millennium BCE and lasted to 538 BCE, and Thamud, which arose around the 1st millennium BCE and lasted to about 300 CE. Additionally, from the beginning of the first millennium BCE, Southern Arabia was the home to a number of kingdoms, such as the Sabaean kingdom (Arabic: سَـبَـأ‎, romanized: Saba',[87] possibly Sheba),[88] and the coastal areas of Eastern Arabia were controlled by the Parthian and Sassanians from 300 BCE.
+According to Arab-Islamic-Jewish traditions, Ishmael was father of the Arabs, to be the ancestor of the Ishmaelites.[89]
+The first written attestation of the ethnonym Arab occurs in an Assyrian inscription of 853 BCE, where Shalmaneser III lists a King Gindibu of mâtu arbâi (Arab land) as among the people he defeated at the Battle of Qarqar. Some of the names given in these texts are Aramaic, while others are the first attestations of Ancient North Arabian dialects. In fact several different ethnonyms are found in Assyrian texts that are conventionally translated "Arab": Arabi, Arubu, Aribi and Urbi. Many of the Qedarite queens were also described as queens of the aribi. The Hebrew Bible occasionally refers to Aravi peoples (or variants thereof), translated as "Arab" or "Arabian." The scope of the term at that early stage is unclear, but it seems to have referred to various desert-dwelling Semitic tribes in the Syrian Desert and Arabia.[citation needed] Arab tribes came into conflict with the Assyrians during the reign of the Assyrian king Ashurbanipal, and he records military victories against the powerful Qedar tribe among others.
+Old Arabic diverges from Central Semitic by the beginning of the 1st millennium BCE.[citation needed]
+Medieval Arab genealogists divided Arabs into three groups:
+And Ishmael and his sons, and the sons of Keturah and their sons, went together and dwelt from Paran to the entering in of Babylon in all the land towards the East facing the desert. And these mingled with each other, and their name was called Arabs, and Ishmaelites.
+Assyrian and Babylonian Royal Inscriptions and North Arabian inscriptions from 9th to 6th century BCE, mention the king of Qedar as king of the Arabs and King of the Ishmaelites.[91][92][93][94]
+Of the names of the sons of Ishmael the names "Nabat, Kedar, Abdeel, Dumah, Massa, and Teman" were mentioned in the Assyrian Royal Inscriptions as tribes of the Ishmaelites. Jesur was mentioned in Greek inscriptions in the 1st century BCE.[95]
+Ibn Khaldun's Muqaddima distinguishes between sedentary Arab Muslims who used to be nomadic, and Bedouin nomadic Arabs of the desert. He used the term "formerly nomadic" Arabs and refers to sedentary Muslims by the region or city they lived in, as in Yemenis.[97] The Christians of Italy and the Crusaders preferred the term Saracens for all the Arabs and Muslims of that time.[98] The Christians of Iberia used the term Moor to describe all the Arabs and Muslims of that time.
+Muslims of Medina referred to the nomadic tribes of the deserts as the A'raab, and considered themselves sedentary, but were aware of their close racial bonds. The term "A'raab" mirrors the term Assyrians used to describe the closely related nomads they defeated in Syria. The Qur'an does not use the word ʿarab, only the nisba adjective ʿarabiy. The Qur'an calls itself ʿarabiy, "Arabic", and Mubin, "clear". The two qualities are connected for example in ayat 43.2–3, "By the clear Book: We have made it an Arabic recitation in order that you may understand". The Qur'an became regarded as the prime example of the al-ʿarabiyya, the language of the Arabs. The term ʾiʿrāb has the same root and refers to a particularly clear and correct mode of speech. The plural noun ʾaʿrāb refers to the Bedouin tribes of the desert who resisted Muhammad, for example in at-Tawba 97,
+al-ʾaʿrābu ʾašaddu kufrān wanifāqān "the Bedouin are the worst in disbelief and hypocrisy".
+Based on this, in early Islamic terminology, ʿarabiy referred to the language, and ʾaʿrāb to the Arab Bedouins, carrying a negative connotation due to the Qur'anic verdict just cited. But after the Islamic conquest of the eighth century, the language of the nomadic Arabs became regarded as the most pure by the grammarians following Abi Ishaq, and the term kalam al-ʿArab, "language of the Arabs", denoted the uncontaminated language of the Bedouins.
+Proto-Arabic, or Ancient North Arabian, texts give a clearer picture of the Arabs' emergence. The earliest are written in variants of epigraphic south Arabian musnad script, including the 8th century BCE Hasaean inscriptions of eastern Saudi Arabia, the 6th century BCE Lihyanite texts of southeastern Saudi Arabia and the Thamudic texts found throughout the Arabian Peninsula and Sinai (not in reality connected with Thamud).
+The Nabataeans were nomadic Arabs who moved into territory vacated by the Edomites – Semites who settled the region centuries before them. Their early inscriptions were in Aramaic, but gradually switched to Arabic, and since they had writing, it was they who made the first inscriptions in Arabic. The Nabataean alphabet was adopted by Arabs to the south, and evolved into modern Arabic script around the 4th century. This is attested by Safaitic inscriptions (beginning in the 1st century BCE) and the many Arabic personal names in Nabataean inscriptions. From about the 2nd century BCE, a few inscriptions from Qaryat al-Faw reveal a dialect no longer considered proto-Arabic, but pre-classical Arabic. Five Syriac inscriptions mentioning Arabs have been found at Sumatar Harabesi, one of which dates to the 2nd century CE.
+Arabs arrived in the Palmyra in the late first millennium BCE.[99] The soldiers of the sheikh Zabdibel, who aided the Seleucids in the battle of Raphia (217 BCE), were described as Arabs; Zabdibel and his men were not actually identified as Palmyrenes in the texts, but the name "Zabdibel" is a Palmyrene name leading to the conclusion that the sheikh hailed from Palmyra.[100] Palmyra was conquered by the Rashidun Caliphate after its 634 capture by the Arab general Khalid ibn al-Walid, who took the city on his way to Damascus; an 18-day march by his army through the Syrian Desert from Mesopotamia.[101] By then Palmyra was limited to the Diocletian camp.[102] After the conquest, the city became part of Homs Province.[103]
+Palmyra prospered as part of the Umayyad Caliphate, and its population grew.[104] It was a key stop on the East-West trade route, with a large souq (Arabic: سُـوق‎, market), built by the Umayyads,[104][105] who also commissioned part of the Temple of Bel as a mosque.[105] During this period, Palmyra was a stronghold of the Banu Kalb tribe.[106] After being defeated by Marwan II during a civil war in the caliphate, Umayyad contender Sulayman ibn Hisham fled to the Banu Kalb in Palmyra, but eventually pledged allegiance to Marwan in 744; Palmyra continued to oppose Marwan until the surrender of the Banu Kalb leader al-Abrash al-Kalbi in 745.[107] That year, Marwan ordered the city's walls demolished.[102][108] In 750 a revolt, led by Majza'a ibn al-Kawthar and Umayyad pretender Abu Muhammad al-Sufyani, against the new Abbasid Caliphate swept across Syria;[109] the tribes in Palmyra supported the rebels.[110] After his defeat Abu Muhammad took refuge in the city, which withstood an Abbasid assault long enough to allow him to escape.[110]
+The Ghassanids, Lakhmids and Kindites were the last major migration of pre-Islamic Arabs out of Yemen to the north. The Ghassanids increased the Semitic presence in the then Hellenized Syria, the majority of Semites were Aramaic peoples. They mainly settled in the Hauran region and spread to modern Lebanon, Palestine and Jordan.
+Greeks and Romans referred to all the nomadic population of the desert in the Near East as Arabi. The Romans called Yemen "Arabia Felix".[111] The Romans called the vassal nomadic states within the Roman Empire Arabia Petraea, after the city of Petra, and called unconquered deserts bordering the empire to the south and east Arabia Magna.
+The Lakhmids as a dynasty inherited their power from the Tanukhids, the mid Tigris region around their capital Al-Hira. They ended up allying with the Sassanids against the Ghassanids and the Byzantine Empire. The Lakhmids contested control of the Central Arabian tribes with the Kindites with the Lakhmids eventually destroying Kinda in 540 after the fall of their main ally Himyar. The Persian Sassanids dissolved the Lakhmid dynasty in 602, being under puppet kings, then under their direct control.[112]
+The Kindites migrated from Yemen along with the Ghassanids and Lakhmids, but were turned back in Bahrain by the Abdul Qais Rabi'a tribe. They returned to Yemen and allied themselves with the Himyarites who installed them as a vassal kingdom that ruled Central Arabia from "Qaryah Dhat Kahl" (the present-day called Qaryat al-Faw). They ruled much of the Northern/Central Arabian peninsula, until they were destroyed by the Lakhmid king Al-Mundhir, and his son 'Amr.
+After the death of Muhammad in 632, Rashidun armies launched campaigns of conquest, establishing the Caliphate, or Islamic Empire, one of the largest empires in history. It was larger and lasted longer than the previous Arab empire of Queen Mawia or the Aramean-Arab Palmyrene Empire. The Rashidun state was a completely new state and unlike the Arab kingdoms of its century such as the Himyarite, Lakhmids or Ghassanids.
+In 661, the Rashidun Caliphate fell into the hands of the Umayyad dynasty and Damascus was established as the empire's capital. The Umayyads were proud of their Arab identity and sponsored the poetry and culture of pre-Islamic Arabia. They established garrison towns at Ramla, Raqqa, Basra, Kufa, Mosul and Samarra, all of which developed into major cities.[114]
+Caliph Abd al-Malik established Arabic as the Caliphate's official language in 686.[115] This reform greatly influenced the conquered non-Arab peoples and fueled the Arabization of the region. However, the Arabs' higher status among non-Arab Muslim converts and the latter's obligation to pay heavy taxes caused resentment. Caliph Umar II strove to resolve the conflict when he came to power in 717. He rectified the disparity, demanding that all Muslims be treated as equals, but his intended reforms did not take effect, as he died after only three years of rule. By now, discontent with the Umayyads swept the region and an uprising occurred in which the Abbasids came to power and moved the capital to Baghdad.
+Umayyads expanded their Empire westwards capturing North Africa from the Byzantines. Before the Arab conquest, North Africa was conquered or settled by various people including Punics, Vandals and Romans. After the Abbasid Revolution, the Umayyads lost most of their territories with the exception of Iberia. Their last holding became known as the Emirate of Córdoba. It wasn't until the rule of the grandson of the founder of this new emirate that the state entered a new phase as the Caliphate of Córdoba. This new state was characterized by an expansion of trade, culture and knowledge, and saw the construction of masterpieces of al-Andalus architecture and the library of Al-Ḥakam II which housed over 400,000 volumes. With the collapse of the Umayyad state in 1031 CE, Islamic Spain was divided into small kingdoms.
+The Abbasids were the descendants of Abbas ibn Abd al-Muttalib, one of the youngest uncles of Muhammad and of the same Banu Hashim clan. The Abbasids led a revolt against the Umayyads and defeated them in the Battle of the Zab effectively ending their rule in all parts of the Empire with the exception of al-Andalus. In 762, the second Abbasid Caliph al-Mansur founded the city of Baghdad and declared it the capital of the Caliphate. Unlike the Umayyads, the Abbasids had the support of non-Arab subjects.[114]
+The Islamic Golden Age was inaugurated by the middle of the 8th century by the ascension of the Abbasid Caliphate and the transfer of the capital from Damascus to the newly founded city of Baghdad. The Abbassids were influenced by the Qur'anic injunctions and hadith such as "The ink of the scholar is more holy than the blood of martyrs" stressing the value of knowledge. During this period the Muslim world became an intellectual centre for science, philosophy, medicine and education as the Abbasids championed the cause of knowledge and established the "House of Wisdom" (Arabic: بيت الحكمة‎) in Baghdad. Rival dynasties such as the Fatimids of Egypt and the Umayyads of al-Andalus were also major intellectual centres with cities such as Cairo and Córdoba rivaling Baghdad.[116]
+The Abbasids ruled for 200 years before they lost their central control when Wilayas began to fracture in the 10th century; afterwards, in the 1190s, there was a revival of their power, which was ended by the Mongols, who conquered Baghdad in 1258 and killed the Caliph Al-Musta'sim. Members of the Abbasid royal family escaped the massacre and resorted to Cairo, which had broken from the Abbasid rule two years earlier; the Mamluk generals taking the political side of the kingdom while Abbasid Caliphs were engaged in civil activities and continued patronizing science, arts and literature.
+The Fatimid caliphate was founded by al-Mahdi Billah, a descendant of Fatimah, the daughter of Muhammad, in the early 10th century. Egypt was the political, cultural, and religious centre of the Fatimid empire. The Fatimid state took shape among the Kutama Berbers, in the West of the North African littoral, in Algeria, in 909 conquering Raqqada, the Aghlabid capital. In 921 the Fatimids established the Tunisian city of Mahdia as their new capital. In 948 they shifted their capital to Al-Mansuriya, near Kairouan in Tunisia, and in 969 they conquered Egypt and established Cairo as the capital of their caliphate.
+Intellectual life in Egypt during the Fatimid period achieved great progress and activity, due to many scholars who lived in or came to Egypt, as well as the number of books available. Fatimid Caliphs gave prominent positions to scholars in their courts, encouraged students, and established libraries in their palaces, so that scholars might expand their knowledge and reap benefits from the work of their predecessors.[117] The Fatimids were also known for their exquisite arts. Many traces of Fatimid architecture exist in Cairo today; the most defining examples include Al-Hakim Mosque and the Al-Azhar University.
+It was not until the 11th century that the Maghreb saw a large influx of ethnic Arabs. Starting with the 11th century, the Arab bedouin Banu Hilal tribes migrated to the West. Having been sent by the Fatimids to punish the Berber Zirids for abandoning Shias, they travelled westwards. The Banu Hilal quickly defeated the Zirids and deeply weakened the neighboring Hammadids. According to some modern historians. their influx was a major factor in the arabization of the Maghreb.[118][119] Although Berbers ruled the region until the 16th century (under such powerful dynasties as the Almoravids, the Almohads, Hafsids, etc.), the arrival of these tribes eventually helped Arabize much of it ethnically, in addition to the linguistic and political impact on local non-Arabs.[citation needed]
+From 1517 to 1918, much of the Arab world was under the suzerainty of the Ottoman Empire. The Ottomans defeated the Mamluk Sultanate in Cairo, and ended the Abbasid Caliphate. Arabs did not feel the change of administration because the Ottomans modeled their rule after the previous Arab administration systems.[citation needed]
+In 1911, Arab intellectuals and politicians from throughout the Levant formed al-Fatat ("the Young Arab Society"), a small Arab nationalist club, in Paris. Its stated aim was "raising the level of the Arab nation to the level of modern nations." In the first few years of its existence, al-Fatat called for greater autonomy within a unified Ottoman state rather than Arab independence from the empire. Al-Fatat hosted the Arab Congress of 1913 in Paris, the purpose of which was to discuss desired reforms with other dissenting individuals from the Arab world. However, as the Ottoman authorities cracked down on the organization's activities and members, al-Fatat went underground and demanded the complete independence and unity of the Arab provinces.[120]
+After World War I, when the Ottoman Empire was overthrown by the British Empire, former Ottoman colonies were divided up between the British and French as League of Nations mandates.
+Arabs in modern times live in the Arab world, which comprises 22 countries in Western Asia, North Africa, and parts of the Horn of Africa. They are all modern states and became significant as distinct political entities after the fall and defeat and dissolution of the Ottoman Empire (1908–1922).
+Arab identity is defined independently of religious identity, and pre-dates the spread of Islam, with historically attested Arab Christian kingdoms and Arab Jewish tribes. Today, however, most Arabs are Muslim, with a minority adhering to other faiths, largely Christianity, but also Druze and Baha'i.[121][122]
+Paternal descent has traditionally been considered the main source of affiliation in the Arab world when it comes to membership into an ethnic group or clan.[123]
+Today, the main unifying characteristic among Arabs is Arabic, a Central Semitic language from the Afroasiatic language family. Modern Standard Arabic serves as the standardized and literary variety of Arabic used in writing. The Arabs are first mentioned in the mid-ninth century BCE as a tribal people dwelling in the central Arabian Peninsula subjugated by Upper Mesopotamia-based state of Assyria. The Arabs appear to have remained largely under the vassalage of the Neo-Assyrian Empire (911–605 BCE), and then the succeeding Neo-Babylonian Empire (605–539 BCE), Persian Achaemenid Empire (539–332 BCE), Greek Macedonian/Seleucid Empire and Parthian Empire.
+Arab tribes, most notably the Ghassanids and Lakhmids begin to appear in the south Syrian deserts and southern Jordan from the mid 3rd century CE onwards, during the mid to later stages of the Roman Empire and Sasanian Empire. Also, before them the Nabataeans of Jordan and arguably the Emessans,[124] Edessans,[125] and Hatrans[126] all appear to have been an Aramaic speaking ethnic Arabs who came to rule much of the pre-Islamic fertile crescent often as vassals of the two rival empires, the Sasanian (Persian) and the Byzantine (Eastern Roman).[127] Thus, although a more limited diffusion of Arab culture and language was felt in some areas by these migrant minority Arabs in pre-Islamic times through Arabic-speaking Christian kingdoms and Jewish tribes, it was only after the rise of Islam in the mid-7th century that Arab culture, people and language began their wholesale spread from the central Arabian Peninsula (including the south Syrian desert) through conquest and trade.
+Arabs in the narrow sense are the indigenous Arabians who trace their roots back to the tribes of Arabia and their immediate descendant groups in the Levant and North Africa. Within the people of the Arabian Peninsula, distinction is made between:
+Arabians are most prevalent in the Arabian Peninsula, but are also found in large numbers in Mesopotamia (Arab tribes in Iraq), the Levant and Sinai (Negev Bedouin, Tarabin bedouin), as well as the Maghreb (Eastern Libya, South Tunisia and South Algeria) and the Sudan region.
+This traditional division of the Arabs of Arabia may have arisen at the time of the First Fitna. Of the Arabian tribes that interacted with Muhammad, the most prominent was the Quraysh. The Quraysh subclan, the Banu Hashim, was the clan of Muhammad. During the early Muslim conquests and the Islamic Golden Age, the political rulers of Islam were exclusively members of the Quraysh.
+The Arab presence in Iran did not begin with the Arab conquest of Persia in 633 CE. For centuries, Iranian rulers had maintained contacts with Arabs outside their borders, dealt with Arab subjects and client states (such as those of Iraq and Yemen), and settled Arab tribesmen in various parts of the Iranian plateau. It follows that the "Arab" conquests and settlements were by no means the exclusive work of Arabs from the Hejaz and the tribesmen of inner Arabia. The Arab infiltration into Iran began before the Muslim conquests and continued as a result of the joint exertions of the civilized Arabs (ahl al-madar) as well as the desert Arabs (ahl al-wabar).[130] The largest group of Iranian Arabs are the Ahwazi Arabs, including Banu Ka'b, Bani Turuf and the Musha'sha'iyyah sect. Smaller groups are the Khamseh nomads in Fars Province and the Arabs in Khorasan.
+The Arabs of the Levant are traditionally divided into Qays and Yaman tribes. This tribal division is likewise taken to date to the Umayyad period. The Yemen trace their origin to South Arabia or Yemen; they include Banu Kalb, Kindah, Ghassanids, and Lakhmids.[131] Since the 1834 Peasants' revolt in Palestine, the Arabic-speaking population of Palestine has shed its formerly tribal structure and emerged as the Palestinians[citation needed].
+Native Jordanians are either descended from Bedouins (of which, 6% live a nomadic lifestyle),[132] or from the many deeply rooted non bedouin communities across the country, most notably Al-Salt city west of Amman which was at the time of Emirate the largest urban settlement east of the Jordan River. Along with indigenous communities in Al Husn, Aqaba, Irbid, Al Karak, Madaba, Jerash, Ajloun, Fuheis and Pella.[133] In Jordan, there is no official census data for how many inhabitants have Palestinian roots but they are estimated to constitute half of the population,[134][135] which in 2008 amounted to about 3 million.[135] Palestinian Central Bureau of Statistics put their number at 3.24 million in 2009.[136]
+The Bedouins of western Egypt and eastern Libya are traditionally divided into Saʿada and Murabtin, the Saʿada having higher social status. This may derive from a historical feudal system in which the Murabtin were vassals to the Saʿada.
+In Sudan, there are numerous Arabic-speaking tribes, including the Shaigya, Ja'alin and Shukria, who are ancestrally related to the Nubians. These groups are collectively known as Sudanese Arabs. In addition, there are other Afroasiatic-speaking populations, such as Copts and Beja.
+The medieval Arab slave trade in the Sudan drove a wedge between the Arabic-speaking groups and the indigenous Nilotic populations. Slavery substantially persists today along these lines.[137] It has contributed to ethnic conflict in the region, such as the Sudanese conflict in South Kordofan and Blue Nile, Northern Mali conflict, or the Boko Haram insurgency.
+The Arabs of the Maghreb are descendants of Arabian tribes of Banu Hilal, the Banu Sulaym and the Maqil native of Middle East[138] and of other tribes native to Saudi Arabia, Yemen and Iraq. Arabs and Arabic-speakers inhabit plains and cities. The Banu Hilal spent almost a century in Egypt before moving to Libya, Tunisia and Algeria, and another century later some moved to Morocco, it is logical to think that they are mixed with inhabitants of Egypt and with Libya.[139]
+The total number of Arabic speakers living in the Arab nations is estimated at 366 million by the CIA Factbook (as of 2014). The estimated number of Arabs in countries outside the Arab League is estimated at 17.5 million, yielding a total of close to 384 million.
+According to the Charter of the Arab League (also known as the Pact of the League of Arab States), the League of Arab States is composed of independent Arab states that are signatories to the Charter.[140]
+Although all Arab states have Arabic as an official language, there are many non-Arabic-speaking populations native to the Arab world. Among these are Berbers, Toubou, Nubians, Jews, Kurds, Armenians.[58] Additionally, many Arab countries in the Persian Gulf have sizable non-Arab immigrant populations (10–30%). Iraq, Bahrain, Kuwait, Qatar, United Arab Emirates and Oman have a Persian speaking minority. The same countries also have Hindi-Urdu speakers and Filipinos as sizable minority. Balochi speakers are a good size minority in Oman. Additionally, countries like Bahrain, UAE, Oman and Kuwait have significant non-Arab and non-Muslim minorities (10–20%) like Hindus and Christians from India, Pakistan, Bangladesh, Nepal and the Philippines.
+The table below shows the distribution of populations in the Arab world, as well as the official language(s) within the various Arab states.[141]
+Arab diaspora refers to descendants of the Arab immigrants who, voluntarily or as refugees, emigrated from their native lands in non-Arab countries, primarily in East Africa, South America, Europe, North America, Australia and parts of South Asia, Southeast Asia, the Caribbean, and West Africa. According to the International Organization for Migration, there are 13 million first-generation Arab migrants in the world, of which 5.8 million reside in Arab countries. Arab expatriates contribute to the circulation of financial and human capital in the region and thus significantly promote regional development. In 2009, Arab countries received a total of 35.1 billion USD in remittance in-flows and remittances sent to Jordan, Egypt and Lebanon from other Arab countries are 40 to 190 per cent higher than trade revenues between these and other Arab countries.[164] The 250,000 strong Lebanese community in West Africa is the largest non-African group in the region.[165][166] Arab traders have long operated in Southeast Asia and along the East Africa's Swahili coast. Zanzibar was once ruled by Omani Arabs.[167] Most of the prominent Indonesians, Malaysians, and Singaporeans of Arab descent are Hadhrami people with origins in southern Yemen in the Hadramawt coastal region.[168]
+There are millions of Arabs living in Europe, mostly concentrated in France (about 6,000,000 in 2005[12]). Most Arabs in France are from the Maghreb but some also come from the Mashreq areas of the Arab world. Arabs in France form the second largest ethnic group after ethnically French people.[169] The modern Arab population of Spain numbers 1,800,000,[170][171][172][173] and there have been Arabs in Spain since the early 8th century when the Umayyad conquest of Hispania created the state of Al-Andalus.[174][175][176] In Germany the Arab population numbers over 1,000,000,[177] in Italy about 680,000,[41] in the United Kingdom between 366,769[178] and 500,000,[179] and in Greece between 250,000 and 750,000[180][failed verification]). In addition, Greece is home to people from Arab countries who have the status of refugees (e.g. refugees of the Syrian civil war).[181] In the Netherlands 180,000,[182] and in Denmark 121,000. Other European countries are also home to Arab populations, including Norway, Austria, Bulgaria, Switzerland, North Macedonia, Romania and Serbia.[183] As of late 2015, Turkey had a total population of 78.7 million, with Syrian refugees accounting for 3.1% of that figure based on conservative estimates[by whom?]. Demographics indicated that the country previously had 1,500,000[184] to 2,000,000 Arab residents,[21] so Turkey's Arab population is now 4.5 to 5.1% of the total population, or approximately 4–5 million people.[21][20]
+Arab immigration to the United States began in sizable numbers during the 1880s. Today, it is estimated that nearly 3.7 million Americans trace their roots to an Arab country.[24][185][186] Arab Americans are found in every state, but more than two thirds of them live in just ten states: California, Michigan, New York, Florida, Texas, New Jersey, Illinois, Ohio, Pennsylvania, and Virginia. Metropolitan Los Angeles, Detroit, and New York City are home to one-third of the population.[24][187] Contrary to popular assumptions or stereotypes, the majority of Arab Americans are native-born, and nearly 82% of Arabs in the U.S. are citizens.[188][189][189][190][191] Arabs immigrants began to arrive in Canada in small numbers in 1882. Their immigration was relatively limited until 1945, after which time it increased progressively, particularly in the 1960s and thereafter.[192] According to the website "Who are Arab Canadians," Montreal, the Canadian city with the largest Arab population, has approximately 267,000 Arab inhabitants.[193]
+Latin America has the largest Arab population outside of the Arab World.[194] Latin America is home to anywhere from 17–25 to 30 million people of Arab descent,[195] which is more than any other diaspora region in the world.[196][197] The Brazilian and Lebanese governments claim there are 7 million Brazilians of Lebanese descent.[198][199] Also, the Brazilian government claims there are 4 million Brazilians of Syrian descent.[198] According to research conducted by IBGE in 2008, covering only the states of Amazonas, Paraíba, São Paulo, Rio Grande do Sul, Mato Grosso and Distrito Federal, 0.9% of white Brazilian respondents said they had family origins in the Middle East.[200][201][202][203][204] Other large Arab communities includes Argentina (about 4,500,000[23][205][206]) The interethnic marriage in the Arab community, regardless of religious affiliation, is very high; most community members have only one parent who has Arab ethnicity.[207] Venezuela (over 1,600,000[26][208]), Colombia (over 1,600,000[27] to 3,200,000[209][210][211]), Mexico (over 1,100,000[30]), Chile (over 800,000[212][37][213][214]), and Central America, particularly El Salvador, and Honduras (between 150,000 and 200,000).[46][38][39] is the fourth largest in the world after those in Israel, Lebanon, and Jordan. Arab Haitians (a large number of whom live in the capital) are more often than not, concentrated in financial areas where the majority of them establish businesses.[215][215][215]
+In 1728, a Russian officer described a group of Arab nomads who populated the Caspian shores of Mughan (in present-day Azerbaijan) and spoke a mixed Turkic-Arabic language.[216] It is believed that these groups migrated to the South Caucasus in the 16th century.[217] The 1888 edition of Encyclopædia Britannica also mentioned a certain number of Arabs populating the Baku Governorate of the Russian Empire.[218] They retained an Arabic dialect at least into the mid-19th century,[219] there are nearly 30 settlements still holding the name Arab (for example, Arabgadim, Arabojaghy, Arab-Yengija, etc.). From the time of the Arab conquest of the South Caucasus, continuous small-scale Arab migration from various parts of the Arab world occurred in Dagestan. The majority of these lived in the village of Darvag, to the north-west of Derbent. The latest of these accounts dates to the 1930s.[217] Most Arab communities in southern Dagestan underwent linguistic Turkicisation, thus nowadays Darvag is a majority-Azeri village.[220][221] According to the History of Ibn Khaldun, the Arabs that were once in Central Asia have been either killed or have fled the Tatar invasion of the region, leaving only the locals.[222] However, today many people in Central Asia identify as Arabs. Most Arabs of Central Asia are fully integrated into local populations, and sometimes call themselves the same as locals (for example, Tajiks, Uzbeks) but they use special titles to show their Arab origin such as Sayyid, Khoja or Siddiqui.[223]
+There are only two communities in India which self-identify as Arabs, the Chaush of the Deccan region and the Chavuse of Gujarat.[224][225] These groups are largely descended from Hadhrami migrants who settled in these two regions in the 18th century. However, neither community still speaks Arabic, although the Chaush have seen re-immigration to the Arab States of the Persian Gulf and thus a re-adoption of Arabic.[226] In South Asia, where Arab ancestry is considered prestigious, many communities have origin myths that claim Arab ancestry. These include the Mappilla of Kerala and the Labbai of Tamil Nadu.[227] Among North Indian and Pakistani Arabs, there are groups who claim the status of Sayyid and have origin myths that allege descent from Muhammad.[228] The South Asian Iraqi biradri may be considered Arabs because records of their ancestors who migrated from Iraq exist in historical documents. There are about 5,000,000 Native Indonesians with Arab ancestry.[229] Arab Indonesians are mainly of Hadrami descent.[230][230] The Sri Lankan Moors are the third largest ethnic group in Sri Lanka, comprising 9.23% of the country's total population.[231] Some sources trace the ancestry of the Sri Lankan Moors to Arab traders who settled in Sri Lanka at some time between the 8th and 15th centuries.[232][233][234]
+Afro-Arabs are individuals and groups from Africa who are of partial Arab descent. Most Afro-Arabs inhabit the Swahili Coast in the African Great Lakes region, although some can also be found in parts of the Arab world.[235][236] Large numbers of Arabs migrated to West Africa, particularly Côte d'Ivoire (home to over 100,000 Lebanese),[237] Senegal (roughly 30,000 Lebanese),[238] Sierra Leone (roughly 10,000 Lebanese today; about 30,000 prior to the outbreak of civil war in 1991), Liberia, and Nigeria.[239] Since the end of the civil war in 2002, Lebanese traders have become re-established in Sierra Leone.[240][241][242] The Arabs of Chad occupy northern Cameroon and Nigeria (where they are sometimes known as Shuwa), and extend as a belt across Chad and into Sudan, where they are called the Baggara grouping of Arab ethnic groups inhabiting the portion of Africa's Sahel. There are 171,000 in Cameroon, 150,000 in Niger[243]), and 107,000 in the Central African Republic.[citation needed]
+Arabs are mostly Muslims with a Sunni majority and a Shia minority, one exception being the Ibadis, who predominate in Oman.[244] Arab Christians generally follow Eastern Churches such as the Greek Orthodox and Greek Catholic churches, though a minority of Protestant Church followers also exists.[245] There are also Arab communities consisting of Druze and Baha'is.[246][247]
+Before the coming of Islam, most Arabs followed a pagan religion with a number of deities, including Hubal,[248] Wadd, Allāt,[249] Manat, and Uzza. A few individuals, the hanifs, had apparently rejected polytheism in favor of monotheism unaffiliated with any particular religion. Some tribes had converted to Christianity or Judaism. The most prominent Arab Christian kingdoms were the Ghassanid and Lakhmid kingdoms.[250] When the Himyarite king converted to Judaism in the late 4th century,[251] the elites of the other prominent Arab kingdom, the Kindites, being Himyirite vassals, apparently also converted (at least partly). With the expansion of Islam, polytheistic Arabs were rapidly Islamized, and polytheistic traditions gradually disappeared.[252][253]
+Today, Sunni Islam dominates in most areas, overwhelmingly so in North Africa and the Horn of Africa. Shia Islam is dominant among the Arab population in Bahrain and southern Iraq while northern Iraq is mostly Sunni. Substantial Shia populations exist in Lebanon, Yemen, Kuwait, Saudi Arabia,[254] northern Syria and Al-Batinah Region in Oman. There are small numbers of Ibadi and non-denominational Muslims too.[244] The Druze community is concentrated in Lebanon, Syria, Israel and Jordan. Many Druze claim independence from other major religions in the area and consider their religion more of a philosophy. Their books of worship are called Kitab Al Hikma (Epistles of Wisdom). They believe in reincarnation and pray to five messengers from God. In Israel, the Druze have a status aparte from the general Arab population, treated as a separate ethno-religious community.
+Christianity had a prominent presence In pre-Islamic Arabia among several Arab communities, including the Bahrani people of Eastern Arabia, the Christian community of Najran, in parts of Yemen, and among certain northern Arabian tribes such as the Ghassanids, Lakhmids, Taghlib, Banu Amela, Banu Judham, Tanukhids and Tayy. In the early Christian centuries, Arabia was sometimes known as Arabia heretica, due to its being "well known as a breeding-ground for heterodox interpretations of Christianity."[255] Christians make up 5.5% of the population of Western Asia and North Africa.[256] A sizeable share of those are Arab Christians proper, and affiliated Arabic-speaking populations of Copts and Maronites. In Lebanon, Christians number about 40.5% of the population.[150] In Syria, Christians make up 10% of the population.[160] In West Bank and in Gaza Strip, Christians make up 8% and 0.7% of the populations, respectively.[257][258] In Egypt, Coptic Christians number about 10% of the population. In Iraq, Christians constitute 0.1% of the population.[259] In Israel, Arab Christians constitute 2.1% (roughly 9% of the Arab population).[260] Arab Christians make up 8% of the population of Jordan.[261] Most North and South American Arabs are Christian,[262] so are about half of the Arabs in Australia who come particularly from Lebanon, Syria and Palestine. One well known member of this religious and ethnic community is Saint Abo, martyr and the patron saint of Tbilisi, Georgia.[263] Arab Christians also live in holy Christian cities such as Nazareth, Bethlehem and the Christian Quarter of the Old City of Jerusalem and many other villages with holy Christian sites.
+Styles
+Features
+Types
+Styles
+Types
+Features
+Headwear
+Clothing
+Theory
+Genres
+Art music
+Folk
+Language
+Prose
+Islamic
+Poetry
+Genres
+Forms
+Arabic prosody
+National literatures of Arab States
+Concepts
+Texts
+Fictional Arab people
+North Arabian deities
+South Arabian  deities
+Arabic culture is the culture of Arab people, from the Atlantic Ocean in the west to the Arabian Sea in the east, and from the Mediterranean Sea. Language, literature, gastronomy, art, architecture, music, spirituality, philosophy, mysticism (etc.) are all part of the cultural heritage of the Arabs.[264]
+Arabs share basic beliefs and values that cross national and social class boundaries. Social attitudes have remained constant because Arab society is more conservative and demands conformity from its members.[265]
+Another important and unifying characteristic of Arabs is a common language. Arabic is a Semitic language of the Afro-Asiatic Family.[266] Evidence of its first use appears in accounts of wars in 853 BCE. It also became widely used in trade and commerce. Arabic also is a liturgical language of 1.7 billion Muslims.[267][268]
+Arabic is one of six official languages of the United Nations.[269] It is revered as the language that God chose to reveal the Quran.[270][271]
+Arabic has developed into at least two distinct forms. Classical Arabic is the form of the Arabic language used in literary texts from Umayyad and Abbasid times (7th to 9th centuries). It is based on the medieval dialects of Arab tribes. Modern Standard Arabic (MSA) is the direct descendant used today throughout the Arab world in writing and in formal speaking, for example, prepared speeches, some radio broadcasts, and non-entertainment content,[272] while the lexis and stylistics of Modern Standard Arabic are different from Classical Arabic. Colloquial Arabic, an informal spoken language, varies by dialect from region to region; various forms of the language are in use today and provide an important force for Arab cohesion.[273]
+Arabic mythology comprises the ancient beliefs of the Arabs.[274] Prior to Islam the Kaaba of Mecca was covered in symbols representing the myriad demons, djinn, demigods, or simply tribal gods and other assorted deities which represented the polytheistic culture of pre-Islamic.[275][276] It has been inferred from this plurality an exceptionally broad context in which mythology could flourish. The most popular beasts and demons of Arabian mythology are Bahamut, Dandan, Falak, Ghoul, Hinn, Jinn, Karkadann, Marid, Nasnas, Qareen, Roc, Shadhavar, Werehyena and other assorted creatures which represented the profoundly polytheistic environment of pre-Islamic.[277]
+The most obvious symbol of Arabian mythology is the Jinn or genie.[278] Jinns are supernatural beings of varying degrees of power. They possess free will (that is, they can choose to be good or evil) and come in two flavors. There are the Marids, usually described as the most powerful type of Jinn. These are the type of genie with the ability to grant wishes to humans. However, granting these wishes is not free. The Quran says that the jinn were created from "mārijin min nar" (smokeless fire or a mixture of fire; scholars explained, this is the part of the flame, which mixed with the blackness of fire).[279][280] They are not purely spiritual, but are also physical in nature, being able to interact in a tactile manner with people and objects and likewise be acted upon. The jinn, humans, and angels make up the known sapient creations of God.[281]
+A ghoul is a monster or evil spirit in Arabic mythology, associated with graveyards and consuming human flesh,[282][283] demonic being believed to inhabit burial grounds and other deserted places. In ancient Arabic folklore, ghūls belonged to a diabolic class of jinn (spirits) and were said to be the offspring of Iblīs, the prince of darkness in Islam. They were capable of constantly changing form, but their presence was always recognizable by their unalterable sign—ass's hooves.[284] which describes the ghūl of Arabic folklore. The ghul is a devilish type of jinn believed to be sired by Iblis.[285]
+Al-Jahiz (born 776, in Basra – December 868/January 869) was an Arab prose writer and author of works of literature, Mu'tazili theology, and politico-religious polemics. A leading scholar in the Abassid Caliphate, his canon includes two hundred books on various subjects, including Arabic grammar, zoology, poetry, lexicography, and rhetoric. Of his writings, only thirty books survive. Al-Jāḥiẓ was also one of the first Arabian writers to suggest a complete overhaul of the language's grammatical system, though this would not be undertaken until his fellow linguist Ibn Maḍāʾ took up the matter two hundred years later.[287]
+There is a small remnant of pre-Islamic poetry, but Arabic literature predominantly emerges in the Middle Ages, during the Golden Age of Islam.[288] Literary Arabic is derived from Classical Arabic, based on the language of the Quran as it was analyzed by Arabic grammarians beginning in the 8th century.[289]
+A large portion of Arabic literature before the 20th century is in the form of poetry, and even prose from this period is either filled with snippets of poetry or is in the form of saj or rhymed prose.[291]
+The ghazal or love poem had a long history being at times tender and chaste and at other times rather explicit.[292] In the Sufi tradition the love poem would take on a wider, mystical and religious importance. Arabic epic literature was much less common than poetry, and presumably originates in oral tradition, written down from the 14th century or so. Maqama or rhymed prose is intermediate between poetry and prose, and also between fiction and non-fiction.[293] Maqama was an incredibly popular form of Arabic literature, being one of the few forms which continued to be written during the decline of Arabic in the 17th and 18th centuries.[294]
+Arabic literature and culture declined significantly after the 13th century, to the benefit of Turkish and Persian. A modern revival took place beginning in the 19th century, alongside resistance against Ottoman rule. The literary revival is known as al-Nahda in Arabic, and was centered in Egypt and Lebanon. Two distinct trends can be found in the nahda period of revival.[295] The first was a neo-classical movement which sought to rediscover the literary traditions of the past, and was influenced by traditional literary genres—such as the maqama—and works like One Thousand and One Nights. In contrast, a modernist movement began by translating Western modernist works—primarily novels—into Arabic.[296] A tradition of modern Arabic poetry was established by writers such as Francis Marrash, Ahmad Shawqi and Hafiz Ibrahim. Iraqi poet Badr Shakir al-Sayyab is considered to be the originator of free verse in Arabic poetry.[297][298][299]
+Arabic cuisine is the cuisine of the Arab people.[300] The cuisines are often centuries old and reflect the culture of great trading in spices, herbs, and foods. The three main regions, also known as the Maghreb, the Mashriq, and the Khaleej have many similarities, but also many unique traditions. These kitchens have been influenced by the climate, cultivating possibilities, as well as trading possibilities. The kitchens of the Maghreb and Levant are relatively young kitchens which were developed over the past centuries. The kitchen from the Khaleej region is a very old kitchen. The kitchens can be divided into the urban and rural kitchens.
+Arab cuisine mostly follows one of three culinary traditions – from the Maghreb, the Levant or the Persian Gulf states. In the Maghreb countries (Morocco, Algeria, Tunisia and Libya) traditional main meals are tajines or dishes using couscous. In the Levant (Palestine, Jordan, Lebanon and Syria) main meals usually start with mezze – small dishes of dips and other items which are eaten with bread. This is typically followed by skewers of grilled lamb or chicken. Gulf cuisine, tends to be more highly spiced with more use of rice. Sometimes a lamb is roasted and served whole.[301]
+One will find the following items in most dishes; cinnamon, fish (in coastal areas), garlic, lamb (or veal), mild to hot sauces, mint, onion, rice, saffron, sesame, yogurt, spices due to heavy trading between the two regions. Tea, thyme (or oregano), turmeric, a variety of fruits (primarily citrus) and vegetables such as cucumbers, eggplants, lettuce, tomato, green pepper, green beans, zucchini and parsley.[301][302]
+Arabic art takes on many forms, though it is jewelry, textiles and architecture that are the most well-known. It is generally split up by different eras, among them being early Arabic, early medieval, late medieval, late Arabic, and finally, current Arabic. One thing to remember is that many times a particular style from one era may continue into the next with few changes, while some have a drastic transformation. This may seem like a strange grouping of art mediums, but they are all closely related.[303][304]
+Arabic writing is done from right to left, and was generally written in dark inks, with certain things embellished with special colored inks (red, green, gold). In early Arabic and early Medieval, writing was typically done on parchment made of animal skin. The ink showed up very well on it, and occasionally the parchment was dyed a separate color and brighter ink was used (this was only for special projects). The name given to the form of writing in early times was called Kufic script.[305]
+Arabic miniatures are small paintings on paper, whether book illustrations or separate works of art. Arabic miniature art dates to the late 7th century. Arabs depended on such art not only to satisfy their artistic taste, but also for scientific explanations.
+Arabesque is a form of artistic decoration consisting of "surface decorations based on rhythmic linear patterns of scrolling and interlacing foliage, tendrils" or plain lines,[306] often combined with other elements. Another definition is "Foliate ornament, typically using leaves, derived from stylised half-palmettes, which were combined with spiralling stems".[307] It usually consists of a single design which can be 'tiled' or seamlessly repeated as many times as desired.[308][309]
+Arabic Architecture has a deep diverse history, it dates to the dawn of the history in pre-Islamic Arabia and includes various styles from the Nabataean architecture to the old yet still used architecture in various regions of the Arab world. Each of it phases largely
+an extension of the earlier phase, it left also heavy impact on the architecture of other nations. Arab Architecture also encompasses a wide range of both secular and religious styles from the foundation of Islam to the present day. Some parts of its religious architectures raised by Muslim Arabs were influenced by cultures of Roman, Byzantine and cultures of other lands which the Arab conquered in the 7th and 8th centuries.[310][311]
+In Sicily, Arab-Norman architecture combined Occidental features, such as the Classical pillars and friezes, with typical Arabic decorations and calligraphy. The principal Islamic architectural types are: the Mosque, the Tomb, the Palace and the Fort. From these four types, the vocabulary of Islamic architecture is derived and used for other buildings such as public baths, fountains and domestic architecture.[312][313]
+Arabic music, while independent and flourishing in the 2010s, has a long history of interaction with many other regional musical styles and genres. It is an amalgam of the music of the Arab people in the Arabian Peninsula and the music of all the peoples that make up the Arab world today.[314] Pre-Islamic Arab music was similar to that of Ancient Middle Eastern music. Most historians agree that there existed distinct forms of music in the Arabian peninsula in the pre-Islamic period between the 5th and 7th century CE. Arab poets of that "Jahili poets", meaning "the poets of the period of ignorance"—used to recite poems with a high notes.[315] It was believed that Jinns revealed poems to poets and music to musicians.[315][315] By the 11th century, Islamic Iberia had become a center for the manufacture of instruments. These goods spread gradually throughout France, influencing French troubadours, and eventually reaching the rest of Europe. The English words lute, rebec, and naker are derived from Arabic oud, rabab, and naqareh.[316][317]
+A number of musical instruments used in classical music are believed to have been derived from Arabic musical instruments: the lute was derived from the Oud, the rebec (ancestor of violin) from the rebab, the guitar from qitara, which in turn was derived from the Persian Tar, naker from naqareh, adufe from al-duff, alboka from al-buq, anafil from al-nafir, exabeba from al-shabbaba (flute), atabal (bass drum) from al-tabl, atambal from al-tinbal,[318] the balaban, the castanet from kasatan, sonajas de azófar from sunuj al-sufr, the conical bore wind instruments,[319] the xelami from the sulami or fistula (flute or musical pipe),[320]
+the shawm and dulzaina from the reed instruments zamr and al-zurna,[321] the gaita from the ghaita, rackett from iraqya or iraqiyya,[322] geige (violin) from ghichak,[323]
+and the theorbo from the tarab.[324]
+During the 1950s and the 1960s, Arabic music began to take on a more Western tone – artists Umm Kulthum, Abdel Halim Hafez, and Shadia along with composers Mohamed Abd al-Wahab and Baligh Hamdi pioneered the use of western instruments in Egyptian music. By the 1970s several other singers had followed suit and a strand of Arabic pop was born. Arabic pop usually consists of Western styled songs with Arabic instruments and lyrics. Melodies are often a mix between Eastern and Western. Beginning in the mid-1980s, Lydia Canaan, musical pioneer widely regarded as the first rock star of the Middle East[325][326][327][328][329][330][331][332]
+Arab polytheism was the dominant religion in pre-Islamic Arabia. Gods and goddesses, including Hubal and the goddesses al-Lāt, Al-'Uzzá and Manāt, were worshipped at local shrines, such as the Kaaba in Mecca, whilst Arabs in the south, in what is today's Yemen, worshipped various gods, some of which represented the Sun or Moon. Different theories have been proposed regarding the role of Allah in Meccan religion.[75][333][334][335] Many of the physical descriptions of the pre-Islamic gods are traced to idols, especially near the Kaaba, which is said to have contained up to 360 of them.[336] Until about the fourth century, almost all Arabs practised polytheistic religions.[337] Although significant Jewish and Christian minorities developed, polytheism remained the dominant belief system in pre-Islamic Arabia.[75][338]
+The religious beliefs and practices of the nomadic bedouin were distinct from those of the settled tribes of towns such as Mecca.[339] Nomadic religious belief systems and practices are believed to have included fetishism, totemism and veneration of the dead but were connected principally with immediate concerns and problems and did not consider larger philosophical questions such as the afterlife.[339] Settled urban Arabs, on the other hand, are thought to have believed in a more complex pantheon of deities.[339] While the Meccans and the other settled inhabitants of the Hejaz worshipped their gods at permanent shrines in towns and oases, the bedouin practised their religion on the move.[340]
+Arabic philosophy refers to philosophical thought in the Arab world. Schools of Arabic thought include Avicennism and Averroism. The first great Arab thinker is widely regarded to be al-Kindi (801–873 A.D.), a Neo-Platonic philosopher, mathematician and scientist who lived in Kufa and Baghdad (modern day Iraq). After being appointed by the Abbasid Caliphs to translate Greek scientific and philosophical texts into Arabic, he wrote a number of original treatises of his own on a range of subjects, from metaphysics and ethics to mathematics and pharmacology.[341]
+Much of his philosophical output focuses on theological subjects such as the nature of God, the soul and prophetic knowledge.[342] Doctrines of the Arabic philosophers of the 9th–12th century who influenced medieval Scholasticism in Europe. The Arabic tradition combines Aristotelianism and Neoplatonism with other ideas introduced through Islam. Influential thinkers include the Persians al-Farabi and Avicenna. The Arabic philosophic literature was translated into Hebrew and Latin, this contributed to the development of modern European philosophy. The Arabic tradition was developed by Moses Maimonides and Ibn Khaldun.[343][344]
+Arabic science underwent considerable development during the 8th to 13th centuries CE, a source of knowledge that later spread throughout Europe and greatly influenced both medical practice and education. These scientific accomplishments occurred after Muhammad united the Arab tribes.[346]
+Within a century after Muhammed's death (632 CE), an empire ruled by Arabs was established. It encompassed a large part of the planet, stretching from southern Europe to North Africa to Central Asia and on to India. In 711 CE, Arab Muslims invaded southern Spain; al-Andalus was a center of Arabic scientific accomplishment. Another center emerged in Baghdad from the Abbasids, who ruled part of the Islamic world during a historic period later characterized as the "Golden Age" (∼750 to 1258 CE).[347]
+This era can be identified as the years between 692 and 945,[348] and ended when the caliphate was marginalized by local Muslim rulers in Baghdad – its traditional seat of power. From 945 onward until the sacking of Baghdad by the Mongols in 1258, the Caliph continued on as a figurehead, with power devolving more to local amirs.[349] The pious scholars of Islam, men and women collectively known as the ulama, were the most influential element of society in the fields of Sharia law, speculative thought and theology.[350] Arabic scientific achievement is not as yet fully understood, but is very large.[351] These achievements encompass a wide range of subject areas, especially mathematics, astronomy, and medicine.[351] Other subjects of scientific inquiry included physics, alchemy and chemistry, cosmology, ophthalmology, geography and cartography, sociology, and psychology.[352][353]
+Al-Battani (c. 858 – 929; born Harran, Bilad al-Sham) was an Arab astronomer, astrologer and mathematician of the Islamic Golden Age. His work is considered instrumental in the development of science and astronomy. One of Al-Battani's best-known achievements in astronomy was the determination of the solar year as being 365 days, 5 hours, 46 minutes and 24 seconds which is only 2 minutes and 22 seconds off.[354]
+Ibn al-Haytham (Alhazen) used experimentation to obtain the results in his Book of Optics (1021), an important development in the history of the scientific method. He combined observations, experiments and rational arguments to support his intromission theory of vision, in which rays of light are emitted from objects rather than from the eyes. He used similar arguments to show that the ancient emission theory of vision supported by Ptolemy and Euclid (in which the eyes emit the rays of light used for seeing), and the ancient intromission theory supported by Aristotle (where objects emit physical particles to the eyes), were both wrong.[355]
+Al-Zahrawi, regarded by many as the greatest surgeon of the middle ages.[356] His surgical treatise "De chirurgia" is the first illustrated surgical guide ever written. It remained the primary source for surgical procedures and instruments in Europe for the next 500 years.[357] The book helped lay the foundation to establish surgery as a scientific discipline independent from medicine, earning al-Zahrawi his name as one of the founders of this field.[358]
+Other notable Arab contributions include among other things: establishing the science of chemistry by Jābir ibn Hayyān,[359][360][361] establishing the science of cryptology and cryptanalysis by al-Kindi,[362][363][364] the development of analytic geometry by Ibn al-Haytham,[365][366] the discovery of the pulmonary circulation by Ibn al-Nafis,[367][368] the discovery of the itch mite parasite by Ibn Zuhr,[369] the first use of irrational numbers as an algebraic objects by Abū Kāmil,[370] the first use of the positional decimal fractions by al-Uqlidisi,[371][372] the development of the Arabic numerals and an early algebraic symbolism in the Maghreb,[373][374] the Thabit number and Thābit theorem by Thābit ibn Qurra,[375] the discovery of several new trigonometric identities by Ibn Yunus and al-Battani,[376][377] the mathematical proof for Ceva's theorem by Ibn Hűd,[378] the first accurate lunar model by Ibn al-Shatir,[citation needed] the invention of the torquetum by Jabir ibn Aflah,[379] the invention of the universal astrolabe and the equatorium by al-Zarqali,[380][381] the first description of the crankshaft by al-Jazari,[382][383] the anticipation of the inertia concept by Averroes,[384] the discovery of the physical reaction by Avempace,[citation needed] the identification of more than 200 new plants by Ibn al-Baitar[385] the Arab Agricultural Revolution, and the Tabula Rogeriana, which was the most accurate world map in pre-modern times by al-Idrisi.[386]
+The birth of the University institution can be traced to this development, as several universities and educational institutions of the Arab world such as the University of Al Quaraouiyine, Al Azhar University, and Al Zaytuna University are considered to be the oldest in the world. Founded by Fatima al Fihri in 859, the University of Al Quaraouiyine in Fez is the oldest existing, continually operating and the first degree awarding educational institution in the world according to UNESCO and Guinness World Records[387][388] and is sometimes referred to as the oldest university.[389]
+There are many scientific Arabic loanwords in Western European languages, including English, mostly via Old French.[390] This includes traditional star names such as Aldebaran, scientific terms like alchemy (whence also chemistry), algebra, algorithm, alcohol, alkali, cipher, zenith, etc.
+Under Ottoman rule, cultural life and science in the Arab world declined. In the 20th and 21st centuries, Arabs who have won important science prizes include Ahmed Zewail and Elias Corey (Nobel Prize), Michael DeBakey and Alim Benabid (Lasker Award), Omar M. Yaghi (Wolf Prize), Huda Zoghbi (Shaw Prize), Zaha Hadid (Pritzker Prize), and Michael Atiyah (both Fields Medal and Abel Prize). Rachid Yazami was one of the co-inventors of the lithium-ion battery,[391] and Tony Fadell was important in the development of the iPod and the iPhone.[392]
+Arabic weddings have changed greatly in the past 100 years. Original traditional Arabic weddings are supposed to be very similar to modern-day Bedouin weddings and rural weddings, and they are in some cases unique from one region to another, even within the same country. The practice of marrying of relatives is a common feature of Arab culture.[393]
+In the Arab world today between 40% and 50% of all marriages are consanguineous or between close family members, though these figures may vary among Arab nations.[394][395] In Egypt, around 40% of the population marry a cousin. A 1992 survey in Jordan found that 32% were married to a first cousin; a further 17.3% were married to more distant relatives.[396] 67% of marriages in Saudi Arabia are between close relatives as are 54% of all marriages in Kuwait, whereas 18% of all Lebanese were between blood relatives.[397][397] Due to the actions of Muhammad and the Rightly Guided Caliphs, marriage between cousins is explicitly allowed in Islam and the Qur'an itself does not discourage or forbid the practice.[398] Nevertheless, opinions vary on whether the phenomenon should be seen as exclusively based on Islamic practices as a 1992 study among Arabs in Jordan did not show significant differences between Christian Arabs or Muslim Arabs when comparing the occurrence of consanguinity.[397]
+E1b1b is the most frequent paternal clade among the populations in the western part of the Arab world (Maghreb, Nile Valley, and the Horn of Africa), whereas haplogroup J is the most frequent paternal clade toward the east (Arabian peninsula and Near East). Other less common haplogroups are R1a, R1b, G, I, L and T.[399][400][401][402][403][404][405][406][407][408][409][410][411]
+Listed here are the human Y-chromosome DNA haplogroups in Arabian Peninsula, Mashriq or Levant, Maghreb and Nile Valley.[412][413][414][415][416][417][418] Yemeni Arabs J (82.3%), E1b1b (12.9%) and E1b1a1-M2 (3.2%).[419][420] Saudi Arabs J-M267 (Y-DNA) (71.02%), J-M172 (2.68%), A (0.83%), B (1.67%), E1b1a (1.50%), E1b1b (11.05%), G (1.34%), H (0.33%), L (1.00%), Q (1.34%), R1a (2.34%), R1b (0.83%), T (2.51%), UP (1.50%).[421][422][423] Emirati Arabs J (45.1%), E1b1b (11.6%), R1a (7.3%), E1b1a1-M2 (5.5%), T (4.9%), R1b (4.3%) and L (3%).[419] Omani Arabs J (47.9%), E1b1b (15.7%), R1a (9.1%), T (8.3%), E1b1a (7.4%), R1b (1.7%), G (1.7%) and L (0.8%).[424] Qatari Arabs J (66.7%), R1a (6.9%), E1b1b (5.6%), E1b1a (2.8%), G (2.8%) and L (2.8%).[425][426] Lebanese Arabs J (45.2%), E1b1b (25.8%), R1a (9.7%), R1b (6.4%), G, I and I (3.2%), (3.2%), (3.2%).[427] Syrian Arabs J (58.3%),[428][429] E1b1b (12.0%), I (5.0%), R1a (10.0%) and R1b 15.0%.[427][429] Palestinian Arabs J (55.2%), E1b1b (20.3%), R1b (8.4%), I (6.3%), G (7%), R1a and T (1.4%), (1.4%).[430][431] Jordanian Arabs J (43.8%), E1b1b (26%), R1b (17.8%), G (4.1%), I (3.4%) and R1a (1.4%).[432] Iraqi Arabs J (50.6%), E1b1b (10.8%), R1b (10.8%), R1a (6.9%) and T (5.9%).[433][434] Egyptian Arabs E1b1b (36.7%) and J (32%), G (8.8%), T (8.2% R1b (4.1%), E1b1a (2.8%) and I (0.7%).[414][435] Sudanese Arabs J (47.1%), E1b1b (16.3%), R1b (15.7%) and I (3.13%).[436][437] Moroccan Arabs E1b1b (75.5%) and J1 (20.4%).[438][439] Tunisian Arabs E1b1b (49.3%), J1 (35.8%), R1b (6.8%) and E1b1a1-M2 (1.4%).[440] Algerian Arabs E1b1b (54%), J1 (35%), R1b (13%).[440] Libyan Arabs E1b1b (35.88%), J (30.53%), E1b1a (8.78%), G (4.20%), R1a/R1b (3.43%) and E (1.53%).[441][442]
+The mtDNA haplogroup J has been observed at notable frequencies among overall populations in the Arab world.[443][444] The maternal clade R0 reaches its highest frequency in the Arabian peninsula,[445] while K and T(specifically subclade T2) is more common in the Levant.[443] In the Nile Valley and Horn of Africa, haplogroups N1 and M1;[445] in the Maghreb, haplogroups H1 and U6 are more significant.[446]
+There are four principal West Eurasian autosomal DNA components that characterize the populations in the Arab world: the Arabian, Levantine, Coptic and Maghrebi components.
+The Arabian component is the main autosomal element in the Persian Gulf region. It is most closely associated with local Arabic-speaking populations.[417] The Arabian component is also found at significant frequencies in parts of the Levant and Northeast Africa.[417][447] The geographical distribution pattern of this component correlates with the pattern of the Islamic expansion, but its presence in Lebanese Christians, Sephardi and Ashkenazi Jews, Cypriots and Armenians might suggest that its spread to the Levant could also represent an earlier event.[417]
+The Levantine component is the main autosomal element in the Near East and Caucasus. It peaks among Druze populations in the Levant. The Levantine component diverged from the Arabian component about 15,500–23,700 ypb.[417]
+The Coptic component is the main autosomal element in Northeast Africa. It peaks among Egyptian Copts in Sudan, and is also found at high frequencies among other Afro-Asiatic-speaking populations in the Nile Valley and the Horn of Africa.[448] The Coptic component is roughly equivalent with the Ethio-Somali component.[449]
+The Maghrebi component is the main autosomal element in the Maghreb. It peaks among the non-Arabized Berber populations in the region.[447] The Maghrebi component diverged from the Coptic/Ethio-Somali, Arabian and Levantine components prior to the Holocene.[447][449]
+A recent genetic study published in the "European Journal of Human Genetics" in Nature (2019) showed that West Asians (Arabs) are closely related to Europeans, Northern Africans and South Asians.[450]

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+A fixed-wing aircraft is a flying machine, such as an airplane (or aeroplane; see spelling differences), which is capable of flight using wings that generate lift caused by the aircraft's forward airspeed and the shape of the wings. Fixed-wing aircraft are distinct from rotary-wing aircraft (in which the wings form a rotor mounted on a spinning shaft or "mast"), and ornithopters (in which the wings flap in a manner similar to that of a bird). The wings of a fixed-wing aircraft are not necessarily rigid; kites, hang gliders, variable-sweep wing aircraft and airplanes that use wing morphing are all examples of fixed-wing aircraft.
+Gliding fixed-wing aircraft, including free-flying gliders of various kinds and tethered kites, can use moving air to gain altitude. Powered fixed-wing aircraft (airplanes) that gain forward thrust from an engine include powered paragliders, powered hang gliders and some ground effect vehicles. Most fixed-wing aircraft are flown by a pilot on board the craft, but some are specifically designed to be unmanned and controlled either remotely or autonomously (using onboard computers).
+Kites were used approximately 2,800 years ago in China, where materials ideal for kite building were readily available. Some authors hold that leaf kites were being flown much earlier in what is now Sulawesi, based on their interpretation of cave paintings on Muna Island off Sulawesi.[1] By at least 549 AD paper kites were being flown, as it was recorded in that year a paper kite was used as a message for a rescue mission.[2] Ancient and medieval Chinese sources list other uses of kites for measuring distances, testing the wind, lifting men, signaling, and communication for military operations.[2]
+Stories of kites were brought to Europe by Marco Polo towards the end of the 13th century, and kites were brought back by sailors from Japan and Malaysia in the 16th and 17th centuries.[3] Although they were initially regarded as mere curiosities, by the 18th and 19th centuries kites were being used as vehicles for scientific research.[3]
+Around 400 BC in Greece, Archytas was reputed to have designed and built the first artificial, self-propelled flying device, a bird-shaped model propelled by a jet of what was probably steam, said to have flown some 200 m (660 ft).[4][5] This machine may have been suspended for its flight.[6][7]
+One of the earliest purported attempts with gliders was by the 11th-century monk Eilmer of Malmesbury, which ended in failure. A 17th-century account states that the 9th-century poet Abbas Ibn Firnas made a similar attempt, though no earlier sources record this event.[8]
+In 1799, Sir George Cayley set forth the concept of the modern airplane as a fixed-wing flying machine with separate systems for lift, propulsion, and control.[9][10] Cayley was building and flying models of fixed-wing aircraft as early as 1803, and he built a successful passenger-carrying glider in 1853.[11] In 1856, Frenchman Jean-Marie Le Bris made the first powered flight, by having his glider "L'Albatros artificiel" pulled by a horse on a beach.[citation needed] In 1884, the American John J. Montgomery made controlled flights in a glider as a part of a series of gliders built between 1883–1886.[12] Other aviators who made similar flights at that time were Otto Lilienthal, Percy Pilcher, and protégés of Octave Chanute.
+In the 1890s, Lawrence Hargrave conducted research on wing structures and developed a box kite that lifted the weight of a man. His box kite designs were widely adopted. Although he also developed a type of rotary aircraft engine, he did not create and fly a powered fixed-wing aircraft.[13]
+Sir Hiram Maxim built a craft that weighed 3.5 tons, with a 110-foot (34-meter) wingspan that was powered by two 360-horsepower (270-kW) steam engines driving two propellers. In 1894, his machine was tested with overhead rails to prevent it from rising. The test showed that it had enough lift to take off. The craft was uncontrollable, which Maxim, it is presumed, realized, because he subsequently abandoned work on it.[14]
+The Wright brothers' flights in 1903 with their Flyer I are recognized by the Fédération Aéronautique Internationale (FAI), the standard setting and record-keeping body for aeronautics, as "the first sustained and controlled heavier-than-air powered flight".[15] By 1905, the Wright Flyer III was capable of fully controllable, stable flight for substantial periods.
+In 1906, Brazilian inventor Alberto Santos Dumont designed, built and piloted an aircraft that set the first world record recognized by the Aéro-Club de France by flying the 14 bis 220 metres (720 ft) in less than 22 seconds.[16] The flight was certified by the FAI.[17]
+The Bleriot VIII design of 1908 was an early aircraft design that had the modern monoplane tractor configuration. It had movable tail surfaces controlling both yaw and pitch, a form of roll control supplied either by wing warping or by ailerons and controlled by its pilot with a joystick and rudder bar. It was an important predecessor of his later Bleriot XI Channel-crossing aircraft of the summer of 1909.[18]
+World War I served as a testbed for the use of the aircraft as a weapon. Aircraft demonstrated their potential as mobile observation platforms, then proved themselves to be machines of war capable of causing casualties to the enemy. The earliest known aerial victory with a synchronized machine gun-armed fighter aircraft occurred in 1915, by German Luftstreitkräfte Leutnant Kurt Wintgens. Fighter aces appeared; the greatest (by number of air victories) was Manfred von Richthofen.
+Following WWI, aircraft technology continued to develop. Alcock and Brown crossed the Atlantic non-stop for the first time in 1919. The first commercial flights took place between the United States and Canada in 1919.
+The so-called Golden Age of Aviation occurred between the two World Wars, during which both updated interpretations of earlier breakthroughs – as with Hugo Junkers' pioneering of all-metal airframes in 1915 leading to giant multi-engined aircraft of up to 60+ meter wingspan sizes by the early 1930s, adoption of the mostly air-cooled radial engine as a practical aircraft powerplant alongside powerful V-12 liquid-cooled aviation engines, and ever-greater instances of long-distance flight attempts – as with a Vickers Vimy in 1919, followed only months later by the U.S. Navy's NC-4 transatlantic flight; culminating in May 1927 with Charles Lindbergh's solo trans-Atlantic flight in the Spirit of St. Louis spurring ever-longer flight attempts, pioneering the way for long-distance flights of the future to become commonplace.
+Airplanes had a presence in all the major battles of World War II. They were an essential component of the military strategies of the period, such as the German Blitzkrieg or the American and Japanese aircraft carrier campaigns of the Pacific.
+Military gliders were developed and used in several campaigns, but they did not become widely used due to the high casualty rate often encountered. The Focke-Achgelis Fa 330 Bachstelze (Wagtail) rotor kite of 1942 was notable for its use by German submarines.
+Before and during the war, both British and German designers were developing jet engines to power airplanes. The first jet aircraft to fly, in 1939, was the German Heinkel He 178. In 1943, the first operational jet fighter, the Messerschmitt Me 262, went into service with the German Luftwaffe and later in the war the British Gloster Meteor entered service but never saw action – top airspeeds of aircraft for that era went as high as 1,130 km/h (702 mph), with the early July 1944 unofficial record flight of the German Me 163B V18 rocket fighter prototype.[19]
+In October 1947, the Bell X-1 was the first aircraft to exceed the speed of sound.[20]
+In 1948–49, aircraft transported supplies during the Berlin Blockade. New aircraft types, such as the B-52, were produced during the Cold War.
+The first jet airliner, the de Havilland Comet, was introduced in 1952, followed by the Soviet Tupolev Tu-104 in 1956. The Boeing 707, the first widely successful commercial jet, was in commercial service for more than 50 years, from 1958 to 2010. The Boeing 747 was the world's biggest passenger aircraft from 1970 until it was surpassed by the Airbus A380 in 2005.
+An airplane (also known as an aeroplane or simply a plane) is a powered fixed-wing aircraft that is propelled forward by thrust from a jet engine or propeller. Planes come in a variety of sizes, shapes, and wing configurations. The broad spectrum of uses for planes includes recreation, transportation of goods and people, military, and research.
+A seaplane is a fixed-wing aircraft capable of taking off and landing (alighting) on water. Seaplanes that can also operate from dry land are a subclass called amphibian aircraft. These aircraft were sometimes called hydroplanes.[21] Seaplanes and amphibians are usually divided into two categories based on their technological characteristics: floatplanes and flying boats.
+Many forms of glider (see below) may be modified by adding a small power plant. These include:
+A ground effect vehicle (GEV) is a craft that attains level flight near the surface of the earth, making use of the ground effect – an aerodynamic interaction between the wings and the earth's surface. Some GEVs are able to fly higher out of ground effect (OGE) when required – these are classed as powered fixed-wing aircraft.[22]
+A glider is a heavier-than-air craft that is supported in flight by the dynamic reaction of the air against its lifting surfaces, and whose free flight does not depend on an engine. A sailplane is a fixed-wing glider designed for soaring – the ability to gain height in updrafts of air and to fly for long periods.
+Gliders are mainly used for recreation, but have also been used for other purposes such as aerodynamics research, warfare and recovering spacecraft.
+A motor glider does have an engine for extending its performance and some have engines powerful enough to take off, but the engine is not used in normal flight.
+As is the case with planes, there are a wide variety of glider types differing in the construction of their wings, aerodynamic efficiency, location of the pilot and controls. Perhaps the most familiar type is the toy paper plane.
+Large gliders are most commonly launched by a tow-plane or by a winch. Military gliders have been used in war to deliver assault troops, and specialized gliders have been used in atmospheric and aerodynamic research. Rocket-powered aircraft and spaceplanes have also made unpowered landings.
+Gliders and sailplanes that are used for the sport of gliding have high aerodynamic efficiency. The highest lift-to-drag ratio is 70:1, though 50:1 is more common. After launch, further energy is obtained through the skillful exploitation of rising air in the atmosphere. Flights of thousands of kilometers at average speeds over 200 km/h have been achieved.
+The most numerous unpowered aircraft are paper airplanes, a handmade type of glider. Like hang gliders and paragliders, they are foot-launched and are in general slower, smaller, and less expensive than sailplanes. Hang gliders most often have flexible wings given shape by a frame, though some have rigid wings. Paragliders and paper airplanes have no frames in their wings.
+Gliders and sailplanes can share a number of features in common with powered aircraft, including many of the same types of fuselage and wing structures. For example, the Horten H.IV was a tailless flying wing glider, and the delta wing-shaped Space Shuttle orbiter flew much like a conventional glider in the lower atmosphere. Many gliders also use similar controls and instruments as powered craft.
+The main application today of glider aircraft is sport and recreation.
+Gliders were developed from the 1920s for recreational purposes. As pilots began to understand how to use rising air, sailplane gliders were developed with a high lift-to-drag ratio. These allowed longer glides to the next source of "lift", and so increase their chances of flying long distances. This gave rise to the popular sport of gliding.
+Early gliders were mainly built of wood and metal but the majority of sailplanes now use composite materials incorporating glass, carbon or aramid fibers. To minimize drag, these types have a streamlined fuselage and long narrow wings having a high aspect ratio. Both single-seat and two-seat gliders are available.
+Initially training was done by short "hops" in primary gliders which are very basic aircraft with no cockpit and minimal instruments.[23] Since shortly after World War II training has always been done in two-seat dual control gliders, but high performance two-seaters are also used to share the workload and the enjoyment of long flights. Originally skids were used for landing, but the majority now land on wheels, often retractable. Some gliders, known as motor gliders, are designed for unpowered flight, but can deploy piston, rotary, jet or electric engines.[24] Gliders are classified by the FAI for competitions into glider competition classes mainly on the basis of span and flaps.
+A class of ultralight sailplanes, including some known as microlift gliders and some known as "airchairs", has been defined by the FAI based on a maximum weight. They are light enough to be transported easily, and can be flown without licensing in some countries. Ultralight gliders have performance similar to hang gliders, but offer some additional crash safety as the pilot can be strapped in an upright seat within a deformable structure. Landing is usually on one or two wheels which distinguishes these craft from hang gliders. Several commercial ultralight gliders have come and gone, but most current development is done by individual designers and home builders.
+Military gliders were used during World War II for carrying troops (glider infantry) and heavy equipment to combat zones. The gliders were towed into the air and most of the way to their target by military transport planes, e.g. C-47 Dakota, or by bombers that had been relegated to secondary activities, e.g. Short Stirling. Once released from the tow near the target, they landed as close to the target as possible. The advantage over paratroopers were that heavy equipment could be landed and that the troops were quickly assembled rather than being dispersed over a drop zone. The gliders were treated as disposable, leading to construction from common and inexpensive materials such as wood, though a few were retrieved and re-used. By the time of the Korean War, transport aircraft had also become larger and more efficient so that even light tanks could be dropped by parachute, causing gliders to fall out of favor.
+Even after the development of powered aircraft, gliders continued to be used for aviation research. The NASA Paresev Rogallo flexible wing was originally developed to investigate alternative methods of recovering spacecraft. Although this application was abandoned, publicity inspired hobbyists to adapt the flexible-wing airfoil for modern hang gliders.
+Initial research into many types of fixed-wing craft, including flying wings and lifting bodies was also carried out using unpowered prototypes.
+A hang glider is a glider aircraft in which the pilot is ensconced in a harness suspended from the airframe, and exercises control by shifting body weight in opposition to a control frame. Most modern hang gliders are made of an aluminum alloy or composite-framed fabric wing. Pilots have the ability to soar for hours, gain thousands of meters of altitude in thermal updrafts, perform aerobatics, and glide cross-country for hundreds of kilometers.
+A paraglider is a lightweight, free-flying, foot-launched glider aircraft with no rigid primary structure.[25] The pilot sits in a harness suspended below a hollow fabric wing whose shape is formed by its suspension lines, the pressure of air entering vents in the front of the wing and the aerodynamic forces of the air flowing over the outside. Paragliding is most often a recreational activity.
+A paper plane is a toy aircraft (usually a glider) made out of paper or paperboard.
+Model glider aircraft are models of aircraft using lightweight materials such as polystyrene and balsa wood. Designs range from simple glider aircraft to accurate scale models, some of which can be very large.
+Glide bombs are bombs with aerodynamic surfaces to allow a gliding flightpath rather than a ballistic one. This enables the carrying aircraft to attack a heavily defended target from a distance.
+A kite is an aircraft tethered to a fixed point so that the wind blows over its wings.[26] Lift is generated when air flows over the kite's wing, producing low pressure above the wing and high pressure below it, and deflecting the airflow downwards. This deflection also generates horizontal drag in the direction of the wind. The resultant force vector from the lift and drag force components is opposed by the tension of the one or more rope lines or tethers attached to the wing.
+Kites are mostly flown for recreational purposes, but have many other uses. Early pioneers such as the Wright Brothers and J.W. Dunne sometimes flew an aircraft as a kite in order to develop it and confirm its flight characteristics, before adding an engine and flight controls, and flying it as an airplane.
+Kites have been used for signaling, for delivery of munitions, and for observation, by lifting an observer above the field of battle, and by using kite aerial photography.
+Kites have been used for scientific purposes, such as Benjamin Franklin's famous experiment proving that lightning is electricity. Kites were the precursors to the traditional aircraft, and were instrumental in the development of early flying craft. Alexander Graham Bell experimented with very large man-lifting kites, as did the Wright brothers and Lawrence Hargrave. Kites had a historical role in lifting scientific instruments to measure atmospheric conditions for weather forecasting.
+Kites can be used to carry radio antennas. This method was used for the reception station of the first transatlantic transmission by Marconi. Captive balloons may be more convenient for such experiments, because kite-carried antennas require a lot of wind, which may be not always possible with heavy equipment and a ground conductor.
+Kites can be used to carry light effects such as lightsticks or battery powered lights.
+Kites can be used to pull people and vehicles downwind. Efficient foil-type kites such as power kites can also be used to sail upwind under the same principles as used by other sailing craft, provided that lateral forces on the ground or in the water are redirected as with the keels, center boards, wheels and ice blades of traditional sailing craft. In the last two decades, several kite sailing sports have become popular, such as kite buggying, kite landboarding, kite boating and kite surfing. Snow kiting has also become popular.
+Kite sailing opens several possibilities not available in traditional sailing:
+Conceptual research and development projects are being undertaken by over a hundred participants to investigate the use of kites in harnessing high altitude wind currents for electricity generation.[27]
+Kite festivals are a popular form of entertainment throughout the world. They include local events, traditional festivals and major international festivals.
+The structural parts of a fixed-wing aircraft are called the airframe. The parts present can vary according to the aircraft's type and purpose. Early types were usually made of wood with fabric wing surfaces, When engines became available for a powered flight around a hundred years ago, their mounts were made of metal. Then as speeds increased more and more parts became metal until by the end of WWII all-metal aircraft were common. In modern times, increasing use of composite materials has been made.
+Typical structural parts include:
+The wings of a fixed-wing aircraft are static planes extending either side of the aircraft. When the aircraft travels forwards,
+air flows over the wings which are shaped to create lift.
+Kites and some light weight gliders and airplanes have flexible wing surfaces which are stretched across a frame and made rigid by the lift forces exerted by the airflow over them. Larger aircraft have rigid wing surfaces which provide additional strength.
+Whether flexible or rigid, most wings have a strong frame to give them their shape and to transfer lift from the wing surface to the rest of the aircraft. The main structural elements are one or more spars running from root to tip, and many ribs running from the leading (front) to the trailing (rear) edge.
+Early airplane engines had little power and light weight was very important. Also, early aerofoil sections were very thin, and could not have strong frame installed within. So until the 1930s, most wings were too light weight to have enough strength and external bracing struts and wires were added. When the available engine power increased during the 1920s and 1930s, wings could be made heavy and strong enough that bracing was not needed anymore. This type of unbraced wing is called a cantilever wing.
+The number and shape of the wings vary widely on different types. A given wing plane may be full-span or divided by a central fuselage into port (left) and starboard (right) wings. Occasionally, even more, wings have been used, with the three-winged triplane achieving some fame in WWI. The four-winged quadruplane and other Multiplane (Aeronautics) designs have had little success.
+A monoplane, which derives from the prefix, mono means one which means it has a single wing plane, a biplane has two stacked one above the other, a tandem wing has two placed one behind the other. When the available engine power increased during the 1920s and 1930s and bracing was no longer needed, the unbraced or cantilever monoplane became the most common form of powered type.
+The wing planform is the shape when seen from above. To be aerodynamically efficient, a wing should be straight with a long span from side to side but have a short chord (high aspect ratio). But to be structurally efficient, and hence lightweight, a wing must have a short span but still enough area to provide lift (low aspect ratio).
+At transonic speeds, near the speed of sound, it helps to sweep the wing backward or forwards to reduce drag from supersonic shock waves as they begin to form. The swept wing is just a straight wing swept backward or forwards.
+The delta wing is a triangle shape which may be used for a number of reasons. As a flexible Rogallo wing it allows a stable shape under aerodynamic forces, and so is often used for kites and other ultralight craft. As a supersonic wing, it combines high strength with low drag and so is often used for fast jets.
+A variable geometry wing can be changed in flight to a different shape. The variable-sweep wing transforms between an efficient straight configuration for takeoff and landing, to a low-drag swept configuration for high-speed flight. Other forms of variable planform have been flown, but none have gone beyond the research stage.
+A fuselage is a long, thin body, usually with tapered or rounded ends to make its shape aerodynamically smooth. The fuselage may contain the flight crew, passengers, cargo or payload, fuel and engines. The pilots of manned aircraft operate them from a cockpit located at the front or top of the fuselage and equipped with controls and usually windows and instruments. A plane may have more than one fuselage, or it may be fitted with booms with the tail located between the booms to allow the extreme rear of the fuselage to be useful for a variety of purposes.
+A flying wing is a tailless aircraft which has no definite fuselage, with most of the crew, payload and equipment being housed inside the main wing structure.[28]:224
+The flying wing configuration was studied extensively in the 1930s and 1940s, notably by Jack Northrop and Cheston L. Eshelman in the United States, and Alexander Lippisch and the Horten brothers in Germany.
+After the war, a number of experimental designs were based on the flying wing concept. Some general interest continued until the early 1950s, but designs did not necessarily offer a great advantage in range and presented a number of technical problems, leading to the adoption of "conventional" solutions like the Convair B-36 and the B-52 Stratofortress. Due to the practical need for a deep wing, the flying wing concept is most practical for designs in the slow-to-medium speed range, and there has been continual interest in using it as a tactical airlifter design.
+Interest in flying wings was renewed in the 1980s due to their potentially low radar reflection cross-sections. Stealth technology relies on shapes which only reflect radar waves in certain directions, thus making the aircraft hard to detect unless the radar receiver is at a specific position relative to the aircraft – a position that changes continuously as the aircraft moves. This approach eventually led to the Northrop B-2 Spirit stealth bomber. In this case the aerodynamic advantages of the flying wing are not the primary needs. However, modern computer-controlled fly-by-wire systems allowed for many of the aerodynamic drawbacks of the flying wing to be minimized, making for an efficient and stable long-range bomber.
+Blended wing body aircraft have a flattened and airfoil shaped body, which produces most of the lift to keep itself aloft, and distinct and separate wing structures, though the wings are smoothly blended in with the body.
+Thus blended wing bodied aircraft incorporate design features from both a futuristic fuselage and flying wing design. The purported advantages of the blended wing body approach are efficient high-lift wings and a wide airfoil-shaped body. This enables the entire craft to contribute to lift generation with the result of potentially increased fuel economy.
+A lifting body is a configuration in which the body itself produces lift. In contrast to a flying wing, which is a wing with minimal or no conventional fuselage, a lifting body can be thought of as a fuselage with little or no conventional wing. Whereas a flying wing seeks to maximize cruise efficiency at subsonic speeds by eliminating non-lifting surfaces, lifting bodies generally minimize the drag and structure of a wing for subsonic, supersonic, and hypersonic flight, or, spacecraft re-entry. All of these flight regimes pose challenges for proper flight stability.
+Lifting bodies were a major area of research in the 1960s and 1970s as a means to build a small and lightweight manned spacecraft. The US built a number of famous lifting body rocket planes to test the concept, as well as several rocket-launched re-entry vehicles that were tested over the Pacific. Interest waned as the US Air Force lost interest in the manned mission, and major development ended during the Space Shuttle design process when it became clear that the highly shaped fuselages made it difficult to fit fuel tankage.
+The classic aerofoil section wing is unstable in flight and difficult to control. Flexible-wing types often rely on an anchor line or the weight of a pilot hanging beneath to maintain the correct attitude. Some free-flying types use an adapted aerofoil that is stable, or other ingenious mechanisms including, most recently, electronic artificial stability.
+But in order to achieve trim, stability and control, most fixed-wing types have an empennage comprising a fin and rudder which act horizontally and a tailplane and elevator which act vertically. This is so common that it is known as the conventional layout. Sometimes there may be two or more fins, spaced out along the tailplane.
+Some types have a horizontal "canard" foreplane ahead of the main wing, instead of behind it.[28]:86[29][30] This foreplane may contribute to the trim, stability or control of the aircraft, or to several of these.
+Kites are controlled by wires running down to the ground. Typically each wire acts as a tether to the part of the kite it is attached to.
+Gliders and airplanes have more complex control systems, especially if they are piloted.
+The main controls allow the pilot to direct the aircraft in the air. Typically these are:
+Other common controls include:
+A craft may have two pilots' seats with dual controls, allowing two pilots to take turns. This is often used for training or for longer flights.
+The control system may allow full or partial automation of flight, such as an autopilot, a wing leveler, or a flight management system. An unmanned aircraft has no pilot but is controlled remotely or via means such as gyroscopes or other forms of autonomous control.
+On manned fixed-wing aircraft, instruments provide information to the pilots, including flight, engines, navigation, communications, and other aircraft systems that may be installed.
+The six basic instruments, sometimes referred to as the "six pack", are as follows:[31]
+Other cockpit instruments might include:
+(Wayback Machine copy)

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+The Beatles were an English rock band formed in Liverpool in 1960. The group, whose best-known line-up comprised John Lennon, Paul McCartney, George Harrison and Ringo Starr, are regarded as the most influential band of all time.[1] They were integral to the development of 1960s counterculture and popular music's recognition as an art form.[2] Rooted in skiffle, beat and 1950s rock and roll, their sound incorporated elements of classical music and traditional pop in innovative ways; the band later explored music styles ranging from ballads and Indian music to psychedelia and hard rock. As pioneers in recording, songwriting and artistic presentation, the Beatles revolutionised many aspects of the music industry and were often publicised as leaders of the era's youth and sociocultural movements.[3]
+Led by primary songwriters Lennon and McCartney, the Beatles built their reputation playing clubs in Liverpool and Hamburg over three years from 1960, initially with Stuart Sutcliffe playing bass. The core trio of Lennon, McCartney and Harrison, together since 1958, went through a succession of drummers, including Pete Best, before asking Starr to join them in 1962. Manager Brian Epstein moulded them into a professional act, and producer George Martin guided and developed their recordings, greatly expanding their domestic success after their first hit, "Love Me Do", in late 1962. As their popularity grew into the intense fan frenzy dubbed "Beatlemania", the band acquired the nickname "the Fab Four", with Epstein, Martin and other members of the band's entourage sometimes given the informal title of "fifth Beatle".
+By early 1964, the Beatles were international stars, leading the "British Invasion" of the United States pop market and breaking numerous sales records. They soon made their film debut with A Hard Day's Night (1964). From 1965 onwards, they produced records of greater complexity, including the albums Rubber Soul (1965), Revolver (1966) and Sgt. Pepper's Lonely Hearts Club Band (1967), and enjoyed further commercial success with The Beatles (also known as "the White Album", 1968) and Abbey Road (1969). In 1968, they founded Apple Corps, a multi-armed multimedia corporation that continues to oversee projects related to the band's legacy. After the group's break-up in 1970, all four members enjoyed success as solo artists. Lennon was shot and killed in December 1980, and Harrison died of lung cancer in November 2001. McCartney and Starr remain musically active.
+The Beatles are the best-selling music act of all time, with estimated sales of 600 million units worldwide.[4] They are the best-selling act in the US, with certified sales of 183 million units. They hold the record for most number-one albums on the UK Albums Chart, most number-one hits on the Billboard Hot 100 chart, and most singles sold in the UK. The group were inducted into the Rock and Roll Hall of Fame in 1988, and all four main members were inducted individually between 1994 and 2015. In 2008, the group topped Billboard's list of the all-time most successful artists on the Billboard Hot 100. The band received seven Grammy Awards, four Brit Awards, an Academy Award (for Best Original Song Score for the 1970 film Let It Be) and fifteen Ivor Novello Awards. Time magazine named them among the 20th century's 100 most important people.
+In March 1957, John Lennon, then aged sixteen, formed a skiffle group with several friends from Quarry Bank High School in Liverpool. They briefly called themselves the Blackjacks, before changing their name to the Quarrymen after discovering that another local group were already using the name.[5] Fifteen-year-old Paul McCartney joined them as a rhythm guitarist shortly after he and Lennon met that July.[6] In February 1958, McCartney invited his friend George Harrison to watch the band. The fifteen-year-old auditioned for Lennon, impressing him with his playing, but Lennon initially thought Harrison was too young for the band. After a month of Harrison's persistence, during a second meeting (arranged by McCartney), he performed the lead guitar part of the instrumental song "Raunchy" on the upper deck of a Liverpool bus,[7] and they enlisted him as their lead guitarist.[8][9]
+By January 1959, Lennon's Quarry Bank friends had left the group, and he began his studies at the Liverpool College of Art.[10] The three guitarists, billing themselves as Johnny and the Moondogs,[11] were playing rock and roll whenever they could find a drummer.[12] Lennon's art school friend Stuart Sutcliffe, who had just sold one of his paintings and was persuaded to purchase a bass guitar with the proceeds, joined in January 1960, and it was he who suggested changing the band's name to Beatals, as a tribute to Buddy Holly and the Crickets.[13][14] They used this name until May, when they became the Silver Beetles, before undertaking a brief tour of Scotland as the backing group for pop singer and fellow Liverpudlian Johnny Gentle. By early July, they had refashioned themselves as the Silver Beatles, and by the middle of August shortened the name to the Beatles.[15]
+Allan Williams, the Beatles' unofficial manager, arranged a residency for them in Hamburg, and for this they auditioned and hired drummer Pete Best in mid-August 1960. The band, now a five-piece, departed Liverpool for Hamburg four days later, contracted to club owner Bruno Koschmider for what would be a 31⁄2-month residency.[16] Beatles historian Mark Lewisohn writes: "They pulled into Hamburg at dusk on 17 August, the time when the red-light area comes to life ... flashing neon lights screamed out the various entertainment on offer, while scantily clad women sat unabashed in shop windows waiting for business opportunities."[17]
+Koschmider had converted a couple of strip clubs in the district into music venues, and he initially placed the Beatles at the Indra Club. After closing Indra due to noise complaints, he moved them to the Kaiserkeller in October.[18] When he learned they had been performing at the rival Top Ten Club in breach of their contract, he gave the band one month's termination notice,[19] and reported the underage Harrison, who had obtained permission to stay in Hamburg by lying to the German authorities about his age.[20] The authorities arranged for Harrison's deportation in late November.[21] One week later, Koschmider had McCartney and Best arrested for arson after they set fire to a condom in a concrete corridor; the authorities deported them.[22] Lennon returned to Liverpool in early December, while Sutcliffe remained in Hamburg until late February with his German fiancée Astrid Kirchherr,[23] who took the first semi-professional photos of the Beatles.[24]
+During the next two years, the Beatles were resident for periods in Hamburg, where they used Preludin both recreationally and to maintain their energy through all-night performances.[25] In 1961, during their second Hamburg engagement, Kirchherr cut Sutcliffe's hair in the "exi" (existentialist) style, later adopted by the other Beatles.[26][27] When Sutcliffe decided to leave the band early that year and resume his art studies in Germany, McCartney took up the bass.[28] Producer Bert Kaempfert contracted what was now a four-piece group until June 1962, and he used them as Tony Sheridan's backing band on a series of recordings for Polydor Records.[14][29] As part of the sessions, the Beatles were signed to Polydor for one year.[30] Credited to "Tony Sheridan & the Beat Brothers", the single "My Bonnie", recorded in June 1961 and released four months later, reached number 32 on the Musikmarkt chart.[31]
+After the Beatles completed their second Hamburg residency, they enjoyed increasing popularity in Liverpool with the growing Merseybeat movement. However, they were also growing tired of the monotony of numerous appearances at the same clubs night after night.[32] In November 1961, during one of the group's frequent performances at The Cavern Club, they encountered Brian Epstein, a local record-store owner and music columnist.[33] He later recalled: "I immediately liked what I heard. They were fresh, and they were honest, and they had what I thought was a sort of presence ... [a] star quality."[34]
+Epstein courted the band over the next couple of months, and they appointed him as their manager in January 1962.[35] Throughout early and mid-1962, Epstein sought to free the Beatles from their contractual obligations to Bert Kaempfert Productions. He eventually negotiated a one-month-early release from their contract in exchange for one last recording session in Hamburg.[36] Tragedy greeted them on their return to Germany in April, when a distraught Kirchherr met them at the airport with news of Sutcliffe's death the previous day from what was later determined as a brain haemorrhage.[37]
+Epstein began negotiations with record labels for a recording contract. To secure a UK record contract, Epstein negotiated an early end to the band's contract with Polydor, in exchange for more recordings backing Tony Sheridan.[38] After a New Year's Day audition, Decca Records rejected the band with the comment "Guitar groups are on the way out, Mr. Epstein."[39] However, three months later, producer George Martin signed the Beatles to EMI's Parlophone label.[37]
+Martin's first recording session with the Beatles took place at EMI's Abbey Road Studios in London on 6 June 1962.[40] Martin immediately complained to Epstein about Best's poor drumming and suggested they use a session drummer in his place.[41] Already contemplating Best's dismissal,[42] the Beatles replaced him in mid-August with Ringo Starr, who left Rory Storm and the Hurricanes to join them.[40] A 4 September session at EMI yielded a recording of "Love Me Do" featuring Starr on drums, but a dissatisfied Martin hired drummer Andy White for the band's third session a week later, which produced recordings of "Love Me Do", "Please Please Me" and "P.S. I Love You".[40]
+Martin initially selected the Starr version of "Love Me Do" for the band's first single, though subsequent re-pressings featured the White version, with Starr on tambourine.[40] Released in early October, "Love Me Do" peaked at number seventeen on the Record Retailer chart.[43] Their television debut came later that month with a live performance on the regional news programme People and Places.[44] After Martin suggested rerecording "Please Please Me" at a faster tempo,[45] a studio session in late November yielded that recording,[46] of which Martin accurately predicted, "You've just made your first No. 1."[47]
+In December 1962, the Beatles concluded their fifth and final Hamburg residency.[48] By 1963, they had agreed that all four band members would contribute vocals to their albums – including Starr, despite his restricted vocal range, to validate his standing in the group.[49] Lennon and McCartney had established a songwriting partnership, and as the band's success grew, their dominant collaboration limited Harrison's opportunities as a lead vocalist.[50] Epstein, to maximise the Beatles' commercial potential, encouraged them to adopt a professional approach to performing.[51] Lennon recalled him saying, "Look, if you really want to get in these bigger places, you're going to have to change – stop eating on stage, stop swearing, stop smoking ...."[39] Lennon said: "We used to dress how we liked, on and off stage. He'd tell us that jeans were not particularly smart and could we possibly manage to wear proper trousers, but he didn't want us suddenly looking square. He'd let us have our own sense of individuality."[39]
+On 11 February 1963, the Beatles recorded ten songs during a single studio session for their debut LP, Please Please Me. The album was supplemented by the four tracks already released on their first two singles. Martin originally considered recording the Beatles' debut LP live at The Cavern Club, but after deciding that the building's acoustics were inadequate, he elected to simulate a "live" album with minimal production in "a single marathon session at Abbey Road".[53] After the moderate success of "Love Me Do", the single "Please Please Me" met with a more emphatic reception. Released in January 1963, two months ahead of the album of the same name, the song reached number one on every UK chart except Record Retailer, where it peaked at number two.[54]
+Recalling how the Beatles "rushed to deliver a debut album, bashing out Please Please Me in a day", AllMusic's Stephen Thomas Erlewine comments, "Decades after its release, the album still sounds fresh, precisely because of its intense origins."[55] Lennon said little thought went into composition at the time; he and McCartney were "just writing songs à la Everly Brothers, à la Buddy Holly, pop songs with no more thought of them than that – to create a sound. And the words were almost irrelevant."[56]
+Released in March 1963, the album initiated a run during which eleven of their twelve studio albums released in the United Kingdom through to 1970 reached number one.[59] The band's third single, "From Me to You", came out in April and was also a chart-topping hit, starting an almost unbroken string of seventeen British number-one singles for the Beatles, including all but one of the eighteen they released over the next six years.[60] Issued in August, the band's fourth single, "She Loves You", achieved the fastest sales of any record in the UK up to that time, selling three-quarters of a million copies in under four weeks.[61] It became their first single to sell a million copies, and remained the biggest-selling record in the UK until 1978.[62][nb 1]
+Their commercial success brought increased media exposure, to which the Beatles responded with an irreverent and comical attitude that defied the expectations of pop musicians at the time, inspiring even more interest.[63] The band toured the UK three times in the first half of the year: a four-week tour that began in February, the Beatles' first nationwide, preceded three-week tours in March and May–June.[64] As their popularity spread, a frenzied adulation of the group took hold. Greeted with riotous enthusiasm by screaming fans, the press dubbed the phenomenon "Beatlemania".[65] Although not billed as tour leaders, the Beatles overshadowed American acts Tommy Roe and Chris Montez during the February engagements and assumed top billing "by audience demand", something no British act had previously accomplished while touring with artists from the US.[66] A similar situation arose during their May–June tour with Roy Orbison.[67]
+In late October, the Beatles began a five-day tour of Sweden, their first time abroad since the final Hamburg engagement of December 1962.[69] On their return to the UK on 31 October, several hundred screaming fans greeted them in heavy rain at Heathrow Airport. Around 50 to 100 journalists and photographers, as well as representatives from the BBC, also joined the airport reception, the first of more than 100 such events.[70] The next day, the band began its fourth tour of Britain within nine months, this one scheduled for six weeks.[71] In mid-November, as Beatlemania intensified, police resorted to using high-pressure water hoses to control the crowd before a concert in Plymouth.[72]
+Please Please Me maintained the top position on the Record Retailer chart for 30 weeks, only to be displaced by its follow-up, With the Beatles,[73] which EMI released on 22 November to record advance orders of 270,000 copies. The LP topped a half-million albums sold in one week.[74] Recorded between July and October, With the Beatles made better use of studio production techniques than its predecessor.[75] It held the top spot for 21 weeks with a chart life of 40 weeks.[76] Erlewine described the LP as "a sequel of the highest order – one that betters the original".[77]
+In a reversal of then standard practice, EMI released the album ahead of the impending single "I Want to Hold Your Hand", with the song excluded to maximise the single's sales.[78] The album caught the attention of music critic William Mann of The Times, who suggested that Lennon and McCartney were "the outstanding English composers of 1963".[75] The newspaper published a series of articles in which Mann offered detailed analyses of the music, lending it respectability.[79] With the Beatles became the second album in UK chart history to sell a million copies, a figure previously reached only by the 1958 South Pacific soundtrack.[80] When writing the sleeve notes for the album, the band's press officer, Tony Barrow, used the superlative the "fabulous foursome", which the media widely adopted as "the Fab Four".[81]
+EMI's American subsidiary, Capitol Records, hindered the Beatles' releases in the United States for more than a year by initially declining to issue their music, including their first three singles. Concurrent negotiations with the independent US label Vee-Jay led to the release of some of the songs in 1963, but not all.[82] Vee-Jay finished preparation for the album Introducing... The Beatles, culled from most of the songs of Parlophone's Please Please Me, but a management shake-up led to the album not being released.[nb 2] Then when it surfaced that the label did not report royalties on their sales, the licence Vee-Jay signed with EMI was voided.[84] A new licence was granted to the Swan label for the single "She Loves You". The record received some airplay in the Tidewater area of Virginia by Gene Loving of radio station WGH and was featured on the "Rate-a-Record" segment of American Bandstand, but it failed to catch on nationally.[85]
+Epstein arranged for a $40,000 US marketing campaign. American chart success began after disc jockey Carroll James of AM radio station WWDC, in Washington, DC, obtained a copy of the British single "I Want to Hold Your Hand" in mid-December 1963 and began playing it on-air.[86] Taped copies of the song soon circulated among other radio stations throughout the US. This caused an increase in demand, leading Capitol to bring forward the release of "I Want to Hold Your Hand" by three weeks.[87] Issued on 26 December, with the band's previously scheduled debut there just weeks away, "I Want to Hold Your Hand" sold a million copies, becoming a number-one hit in the US by mid-January.[88] In its wake, Vee-Jay released Introducing... The Beatles[89] to go along with Capitol's debut album, Meet the Beatles!, while Swan reactivated production of "She Loves You".[90]
+On 7 February 1964, the Beatles left the UK with an estimated 4,000 fans gathered at Heathrow, waving and screaming as the aircraft took off.[91] Upon landing at New York's John F. Kennedy Airport, an uproarious crowd estimated at 3,000 greeted them.[92] They gave their first live US television performance two days later on The Ed Sullivan Show, watched by approximately 73 million viewers in over 23 million households,[93] or 34 percent of the American population. Biographer Jonathan Gould writes that, according to the Nielsen rating service, it was "the largest audience that had ever been recorded for an American television program".[94] The next morning, the Beatles awoke to a largely negative critical consensus in the US,[95] but a day later at their first US concert, Beatlemania erupted at the Washington Coliseum.[96] Back in New York the following day, the Beatles met with another strong reception during two shows at Carnegie Hall.[93] The band flew to Florida, where they appeared on the weekly Ed Sullivan Show a second time, before another 70 million viewers, before returning to the UK on 22 February.[97]
+The Beatles' first visit to the US took place when the nation were still mourning the assassination of President John F. Kennedy the previous November.[98] Commentators often suggest that for many, particularly the young, the Beatles' performances reignited the sense of excitement and possibility that momentarily faded in the wake of the assassination, and helped make way for the revolutionary social changes to come in the decade.[99] Their hairstyle, unusually long for the era and mocked by many adults,[14] became an emblem of rebellion to the burgeoning youth culture.[100]
+The group's popularity generated unprecedented interest in British music, and many other UK acts subsequently made their American debuts, successfully touring over the next three years in what was termed the British Invasion.[101] The Beatles' success in the US opened the door for a successive string of British beat groups and pop acts such as the Dave Clark Five, the Animals, Petula Clark, the Kinks, and the Rolling Stones to achieve success in America.[102] During the week of 4 April 1964, the Beatles held twelve positions on the Billboard Hot 100 singles chart, including the top five.[103][nb 3]
+Capitol Records' lack of interest throughout 1963 did not go unnoticed, and a competitor, United Artists Records, encouraged their film division to offer the Beatles a three-motion-picture deal, primarily for the commercial potential of the soundtracks in the US.[105] Directed by Richard Lester, A Hard Day's Night involved the band for six weeks in March–April 1964 as they played themselves in a musical comedy.[106] The film premiered in London and New York in July and August, respectively, and was an international success, with some critics drawing a comparison with the Marx Brothers.[107]
+United Artists released a full soundtrack album for the North American market, combining Beatles songs and Martin's orchestral score; elsewhere, the group's third studio LP, A Hard Day's Night, contained songs from the film on side one and other new recordings on side two.[108] According to Erlewine, the album saw them "truly coming into their own as a band. All of the disparate influences on their first two albums coalesced into a bright, joyous, original sound, filled with ringing guitars and irresistible melodies."[109] That "ringing guitar" sound was primarily the product of Harrison's 12-string electric Rickenbacker, a prototype given to him by the manufacturer, which made its debut on the record.[110][nb 4]
+Touring internationally in June and July, the Beatles staged 37 shows over 27 days in Denmark, the Netherlands, Hong Kong, Australia and New Zealand.[111][nb 5] In August and September they returned to the US, with a 30-concert tour of 23 cities.[113] Generating intense interest once again, the month-long tour attracted between 10,000 and 20,000 fans to each 30-minute performance in cities from San Francisco to New York.[113]
+In August, journalist Al Aronowitz arranged for the Beatles to meet Bob Dylan.[114] Visiting the band in their New York hotel suite, Dylan introduced them to cannabis.[115] Gould points out the musical and cultural significance of this meeting, before which the musicians' respective fanbases were "perceived as inhabiting two separate subcultural worlds": Dylan's audience of "college kids with artistic or intellectual leanings, a dawning political and social idealism, and a mildly bohemian style" contrasted with their fans, "veritable 'teenyboppers' – kids in high school or grade school whose lives were totally wrapped up in the commercialised popular culture of television, radio, pop records, fan magazines, and teen fashion. To many of Dylan's followers in the folk music scene, the Beatles were seen as idolaters, not idealists."[116] Within six months of the meeting, according to Gould, "Lennon would be making records on which he openly imitated Dylan's nasal drone, brittle strum, and introspective vocal persona"; and six months after that, Dylan began performing with a backing band and electric instrumentation, and "dressed in the height of Mod fashion".[117] As a result, Gould continues, the traditional division between folk and rock enthusiasts "nearly evaporated", as the Beatles' fans began to mature in their outlook and Dylan's audience embraced the new, youth-driven pop culture.[117]
+During the 1964 US tour, the group were confronted with racial segregation in the country at the time.[118][119] When informed that the venue for their 11 September concert, the Gator Bowl in Jacksonville, Florida, was segregated, the Beatles said they would refuse to perform unless the audience was integrated.[120][118][119] Lennon stated: "We never play to segregated audiences and we aren't going to start now ... I'd sooner lose our appearance money."[118] City officials relented and agreed to allow an integrated show.[118] The group also cancelled their reservations at the whites-only Hotel George Washington in Jacksonville.[119] For their subsequent US tours in 1965 and 1966, the Beatles included clauses in contracts stipulating that shows be integrated.[119][121]
+According to Gould, Beatles for Sale, the Beatles' fourth studio LP, evidenced a growing conflict between the commercial pressures of their global success and their creative ambitions.[122] They had intended the album, recorded between August and October 1964,[123] to continue the format established by A Hard Day's Night which, unlike their first two LPs, contained only original songs.[122] They had nearly exhausted their backlog of songs on the previous album, however, and given the challenges constant international touring posed to their songwriting efforts, Lennon admitted, "Material's becoming a hell of a problem".[124] As a result, six covers from their extensive repertoire were chosen to complete the album. Released in early December, its eight original compositions stood out, demonstrating the growing maturity of the Lennon–McCartney songwriting partnership.[122]
+In early 1965, following a dinner with Lennon, Harrison and their wives, Harrison's dentist, John Riley, secretly added LSD to their coffee.[125] Lennon described the experience: "It was just terrifying, but it was fantastic. I was pretty stunned for a month or two."[126] He and Harrison subsequently became regular users of the drug, joined by Starr on at least one occasion. Harrison's use of psychedelic drugs encouraged his path to meditation and Hinduism. He commented: "For me, it was like a flash. The first time I had acid, it just opened up something in my head that was inside of me, and I realized a lot of things. I didn't learn them because I already knew them, but that happened to be the key that opened the door to reveal them. From the moment I had that, I wanted to have it all the time – these thoughts about the yogis and the Himalayas, and Ravi's music."[127][128] McCartney was initially reluctant to try it, but eventually did so in late 1966.[129] He became the first Beatle to discuss LSD publicly, declaring in a magazine interview that "it opened my eyes" and "made me a better, more honest, more tolerant member of society".[130]
+Controversy erupted in June 1965 when Queen Elizabeth II appointed all four Beatles Members of the Order of the British Empire (MBE) after Prime Minister Harold Wilson nominated them for the award.[131] In protest – the honour was at that time primarily bestowed upon military veterans and civic leaders – some conservative MBE recipients returned their insignia.[132]
+In July, the Beatles' second film, Help!, was released, again directed by Lester. Described as "mainly a relentless spoof of Bond",[133] it inspired a mixed response among both reviewers and the band. McCartney said: "Help! was great but it wasn't our film – we were sort of guest stars. It was fun, but basically, as an idea for a film, it was a bit wrong."[134] The soundtrack was dominated by Lennon, who wrote and sang lead on most of its songs, including the two singles: "Help!" and "Ticket to Ride".[135] The accompanying Help! album, the group's fifth studio LP, mirrored A Hard Day's Night by featuring soundtrack songs on side one and additional songs from the same sessions on side two.[136] The LP contained all original material save for two covers, "Act Naturally" and "Dizzy Miss Lizzy"; they were the last covers the band would include on an album, except for Let It Be's brief rendition of the traditional Liverpool folk song "Maggie Mae".[137] The band expanded their use of vocal overdubs on Help! and incorporated classical instruments into some arrangements, including a string quartet on the pop ballad "Yesterday".[138] Composed by and sung by McCartney – none of the other Beatles perform on the recording[139] – "Yesterday" inspired the most cover versions of any song ever written.[140] With Help!, the Beatles became the first rock group to be nominated for a Grammy Award for Album of the Year.[141]
+The group's third US tour opened with a performance before a world-record crowd of 55,600 at New York's Shea Stadium on 15 August 1965 – "perhaps the most famous of all Beatles' concerts", in Lewisohn's description.[142] A further nine successful concerts followed in other American cities. At a show in Atlanta, the Beatles gave one of the first live performances ever to make use of a foldback system of on-stage monitor speakers.[143] Towards the end of the tour, they met with Elvis Presley, a foundational musical influence on the band, who invited them to his home in Beverly Hills.[144][145] September saw the launch of an American Saturday-morning cartoon series, The Beatles, that echoed A Hard Day's Night's slapstick antics over its two-year original run.[146] The series was a historical milestone as the first weekly television series to feature animated versions of real, living people.[147]
+In mid-October 1965, the Beatles entered the recording studio; for the first time when making an album, they had an extended period without other major commitments.[148] Until this time, according to George Martin, "we had been making albums rather like a collection of singles. Now we were really beginning to think about albums as a bit of art on their own."[149] Released in December, Rubber Soul was hailed by critics as a major step forward in the maturity and complexity of the band's music.[150] Their thematic reach was beginning to expand as they embraced deeper aspects of romance and philosophy.[151] NEMS executive Peter Brown attributed the new musical direction to "the Beatles' now habitual use of marijuana",[152] an assertion confirmed by the band – Lennon referred to it as "the pot album",[153] and Starr said: "Grass was really influential in a lot of our changes, especially with the writers. And because they were writing different material, we were playing differently."[153] After Help!'s foray into the world of classical music with flutes and strings, Harrison's introduction of a sitar on "Norwegian Wood (This Bird Has Flown)" marked a further progression outside the traditional boundaries of popular music. As their lyrics grew more artful, fans began to study them for deeper meaning. Of "Norwegian Wood" Lennon commented: "I was trying to be sophisticated in writing about an affair ... but in such a smokescreen way that you couldn't tell."[154]
+While many of Rubber Soul's prominent songs were the product of Lennon and McCartney's collaborative songwriting,[155] it also featured distinct compositions from each,[156] though they continued to share official credit. The song "In My Life", of which each later claimed lead authorship, is considered a highlight of the entire Lennon–McCartney catalogue.[157] Harrison called Rubber Soul his "favourite album"[153] and Starr referred to it as "the departure record".[158] McCartney has said, "We'd had our cute period, and now it was time to expand."[159] However, recording engineer Norman Smith later stated that the studio sessions revealed signs of growing conflict within the group – "the clash between John and Paul was becoming obvious", he wrote, and "as far as Paul was concerned, George could do no right".[160] In 2003, Rolling Stone ranked Rubber Soul fifth among "The 500 Greatest Albums of All Time",[161] and AllMusic's Richie Unterberger describes it as "one of the classic folk-rock records".[162]
+Capitol Records, from December 1963 when it began issuing Beatles recordings for the US market, exercised complete control over format,[82] compiling distinct US albums from the band's recordings and issuing songs of their choosing as singles.[163][nb 6] In June 1966, Yesterday and Today, one of Capitol's compilation albums, caused an uproar with its cover, which portrayed the grinning Beatles dressed in butcher's overalls, accompanied by raw meat and mutilated plastic baby dolls. According to Beatles biographer Bill Harry, it has been incorrectly suggested that this was meant as a satirical response to the way Capitol had "butchered" the US versions of the band's albums.[165] Thousands of copies of the LP had a new cover pasted over the original; an unpeeled "first-state" copy fetched $10,500 at a December 2005 auction.[166] In England, meanwhile, Harrison met sitar maestro Ravi Shankar, who agreed to train him on the instrument.[167]
+During a tour of the Philippines the month after the Yesterday and Today furore, the Beatles unintentionally snubbed the nation's first lady, Imelda Marcos, who had expected them to attend a breakfast reception at the Presidential Palace.[168] When presented with the invitation, Epstein politely declined on the band members' behalf, as it had never been his policy to accept such official invitations.[169] They soon found that the Marcos regime was unaccustomed to taking no for an answer. The resulting riots endangered the group and they escaped the country with difficulty.[170] Immediately afterwards, the band members visited India for the first time.[171]
+– John Lennon, 1966[172]
+Almost as soon as they returned home, the Beatles faced a fierce backlash from US religious and social conservatives (as well as the Ku Klux Klan) over a comment Lennon had made in a March interview with British reporter Maureen Cleave.[173] "Christianity will go", Lennon had said. "It will vanish and shrink. I needn't argue about that; I'm right and I will be proved right ... Jesus was alright but his disciples were thick and ordinary. It's them twisting it that ruins it for me."[174] His comments went virtually unnoticed in England, but when US teenage fan magazine Datebook printed them five months later, it sparked a controversy with Christians in America's conservative Bible Belt region.[173] The Vatican issued a protest, and bans on Beatles' records were imposed by Spanish and Dutch stations and South Africa's national broadcasting service.[175] Epstein accused Datebook of having taken Lennon's words out of context. At a press conference Lennon pointed out, "If I'd said television was more popular than Jesus, I might have got away with it."[176] He claimed that he was referring to how other people viewed their success, but at the prompting of reporters, he concluded: "If you want me to apologise, if that will make you happy, then okay, I'm sorry."[176]
+Released in August, a week before the Beatles' final tour, Revolver marked another artistic step forward for the group.[177] The album featured sophisticated songwriting, studio experimentation, and a greatly expanded repertoire of musical styles, ranging from innovative classical string arrangements to psychedelic rock.[177] Abandoning the customary group photograph, its Aubrey Beardsley-inspired cover – designed by Klaus Voormann, a friend of the band since their Hamburg days – was a monochrome collage and line drawing caricature of the group.[177] The album was preceded by the single "Paperback Writer", backed by "Rain".[178] Short promotional films were made for both songs; described by cultural historian Saul Austerlitz as "among the first true music videos",[179] they aired on The Ed Sullivan Show and Top of the Pops in June.[180]
+Among the experimental songs that Revolver featured was "Tomorrow Never Knows", the lyrics for which Lennon drew from Timothy Leary's The Psychedelic Experience: A Manual Based on the Tibetan Book of the Dead. Its creation involved eight tape decks distributed about the EMI building, each staffed by an engineer or band member, who randomly varied the movement of a tape loop while Martin created a composite recording by sampling the incoming data.[181] McCartney's "Eleanor Rigby" made prominent use of a string octet; Gould describes it as "a true hybrid, conforming to no recognisable style or genre of song".[182] Harrison was developing as a songwriter, and three of his compositions earned a place on the record.[183] In 2003, Rolling Stone ranked Revolver as the third greatest album of all time.[161]
+As preparations were made for a tour of the US, the Beatles knew that their music would hardly be heard. Having originally used Vox AC30 amplifiers, they later acquired more powerful 100-watt amplifiers, specially designed by Vox for them as they moved into larger venues in 1964, but these were still inadequate. Struggling to compete with the volume of sound generated by screaming fans, the band had grown increasingly bored with the routine of performing live.[184] Recognising that their shows were no longer about the music, they decided to make the August tour their last.[185]
+The band performed none of their new songs on the tour.[186] In Chris Ingham's description, they were very much "studio creations ... and there was no way a four-piece rock 'n' roll group could do them justice, particularly through the desensitising wall of the fans' screams. 'Live Beatles' and 'Studio Beatles' had become entirely different beasts."[187] The band's concert at San Francisco's Candlestick Park on 29 August was their last commercial concert.[188] It marked the end of four years dominated by almost nonstop touring that included over 1,400 concert appearances internationally.[189]
+Freed from the burden of touring, the Beatles embraced an increasingly experimental approach as they recorded Sgt. Pepper's Lonely Hearts Club Band, beginning in late November 1966.[191] According to engineer Geoff Emerick, the album's recording took over 700 hours.[192] He recalled the band's insistence "that everything on Sgt. Pepper had to be different. We had microphones right down in the bells of brass instruments and headphones turned into microphones attached to violins. We used giant primitive oscillators to vary the speed of instruments and vocals and we had tapes chopped to pieces and stuck together upside down and the wrong way around."[193] Parts of "A Day in the Life" featured a 40-piece orchestra.[193] The sessions initially yielded the non-album double A-side single "Strawberry Fields Forever"/"Penny Lane" in February 1967;[194] the Sgt. Pepper LP followed with a rush-release in May.[195] The musical complexity of the records, created using relatively primitive four-track recording technology, astounded contemporary artists.[190]  Among music critics, acclaim for the album was virtually universal.[196] Gould writes:
+The overwhelming consensus is that the Beatles had created a popular masterpiece: a rich, sustained, and overflowing work of collaborative genius whose bold ambition and startling originality dramatically enlarged the possibilities and raised the expectations of what the experience of listening to popular music on record could be. On the basis of this perception, Sgt. Pepper became the catalyst for an explosion of mass enthusiasm for album-formatted rock that would revolutionise both the aesthetics and the economics of the record business in ways that far outstripped the earlier pop explosions triggered by the Elvis phenomenon of 1956 and the Beatlemania phenomenon of 1963.[197]
+In the wake of Sgt. Pepper, the underground and mainstream press widely publicised the Beatles as leaders of youth culture, as well as "lifestyle revolutionaries".[3] The album was the first major pop/rock LP to include its complete lyrics, which appeared on the back cover.[198][199] Those lyrics were the subject of critical analysis; for instance, in late 1967 the album was the subject of a scholarly inquiry by American literary critic and professor of English Richard Poirier, who observed that his students were "listening to the group's music with a degree of engagement that he, as a teacher of literature, could only envy".[200][nb 7] The elaborate cover also attracted considerable interest and study.[201] A collage designed by pop artists Peter Blake and Jann Haworth, it depicted the group as the fictional band referred to in the album's title track[202] standing in front of a crowd of famous people.[203] The heavy moustaches worn by the group reflected the growing influence of hippie style,[204] while cultural historian Jonathan Harris describes their "brightly coloured parodies of military uniforms" as a knowingly "anti-authoritarian and anti-establishment" display.[205]
+Sgt. Pepper topped the UK charts for 23 consecutive weeks, with a further four weeks at number one in the period through to February 1968.[206] With 2.5 million copies sold within three months of its release,[207] Sgt. Pepper's initial commercial success exceeded that of all previous Beatles albums.[208] It sustained its immense popularity into the 21st century while breaking numerous sales records.[209]  In 2003, Rolling Stone ranked Sgt. Pepper foremost on its list of the greatest albums of all time.[161]
+Two Beatles film projects were conceived within weeks of completing Sgt. Pepper: Magical Mystery Tour, a one-hour television film, and Yellow Submarine, an animated feature-length film produced by United Artists.[210] The group began recording music for the former in late April 1967, but the project then lay dormant as they focused on recording songs for the latter.[211] On 25 June, the Beatles performed their forthcoming single "All You Need Is Love" to an estimated 350 million viewers on Our World, the first live global television link.[212] Released a week later, during the Summer of Love, the song was adopted as a flower power anthem.[213] The Beatles' use of psychedelic drugs was at its height during that summer.[214] In July and August, the group pursued interests related to similar utopian-based ideology, including a week-long investigation into the possibility of starting an island-based commune off the coast of Greece.[215]
+On 24 August, the group were introduced to Maharishi Mahesh Yogi in London. The next day, they travelled to Bangor for his Transcendental Meditation retreat. On 27 August, their manager's assistant, Peter Brown, phoned to inform them that Epstein had died.[216] The coroner ruled the death an accidental carbitol overdose, although it was widely rumoured to be a suicide.[217][nb 8] His death left the group disoriented and fearful about the future.[219] Lennon recalled: "We collapsed. I knew that we were in trouble then. I didn't really have any misconceptions about our ability to do anything other than play music, and I was scared. I thought, 'We've fuckin' had it now.'"[220] Harrison's then-wife Pattie Boyd remembered that "Paul and George were in complete shock. I don't think it could have been worse if they had heard that their own fathers had dropped dead."[221] During a band meeting in September, McCartney recommended that the band proceed with Magical Mystery Tour.[211]
+The Magical Mystery Tour soundtrack was released in the UK as a six-track double extended play (EP) in early December 1967.[82][222] It was the first example of a double EP in the UK.[223][224] The record carried on the psychedelic vein of Sgt. Pepper,[225] however, in line with the band's wishes, the packaging reinforced the idea that the release was a film soundtrack rather than a follow-up to Sgt. Pepper.[222]  In the US, the soundtrack appeared as an identically titled LP that also included five tracks from the band's recent singles.[104] In its first three weeks, the album set a record for the highest initial sales of any Capitol LP, and it is the only Capitol compilation later to be adopted in the band's official canon of studio albums.[226]
+Magical Mystery Tour first aired on Boxing Day to an audience of approximately 15 million.[227] Largely directed by McCartney, the film was the band's first critical failure in the UK.[228] It was dismissed as "blatant rubbish" by the Daily Express; the Daily Mail called it "a colossal conceit"; and The Guardian labelled the film "a kind of fantasy morality play about the grossness and warmth and stupidity of the audience".[229] Gould describes it as "a great deal of raw footage showing a group of people getting on, getting off, and riding on a bus".[229] Although the viewership figures were respectable, its slating in the press led US television networks to lose interest in broadcasting the film.[230]
+The group were less involved with Yellow Submarine, which only featured the band appearing as themselves for a short live-action segment.[231] Premiering in July 1968, the film featured cartoon versions of the band members and a soundtrack with eleven of their songs, including four unreleased studio recordings that made their debut in the film.[232] Critics praised the film for its music, humour and innovative visual style.[233] A soundtrack LP was issued seven months later; it contained those four new songs, the title track (already issued on Revolver), "All You Need Is Love" (already issued as a single and on the US Magical Mystery Tour LP) and seven instrumental pieces composed by Martin.[234]
+In February 1968, the Beatles travelled to Maharishi Mahesh Yogi's ashram in Rishikesh, India, to take part in a three-month meditation "Guide Course". Their time in India marked one of the band's most prolific periods, yielding numerous songs, including a majority of those on their next album.[235] However, Starr left after only ten days, unable to stomach the food, and McCartney eventually grew bored and departed a month later.[236] For Lennon and Harrison, creativity turned to question when an electronics technician known as Magic Alex suggested that the Maharishi was attempting to manipulate them.[237] When he alleged that the Maharishi had made sexual advances to women attendees, a persuaded Lennon left abruptly just two months into the course, bringing an unconvinced Harrison and the remainder of the group's entourage with him.[236] In anger, Lennon wrote a scathing song titled "Maharishi", renamed "Sexy Sadie" to avoid potential legal issues. McCartney said, "We made a mistake. We thought there was more to him than there was."[237]
+In May, Lennon and McCartney traveled to New York for the public unveiling of the Beatles' new business venture, Apple Corps.[238] It was initially formed several months earlier as part of a plan to create a tax-effective business structure, but the band then desired to extend the corporation to other pursuits, including record distribution, peace activism, and education.[239] McCartney described Apple as "rather like a Western communism".[240] The enterprise drained the group financially with a series of unsuccessful projects[241] handled largely by members of the Beatles' entourage, who were given their jobs regardless of talent and experience.[242] Among its numerous subsidiaries were Apple Electronics, established to foster technological innovations with Magic Alex at the head, and Apple Retailing, which opened the short-lived Apple Boutique in London.[243] Harrison later said, "Basically, it was chaos ... John and Paul got carried away with the idea and blew millions, and Ringo and I just had to go along with it."[240]
+From late May to mid-October 1968, the group recorded what became The Beatles, a double LP commonly known as "the White Album" for its virtually featureless cover.[245] During this time, relations between the members grew openly divisive.[246] Starr quit for two weeks, leaving his bandmates to record "Back in the U.S.S.R." and "Dear Prudence" as a trio.[247] Lennon had lost interest in collaborating with McCartney,[248] whose contribution "Ob-La-Di, Ob-La-Da" he scorned as "granny music shit".[249] Tensions were further aggravated by Lennon's romantic preoccupation with avant-garde artist Yoko Ono, whom he insisted on bringing to the sessions despite the group's well-established understanding that girlfriends were not allowed in the studio.[250] McCartney has recalled that the album "wasn't a pleasant one to make".[251] He and Lennon identified the sessions as the start of the band's break-up.[252][253]
+With the record, the band executed a wider range of musical styles[254] and broke with their recent tradition of incorporating several musical styles in one song by keeping each piece of music consistently faithful to a select genre.[255] During the sessions, the group upgraded to an eight-track tape console, which made it easier for them to layer tracks piecemeal, while the members often recorded independently of each other, affording the album a reputation as a collection of solo recordings rather than a unified group effort.[256] Describing the double album, Lennon later said: "Every track is an individual track; there isn't any Beatle music on it. [It's] John and the band, Paul and the band, George and the band."[257] The sessions also produced the Beatles' longest song yet, "Hey Jude", released in August as a non-album single with "Revolution".[258]
+Issued in November, the White Album was the band's first Apple Records album release, although EMI continued to own their recordings.[259] The record attracted more than 2 million advance orders, selling nearly 4 million copies in the US in little over a month, and its tracks dominated the playlists of American radio stations.[260] Its lyric content was the focus of much analysis by the counterculture.[261] Despite its popularity, reviewers were largely confused by the album's content, and it failed to inspire the level of critical writing that Sgt. Pepper had.[260] General critical opinion eventually turned in favour of the White Album, and in 2003, Rolling Stone ranked it as the tenth greatest album of all time.[161]
+Although Let It Be was the Beatles' final album release, it was largely recorded before Abbey Road. The project's impetus came from an idea Martin attributes to McCartney, who suggested they "record an album of new material and rehearse it, then perform it before a live audience for the very first time – on record and on film".[262] Originally intended for a one-hour television programme to be called Beatles at Work, in the event much of the album's content came from studio work beginning in January 1969, many hours of which were captured on film by director Michael Lindsay-Hogg.[262][263] Martin said that the project was "not at all a happy recording experience. It was a time when relations between the Beatles were at their lowest ebb."[262] Lennon described the largely impromptu sessions as "hell ... the most miserable ... on Earth", and Harrison, "the low of all-time".[264] Irritated by McCartney and Lennon, Harrison walked out for five days. Upon returning, he threatened to leave the band unless they "abandon[ed] all talk of live performance" and instead focused on finishing a new album, initially titled Get Back, using songs recorded for the TV special.[265] He also demanded they cease work at Twickenham Film Studios, where the sessions had begun, and relocate to the newly finished Apple Studio. His bandmates agreed, and it was decided to salvage the footage shot for the TV production for use in a feature film.[266]
+To alleviate tensions within the band and improve the quality of their live sound, Harrison invited keyboardist Billy Preston to participate in the last nine days of sessions.[267] Preston received label billing on the "Get Back" single – the only musician ever to receive that acknowledgment on an official Beatles release.[268] After the rehearsals, the band could not agree on a location to film a concert, rejecting several ideas, including a boat at sea, a lunatic asylum, the Tunisian desert, and the Colosseum.[262] Ultimately, what would be their final live performance was filmed on the rooftop of the Apple Corps building at 3 Savile Row, London, on 30 January 1969.[269] Five weeks later, engineer Glyn Johns, whom Lewisohn describes as Get Back's "uncredited producer", began work assembling an album, given "free rein" as the band "all but washed their hands of the entire project".[270]
+New strains developed between the band members regarding the appointment of a financial adviser, the need for which had become evident without Epstein to manage business affairs. Lennon, Harrison and Starr favoured Allen Klein, who had managed the Rolling Stones and Sam Cooke;[271] McCartney wanted Lee and John Eastman – father and brother, respectively, of Linda Eastman,[272] whom McCartney married on 12 March.[273] Agreement could not be reached, so both Klein and the Eastmans were temporarily appointed: Klein as the Beatles' business manager and the Eastmans as their lawyers.[274][275] Further conflict ensued, however, and financial opportunities were lost.[271] On 8 May, Klein was named sole manager of the band,[276] the Eastmans having previously been dismissed as the Beatles' lawyers. McCartney refused to sign the management contract with Klein, but he was out-voted by the other Beatles.[277]
+Martin stated that he was surprised when McCartney asked him to produce another album, as the Get Back sessions had been "a miserable experience" and he had "thought it was the end of the road for all of us".[278] The primary recording sessions for Abbey Road began on 2 July.[279] Lennon, who rejected Martin's proposed format of a "continuously moving piece of music", wanted his and McCartney's songs to occupy separate sides of the album.[280] The eventual format, with individually composed songs on the first side and the second consisting largely of a medley, was McCartney's suggested compromise.[280] Emerick noted that the replacement of the studio's valve mixing console with a transistorised one yielded a less punchy sound, leaving the group frustrated at the thinner tone and lack of impact and contributing to its "kinder, gentler" feel relative to their previous albums.[281]
+On 4 July, the first solo single by a Beatle was released: Lennon's "Give Peace a Chance", credited to the Plastic Ono Band. The completion and mixing of "I Want You (She's So Heavy)" on 20 August was the last occasion on which all four Beatles were together in the same studio.[282] On 8 September, while Starr was in hospital, the other band members met to discuss recording a new album. They considered a different approach to songwriting by ending the Lennon–McCartney pretense and having four compositions apiece from Lennon, McCartney and Harrison, with two from Starr and a lead single around Christmas.[283] On 20 September, Lennon announced his departure to the rest of the group but agreed to withhold a public announcement to avoid undermining sales of the forthcoming album.[284]
+Released on 26 September, Abbey Road sold four million copies within three months and topped the UK charts for a total of seventeen weeks.[285] Its second track, the ballad "Something", was issued as a single – the only Harrison composition that appeared as a Beatles A-side.[286] Abbey Road received mixed reviews, although the medley met with general acclaim.[285] Unterberger considers it "a fitting swan song for the group", containing "some of the greatest harmonies to be heard on any rock record".[287] Musicologist and author Ian MacDonald calls the album "erratic and often hollow", despite the "semblance of unity and coherence" offered by the medley.[288] Martin singled it out as his favourite Beatles album; Lennon said it was "competent" but had "no life in it".[281]
+For the still unfinished Get Back album, one last song, Harrison's "I Me Mine", was recorded on 3 January 1970. Lennon, in Denmark at the time, did not participate.[289] In March, rejecting the work Johns had done on the project, now retitled Let It Be, Klein gave the session tapes to American producer Phil Spector, who had recently produced Lennon's solo single "Instant Karma!"[290] In addition to remixing the material, Spector edited, spliced and overdubbed several of the recordings that had been intended as "live". McCartney was unhappy with the producer's approach and particularly dissatisfied with the lavish orchestration on "The Long and Winding Road", which involved a fourteen-voice choir and 36-piece instrumental ensemble.[291] McCartney's demands that the alterations to the song be reverted were ignored,[292] and he publicly announced his departure from the band on 10 April, a week before the release of his first, self-titled solo album.[291][293]
+On 8 May 1970, Let It Be was released. Its accompanying single, "The Long and Winding Road", was the Beatles' last; it was released in the US, but not in the UK.[178] The Let It Be documentary film followed later that month, and would win the 1970 Academy Award for Best Original Song Score.[294] Sunday Telegraph critic Penelope Gilliatt called it "a very bad film and a touching one ... about the breaking apart of this reassuring, geometrically perfect, once apparently ageless family of siblings".[295] Several reviewers stated that some of the performances in the film sounded better than their analogous album tracks.[296] Describing Let It Be as the "only Beatles album to occasion negative, even hostile reviews", Unterberger calls it "on the whole underrated"; he singles out "some good moments of straight hard rock in 'I've Got a Feeling' and 'Dig a Pony'", and praises "Let It Be", "Get Back", and "the folky 'Two of Us', with John and Paul harmonising together".[297]
+McCartney filed suit for the dissolution of the Beatles' contractual partnership on 31 December 1970.[298] Legal disputes continued long after their break-up, and the dissolution was not formalised until 29 December 1974,[299] when Lennon signed the paperwork terminating the partnership while on vacation with his family at Walt Disney World Resort in Florida.[300]
+Lennon, McCartney, Harrison and Starr all released solo albums in 1970. Their solo records sometimes involved one or more of the others;[301] Starr's Ringo (1973) was the only album to include compositions and performances by all four ex-Beatles, albeit on separate songs. With Starr's participation, Harrison staged the Concert for Bangladesh in New York City in August 1971.[302] Other than an unreleased jam session in 1974, later bootlegged as A Toot and a Snore in '74, Lennon and McCartney never recorded together again.[303]
+Two double-LP sets of the Beatles' greatest hits, compiled by Klein, 1962–1966 and 1967–1970, were released in 1973, at first under the Apple Records imprint.[304] Commonly known as the "Red Album" and "Blue Album", respectively, each has earned a Multi-Platinum certification in the US and a Platinum certification in the UK.[305][306] Between 1976 and 1982, EMI/Capitol released a wave of compilation albums without input from the ex-Beatles, starting with the double-disc compilation Rock 'n' Roll Music.[307] The only one to feature previously unreleased material was The Beatles at the Hollywood Bowl (1977); the first officially issued concert recordings by the group, it contained selections from two shows they played during their 1964 and 1965 US tours.[308][nb 9]
+The music and enduring fame of the Beatles were commercially exploited in various other ways, again often outside their creative control. In April 1974, the musical John, Paul, George, Ringo ... and Bert, written by Willy Russell and featuring singer Barbara Dickson, opened in London. It included, with permission from Northern Songs, eleven Lennon-McCartney compositions and one by Harrison, "Here Comes the Sun". Displeased with the production's use of his song, Harrison withdrew his permission to use it.[310] Later that year, the off-Broadway musical Sgt. Pepper's Lonely Hearts Club Band on the Road opened.[311] All This and World War II (1976) was an unorthodox nonfiction film that combined newsreel footage with covers of Beatles songs by performers ranging from Elton John and Keith Moon to the London Symphony Orchestra.[312] The Broadway musical Beatlemania, an unauthorised nostalgia revue, opened in early 1977 and proved popular, spinning off five separate touring productions.[313] In 1979, the band sued the producers, settling for several million dollars in damages.[313] Sgt. Pepper's Lonely Hearts Club Band (1978), a musical film starring the Bee Gees and Peter Frampton, was a commercial failure and an "artistic fiasco", according to Ingham.[314]
+Accompanying the wave of Beatles nostalgia and persistent reunion rumours in the US during the 1970s, several entrepreneurs made public offers to the Beatles for a reunion concert.[315] Promoter Bill Sargent first offered the Beatles $10 million for a reunion concert in 1974. He raised his offer to $30 million in January 1976 and then to $50 million the following month.[316][317] On 24 April 1976, during a broadcast of Saturday Night Live, producer Lorne Michaels offered the Beatles $3,000 to reunite on the show. Lennon and McCartney were watching the live broadcast at Lennon's apartment at the Dakota in New York, which was within driving distance of the NBC studio where the show was being broadcast. The former bandmates briefly entertained the idea of going to the studio and surprising Michaels by accepting his offer, but decided not to.[318]
+In December 1980, Lennon was shot and killed outside his New York City apartment. Harrison rewrote the lyrics of his song "All Those Years Ago" in Lennon's honour. With Starr on drums and McCartney and his wife, Linda, contributing backing vocals, the song was released as a single in May 1981.[319] McCartney's own tribute, "Here Today", appeared on his Tug of War album in April 1982.[320] In 1987, Harrison's Cloud Nine album included "When We Was Fab", a song about the Beatlemania era.[321]
+When the Beatles' studio albums were released on CD by EMI and Apple Corps in 1987, their catalogue was standardised throughout the world, establishing a canon of the twelve original studio LPs as issued in the UK plus the US LP version of Magical Mystery Tour.[322] All the remaining material from the singles and EPs that had not appeared on these thirteen studio albums was gathered on the two-volume compilation Past Masters (1988). Except for the Red and Blue albums, EMI deleted all its other Beatles compilations – including the Hollywood Bowl record – from its catalogue.[308]
+In 1988, the Beatles were inducted into the Rock and Roll Hall of Fame, their first year of eligibility. Harrison and Starr attended the ceremony with Lennon's widow, Yoko Ono, and his two sons, Julian and Sean.[323][324] McCartney declined to attend, citing unresolved "business differences" that would make him "feel like a complete hypocrite waving and smiling with them at a fake reunion".[324] The following year, EMI/Capitol settled a decade-long lawsuit filed by the band over royalties, clearing the way to commercially package previously unreleased material.[325][326]
+Live at the BBC, the first official release of unissued Beatles performances in seventeen years, appeared in 1994.[327] That same year McCartney, Harrison and Starr collaborated on the Anthology project. Anthology was the culmination of work begun in 1970, when Apple Corps director Neil Aspinall, their former road manager and personal assistant, had started to gather material for a documentary with the working title The Long and Winding Road.[328] Documenting their history in the band's own words, the Anthology project included the release of several unissued Beatles recordings. McCartney, Harrison and Starr also added new instrumental and vocal parts to songs recorded as demos by Lennon in the late 1970s.[329]
+During 1995–96, the project yielded a television miniseries, an eight-volume video set, and three two-CD/three-LP box sets featuring artwork by Klaus Voormann. Two songs based on Lennon demos, "Free as a Bird" and "Real Love", were issued as new Beatles singles. The releases were commercially successful and the television series was viewed by an estimated 400 million people.[330] In 1999, to coincide with the re-release of the 1968 film Yellow Submarine, an expanded soundtrack album, Yellow Submarine Songtrack, was issued.[331]
+The Beatles' 1, a compilation album of the band's British and American number-one hits, was released on 13 November 2000. It became the fastest-selling album of all time, with 3.6 million sold in its first week[332] and 13 million within a month.[333] It topped albums charts in at least 28 countries.[334] The compilation had sold 31 million copies globally by April 2009.[335]
+Harrison died from metastatic lung cancer in November 2001.[336][337][338] McCartney and Starr were among the musicians who performed at the Concert for George, organised by Eric Clapton and Harrison's widow, Olivia. The tribute event took place at the Royal Albert Hall on the first anniversary of Harrison's death.[339]
+In 2003, Let It Be... Naked, a reconceived version of the Let It Be album, with McCartney supervising production, was released. One of the main differences from the Spector-produced version was the omission of the original string arrangements.[340] It was a top ten hit in both Britain and America. The US album configurations from 1964 to 1965 were released as box sets in 2004 and 2006; The Capitol Albums, Volume 1 and Volume 2 included both stereo and mono versions based on the mixes that were prepared for vinyl at the time of the music's original American release.[341]
+As a soundtrack for Cirque du Soleil's Las Vegas Beatles stage revue, Love, George Martin and his son Giles remixed and blended 130 of the band's recordings to create what Martin called "a way of re-living the whole Beatles musical lifespan in a very condensed period".[342] The show premiered in June 2006, and the Love album was released that November.[343] In April 2009, Starr performed three songs with McCartney at a benefit concert held at New York's Radio City Music Hall and organised by McCartney.[344]
+On 9 September 2009, the Beatles' entire back catalogue was reissued following an extensive digital remastering process that lasted four years.[322] Stereo editions of all twelve original UK studio albums, along with Magical Mystery Tour and the Past Masters compilation, were released on compact disc both individually and as a box set.[345] A second collection, The Beatles in Mono, included remastered versions of every Beatles album released in true mono along with the original 1965 stereo mixes of Help! and Rubber Soul (both of which Martin had remixed for the 1987 editions).[346] The Beatles: Rock Band, a music video game in the Rock Band series, was issued on the same day.[347] In December 2009, the band's catalogue was officially released in FLAC and MP3 format in a limited edition of 30,000 USB flash drives.[348]
+Owing to a long-running royalty disagreement, the Beatles were among the last major artists to sign deals with online music services.[349] Residual disagreement emanating from Apple Corps' dispute with Apple, Inc., iTunes' owners, over the use of the name "Apple" was also partly responsible for the delay, although in 2008, McCartney stated that the main obstacle to making the Beatles' catalogue available online was that EMI "want[s] something we're not prepared to give them".[350] In 2010, the official canon of thirteen Beatles studio albums, Past Masters, and the "Red" and "Blue" greatest-hits albums were made available on iTunes.[351]
+In 2012, EMI's recorded music operations were sold to Universal Music Group. In order for Universal Music to acquire EMI, the European Union, for antitrust reasons, forced EMI to spin off assets including Parlophone. Universal was allowed to keep the Beatles' recorded music catalogue, managed by Capitol Records under its Capitol Music Group division.[352] The entire original Beatles album catalogue was also reissued on vinyl in 2012; available either individually or as a box set.[353]
+In 2013, a second volume of BBC recordings, titled On Air – Live at the BBC Volume 2, was released. That December saw the release of another 59 Beatles recordings on iTunes. The set, titled The Beatles Bootleg Recordings 1963, had the opportunity to gain a 70-year copyright extension conditional on the songs being published at least once before the end of 2013. Apple Records released the recordings on 17 December to prevent them from going into the public domain and had them taken down from iTunes later that same day. Fan reactions to the release were mixed, with one blogger saying "the hardcore Beatles collectors who are trying to obtain everything will already have these."[354][355]
+On 26 January 2014, McCartney and Starr performed together at the 56th Annual Grammy Awards, held at the Staples Center in Los Angeles.[356] The following day, The Night That Changed America: A Grammy Salute to The Beatles television special was taped in the Los Angeles Convention Center's West Hall. It aired on 9 February, the exact date of – and at the same time, and on the same network as – the original broadcast of the Beatles' first US television appearance on The Ed Sullivan Show, 50 years earlier. The special included performances of Beatles songs by current artists as well as by McCartney and Starr, archival footage, and interviews with the two surviving ex-Beatles carried out by David Letterman at the Ed Sullivan Theater.[357][358] In December 2015, the Beatles released their catalogue for streaming on various streaming music services including Spotify and Apple Music.[359]
+In September 2016, the documentary film The Beatles: Eight Days a Week was released. Directed by Ron Howard, it chronicled the Beatles' career during their touring years from 1962 to 1966, from their performances in Liverpool's the Cavern Club in 1961 to their final concert in San Francisco in 1966. The film was released theatrically on 15 September in the UK and the US, and started streaming on Hulu on 17 September. It received several awards and nominations, including for Best Documentary at the 70th British Academy Film Awards and the Outstanding Documentary or Nonfiction Special at the 69th Primetime Creative Arts Emmy Awards.[360] An expanded, remixed and remastered version of The Beatles at the Hollywood Bowl was released on 9 September, to coincide with the release of the film.[361][362]
+On 18 May 2017, Sirius XM Radio launched a 24/7 radio channel, The Beatles Channel. A week later, Sgt. Pepper's Lonely Hearts Club Band was reissued with new stereo mixes and unreleased material for the album's 50th anniversary.[363] Similar box sets were released for The Beatles in November 2018,[364] and Abbey Road in September 2019.[365] On the first week of October 2019, Abbey Road returned to number one on the UK Albums Chart. The Beatles broke their own record for the album with the longest gap between topping the charts as Abbey Road hit the top spot 50 years after its original release.[366]
+In August 2021, The Beatles: Get Back, a new documentary film directed by Peter Jackson utilising footage captured for what became the Let It Be film, will be released by Walt Disney Studios Motion Pictures in the US and Canada, with a global release to follow.[367]
+In Icons of Rock: An Encyclopedia of the Legends Who Changed Music Forever, Scott Schinder and Andy Schwartz describe the Beatles' musical evolution:
+In their initial incarnation as cheerful, wisecracking moptops, the Fab Four revolutionised the sound, style, and attitude of popular music and opened rock and roll's doors to a tidal wave of British rock acts. Their initial impact would have been enough to establish the Beatles as one of their era's most influential cultural forces, but they didn't stop there. Although their initial style was a highly original, irresistibly catchy synthesis of early American rock and roll and R&B, the Beatles spent the rest of the 1960s expanding rock's stylistic frontiers, consistently staking out new musical territory on each release. The band's increasingly sophisticated experimentation encompassed a variety of genres, including folk-rock, country, psychedelia, and baroque pop, without sacrificing the effortless mass appeal of their early work.[368]
+In The Beatles as Musicians, Walter Everett describes Lennon and McCartney's contrasting motivations and approaches to composition: "McCartney may be said to have constantly developed – as a means to entertain – a focused musical talent with an ear for counterpoint and other aspects of craft in the demonstration of a universally agreed-upon common language that he did much to enrich. Conversely, Lennon's mature music is best appreciated as the daring product of a largely unconscious, searching but undisciplined artistic sensibility."[369]
+Ian MacDonald describes McCartney as "a natural melodist – a creator of tunes capable of existing apart from their harmony". His melody lines are characterised as primarily "vertical", employing wide, consonant intervals which express his "extrovert energy and optimism". Conversely, Lennon's "sedentary, ironic personality" is reflected in a "horizontal" approach featuring minimal, dissonant intervals and repetitive melodies which rely on their harmonic accompaniment for interest: "Basically a realist, he instinctively kept his melodies close to the rhythms and cadences of speech, colouring his lyrics with bluesy tone and harmony rather than creating tunes that made striking shapes of their own."[370] MacDonald praises Harrison's lead guitar work for the role his "characterful lines and textural colourings" play in supporting Lennon and McCartney's parts, and describes Starr as "the father of modern pop/rock drumming".[371]
+The band's earliest influences include Elvis Presley, Carl Perkins, Little Richard and Chuck Berry.[372] During the Beatles' co-residency with Little Richard at the Star-Club in Hamburg, from April to May 1962, he advised them on the proper technique for performing his songs.[373] Of Presley, Lennon said, "Nothing really affected me until I heard Elvis. If there hadn't been Elvis, there would not have been the Beatles."[374] Other early influences include Buddy Holly, Eddie Cochran, Roy Orbison[375] and the Everly Brothers.[376]
+The Beatles continued to absorb influences long after their initial success, often finding new musical and lyrical avenues by listening to their contemporaries, including Bob Dylan, the Who, Frank Zappa, the Lovin' Spoonful, the Byrds and the Beach Boys, whose 1966 album Pet Sounds amazed and inspired McCartney.[377][378][379][380] Referring to the Beach Boys' creative leader, Martin later stated: "No one made a greater impact on the Beatles than Brian [Wilson]."[381] Ravi Shankar, with whom Harrison studied for six weeks in India in late 1966, had a significant effect on his musical development during the band's later years.[382]
+Originating as a skiffle group, the Beatles quickly embraced 1950s rock and roll and helped pioneer the Merseybeat genre,[383] and their repertoire ultimately expanded to include a broad variety of pop music.[384] Reflecting the range of styles they explored, Lennon said of Beatles for Sale, "You could call our new one a Beatles country-and-western LP",[385] while Gould credits Rubber Soul as "the instrument by which legions of folk-music enthusiasts were coaxed into the camp of pop".[386]
+Although the 1965 song "Yesterday" was not the first pop record to employ orchestral strings, it marked the group's first recorded use of classical music elements. Gould observes: "The more traditional sound of strings allowed for a fresh appreciation of their talent as composers by listeners who were otherwise allergic to the din of drums and electric guitars."[387] They continued to experiment with string arrangements to various effect; Sgt. Pepper's "She's Leaving Home", for instance, is "cast in the mold of a sentimental Victorian ballad", Gould writes, "its words and music filled with the clichés of musical melodrama".[388]
+The band's stylistic range expanded in another direction with their 1966 B-side "Rain", described by Martin Strong as "the first overtly psychedelic Beatles record".[389] Other psychedelic numbers followed, such as "Tomorrow Never Knows" (recorded before "Rain"), "Strawberry Fields Forever", "Lucy in the Sky with Diamonds" and "I Am the Walrus". The influence of Indian classical music was evident in Harrison's "The Inner Light", "Love You To" and "Within You Without You" – Gould describes the latter two as attempts "to replicate the raga form in miniature".[390]
+Innovation was the most striking feature of their creative evolution, according to music historian and pianist Michael Campbell: "'A Day in the Life' encapsulates the art and achievement of the Beatles as well as any single track can. It highlights key features of their music: the sound imagination, the persistence of tuneful melody, and the close coordination between words and music. It represents a new category of song – more sophisticated than pop ... and uniquely innovative. There literally had never before been a song – classical or vernacular – that had blended so many disparate elements so imaginatively."[391] Philosophy professor Bruce Ellis Benson agrees: "the Beatles ... give us a wonderful example of how such far-ranging influences as Celtic music, rhythm and blues, and country and western could be put together in a new way."[392]
+Author Dominic Pedler describes the way they crossed musical styles: "Far from moving sequentially from one genre to another (as is sometimes conveniently suggested) the group maintained in parallel their mastery of the traditional, catchy chart hit while simultaneously forging rock and dabbling with a wide range of peripheral influences from country to vaudeville. One of these threads was their take on folk music, which would form such essential groundwork for their later collisions with Indian music and philosophy."[393] As the personal relationships between the band members grew increasingly strained, their individual tastes became more apparent. The minimalistic cover artwork for the White Album contrasted with the complexity and diversity of its music, which encompassed Lennon's "Revolution 9" (whose musique concrète approach was influenced by Yoko Ono), Starr's country song "Don't Pass Me By", Harrison's rock ballad "While My Guitar Gently Weeps", and the "proto-metal roar" of McCartney's "Helter Skelter".[394]
+George Martin's close involvement in his role as producer made him one of the leading candidates for the informal title of the "fifth Beatle".[395] He applied his classical musical training in various ways, and functioned as "an informal music teacher" to the progressing songwriters, according to Gould.[396] Martin suggested to a sceptical McCartney that the arrangement of "Yesterday" should feature a string quartet accompaniment, thereby introducing the Beatles to a "hitherto unsuspected world of classical instrumental colour", in MacDonald's description.[397] Their creative development was also facilitated by Martin's willingness to experiment in response to their suggestions, such as adding "something baroque" to a particular recording.[398] In addition to scoring orchestral arrangements for recordings, Martin often performed on them, playing instruments including piano, organ and brass.[399]
+Collaborating with Lennon and McCartney required Martin to adapt to their different approaches to songwriting and recording. MacDonald comments, "while [he] worked more naturally with the conventionally articulate McCartney, the challenge of catering to Lennon's intuitive approach generally spurred him to his more original arrangements, of which 'Being for the Benefit of Mr. Kite!' is an outstanding example."[400] Martin said of the two composers' distinct songwriting styles and his stabilising influence:
+Compared with Paul's songs, all of which seemed to keep in some sort of touch with reality, John's had a psychedelic, almost mystical quality ... John's imagery is one of the best things about his work – 'tangerine trees', 'marmalade skies', 'cellophane flowers' ... I always saw him as an aural Salvador Dalí, rather than some drug-ridden record artist. On the other hand, I would be stupid to pretend that drugs didn't figure quite heavily in the Beatles' lives at that time ... they knew that I, in my schoolmasterly role, didn't approve ... Not only was I not into it myself, I couldn't see the need for it; and there's no doubt that, if I too had been on dope, Pepper would never have been the album it was. Perhaps it was the combination of dope and no dope that worked, who knows?[401]
+Harrison echoed Martin's description of his stabilising role: "I think we just grew through those years together, him as the straight man and us as the loonies; but he was always there for us to interpret our madness – we used to be slightly avant-garde on certain days of the week, and he would be there as the anchor person, to communicate that through the engineers and on to the tape."[402]
+Making innovative use of technology while expanding the possibilities of recorded music, the Beatles urged experimentation by Martin and his recording engineers. Seeking ways to put chance occurrences to creative use, accidental guitar feedback, a resonating glass bottle, a tape loaded the wrong way round so that it played backwards – any of these might be incorporated into their music.[403] Their desire to create new sounds on every new recording, combined with Martin's arranging abilities and the studio expertise of EMI staff engineers Norman Smith, Ken Townsend and Geoff Emerick, all contributed significantly to their records from Rubber Soul and, especially, Revolver onwards.[403]
+Along with innovative studio techniques such as sound effects, unconventional microphone placements, tape loops, double tracking and vari-speed recording, the Beatles augmented their songs with instruments that were unconventional in rock music at the time. These included string and brass ensembles as well as Indian instruments such as the sitar in "Norwegian Wood" and the swarmandal in "Strawberry Fields Forever".[404] They also used novel electronic instruments such as the Mellotron, with which McCartney supplied the flute voices on the "Strawberry Fields Forever" intro,[405] and the clavioline, an electronic keyboard that created the unusual oboe-like sound on "Baby, You're a Rich Man".[406]
+Former Rolling Stone associate editor Robert Greenfield compared the Beatles to Picasso, as "artists who broke through the constraints of their time period to come up with something that was unique and original ... [I]n the form of popular music, no one will ever be more revolutionary, more creative and more distinctive ..."[347] The British poet Philip Larkin described their work as "an enchanting and intoxicating hybrid of Negro rock-and-roll with their own adolescent romanticism", and "the first advance in popular music since the War".[408]
+They not only sparked the British Invasion of the US,[409] they became a globally influential phenomenon as well.[410] From the 1920s, the US had dominated popular entertainment culture throughout much of the world, via Hollywood films, jazz, the music of Broadway and Tin Pan Alley and, later, the rock and roll that first emerged in Memphis, Tennessee.[333] The Beatles are regarded as British cultural icons, with young adults from abroad naming the band among a group of people that they most associated with UK culture.[411][412]
+Their musical innovations and commercial success inspired musicians worldwide.[410] Many artists have acknowledged the Beatles' influence and enjoyed chart success with covers of their songs.[413] On radio, their arrival marked the beginning of a new era; in 1968 the programme director of New York's WABC radio station forbade his DJs from playing any "pre-Beatles" music, marking the defining line of what would be considered oldies on American radio.[414] They helped to redefine the album as something more than just a few hits padded out with "filler",[415] and they were primary innovators of the modern music video.[416] The Shea Stadium show with which they opened their 1965 North American tour attracted an estimated 55,600 people,[142] then the largest audience in concert history; Spitz describes the event as a "major breakthrough ... a giant step toward reshaping the concert business".[417] Emulation of their clothing and especially their hairstyles, which became a mark of rebellion, had a global impact on fashion.[100]
+According to Gould, the Beatles changed the way people listened to popular music and experienced its role in their lives. From what began as the Beatlemania fad, the group's popularity grew into what was seen as an embodiment of sociocultural movements of the decade. As icons of the 1960s counterculture, Gould continues, they became a catalyst for bohemianism and activism in various social and political arenas, fuelling movements such as women's liberation, gay liberation and environmentalism.[418] According to Peter Lavezzoli, after the "more popular than Jesus" controversy in 1966, the Beatles felt considerable pressure to say the right things and "began a concerted effort to spread a message of wisdom and higher consciousness".[167]
+Other commentators such as Mikal Gilmore and Todd Leopold have traced the inception of their socio-cultural impact earlier, interpreting even the Beatlemania period, particularly on their first visit to the US, as a key moment in the development of generational awareness.[98][419] Referring to their appearance on the Ed Sullivan Show Leopold states: "In many ways, the Sullivan appearance marked the beginning of a cultural revolution ... The Beatles were like aliens dropped into the United States of 1964."[419] According to Gilmore:
+Elvis Presley had shown us how rebellion could be fashioned into eye-opening style; the Beatles were showing us how style could have the impact of cultural revelation – or at least how a pop vision might be forged into an unimpeachable consensus.[98]
+Established in 2009, Global Beatles Day is an annual holiday on 25 June each year that honours and celebrates the ideals of the Beatles.[420] The date was chosen to commemorate the date the group participated in the BBC programme Our World in 1967, performing "All You Need Is Love" broadcast to an international audience.[421]
+In 1965, Queen Elizabeth II appointed Lennon, McCartney, Harrison and Starr Members of the Order of the British Empire (MBE).[131] The Beatles won the 1971 Academy Award for Best Original Song Score for the film Let It Be (1970).[294] The recipients of seven Grammy Awards[422] and fifteen Ivor Novello Awards,[423] the Beatles have six Diamond albums, as well as 20 Multi-Platinum albums, 16 Platinum albums and six Gold albums in the US.[305] In the UK, the Beatles have four Multi-Platinum albums, four Platinum albums, eight Gold albums and one Silver album.[306] They were inducted into the Rock and Roll Hall of Fame in 1988.[323]
+The best-selling band in history, the Beatles have sold more than 800 million physical and digital albums as of 2013[update].[424][nb 10] They have had more number-one albums on the UK charts, fifteen,[426] and sold more singles in the UK, 21.9 million, than any other act.[427] In 2004, Rolling Stone ranked the Beatles as the most significant and influential rock music artists of the last 50 years.[428] They ranked number one on Billboard magazine's list of the all-time most successful Hot 100 artists, released in 2008 to celebrate the US singles chart's 50th anniversary.[429] As of 2017[update], they hold the record for most number-one hits on the Billboard Hot 100, with twenty.[430] The Recording Industry Association of America certifies that the Beatles have sold 178 million units in the US, more than any other artist.[431] They were collectively included in Time magazine's compilation of the 20th century's 100 most influential people.[432] In 2014, they received the Grammy Lifetime Achievement Award.[433]
+On 16 January each year, beginning in 2001, people celebrate World Beatles Day under UNESCO. This date has direct relation to the opening of The Cavern Club in 1957.[434][435]
+Five asteroids, 4147 Lennon, 4148 McCartney, 4149 Harrison, 4150 Starr and 8749 Beatles are named after the Beatles. In 2007, the Beatles became the first band to feature on a series of UK postage stamps issued by the Royal Mail.[436]
+Principal members
+Early members
+Touring musician
+The Beatles have a core catalogue consisting of 13 studio albums and one compilation.[437]
+Core catalogue
+See also
+Through 1969, the Beatles' catalogue was published almost exclusively by Northern Songs Ltd, a company formed in February 1963 by music publisher Dick James specifically for Lennon and McCartney, though it later acquired songs by other artists. The company was organised with James and his partner, Emmanuel Silver, owning a controlling interest, variously described as 51% or 50% plus one share. McCartney had 20%. Reports again vary concerning Lennon's portion – 19 or 20% – and Brian Epstein's – 9 or 10% – which he received in lieu of a 25% band management fee.[438][439][440] In 1965, the company went public. Five million shares were created, of which the original principals retained 3.75 million. James and Silver each received 937,500 shares (18.75% of 5 million); Lennon and McCartney each received 750,000 shares (15%); and Epstein's management company, NEMS Enterprises, received 375,000 shares (7.5%). Of the 1.25 million shares put up for sale, Harrison and Starr each acquired 40,000.[441] At the time of the stock offering, Lennon and McCartney renewed their three-year publishing contracts, binding them to Northern Songs until 1973.[442]
+Harrison created Harrisongs to represent his Beatles compositions, but signed a three-year contract with Northern Songs that gave it the copyright to his work through March 1968, which included "Taxman" and "Within You Without You".[443] The songs on which Starr received co-writing credit before 1968, such as "What Goes On" and "Flying", were also Northern Songs copyrights.[444] Harrison did not renew his contract with Northern Songs when it ended, signing instead with Apple Publishing while retaining the copyright to his work from that point on. Harrison thus owns the rights to his later Beatles songs such as "While My Guitar Gently Weeps" and "Something". That year, as well, Starr created Startling Music, which holds the rights to his Beatles compositions, "Don't Pass Me By" and "Octopus's Garden".[445][446]
+In March 1969, James arranged to sell his and his partner's shares of Northern Songs to the British broadcasting company Associated Television (ATV), founded by impresario Lew Grade, without first informing the Beatles. The band then made a bid to gain a controlling interest by attempting to work out a deal with a consortium of London brokerage firms that had accumulated a 14% holding.[447] The deal collapsed over the objections of Lennon, who declared, "I'm sick of being fucked about by men in suits sitting on their fat arses in the City."[448] By the end of May, ATV had acquired a majority stake in Northern Songs, controlling nearly the entire Lennon–McCartney catalogue, as well as any future material until 1973.[449] In frustration, Lennon and McCartney sold their shares to ATV in late October 1969.[450]
+In 1981, financial losses by ATV's parent company, Associated Communications Corporation (ACC), led it to attempt to sell its music division. According to authors Brian Southall and Rupert Perry, Grade contacted McCartney, offering ATV Music and Northern Songs for $30 million.[451] According to an account McCartney gave in 1995, he met with Grade and explained he was interested solely in the Northern Songs catalogue if Grade were ever willing to "separate off" that portion of ATV Music. Soon afterwards, Grade offered to sell him Northern Songs for £20 million, giving the ex-Beatle "a week or so" to decide. By McCartney's account, he and Ono countered with a £5 million bid that was rejected.[452] According to reports at the time, Grade refused to separate Northern Songs and turned down an offer of £21–25 million from McCartney and Ono for Northern Songs. In 1982, ACC was acquired in a takeover by Australian business magnate Robert Holmes à Court for £60 million.[453]
+In 1985, Michael Jackson purchased ATV for a reported $47.5 million. The acquisition gave him control over the publishing rights to more than 200 Beatles songs, as well as 40,000 other copyrights.[454] In 1995, in a deal that earned him a reported $110 million, Jackson merged his music publishing business with Sony, creating a new company, Sony/ATV Music Publishing, in which he held a 50% stake. The merger made the new company, then valued at over half a billion dollars, the third-largest music publisher in the world.[455] In 2016, Sony acquired Jackson's share of Sony/ATV from the Jackson estate for $750 million.[456]
+Despite the lack of publishing rights to most of their songs, Lennon's estate and McCartney continue to receive their respective shares of the writers' royalties, which together are 331⁄3% of total commercial proceeds in the US and which vary elsewhere around the world between 50 and 55%.[457] Two of Lennon and McCartney's earliest songs – "Love Me Do" and "P.S. I Love You" – were published by an EMI subsidiary, Ardmore & Beechwood, before they signed with James. McCartney acquired their publishing rights from Ardmore[458] in 1978,[459] and they are the only two Beatles songs owned by McCartney's company MPL Communications.[460] On 18 January 2017, McCartney filed a suit in the United States district court against Sony/ATV Music Publishing seeking to reclaim ownership of his share of the Lennon–McCartney song catalogue beginning in 2018. Under US copyright law, for works published before 1978 the author can reclaim copyrights assigned to a publisher after 56 years.[461][462] McCartney and Sony agreed to a confidential settlement in June 2017.[463][464]
+Fictionalised
+Documentaries and filmed performances
+1963
+1964
+1965
+1966

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+Homosexuality is romantic attraction, sexual attraction, or sexual behavior between members of the same sex or gender.[1]  As a sexual orientation, homosexuality is "an enduring pattern of emotional, romantic, and/or sexual attractions" to people of the same sex. It "also refers to a person's sense of identity based on those attractions, related behaviors, and membership in a community of others who share those attractions."[2][3]
+Along with bisexuality and heterosexuality, homosexuality is one of the three main categories of sexual orientation within the heterosexual–homosexual continuum.[2] Scientists do not know the exact cause of sexual orientation, but they theorize that it is caused by a complex interplay of genetic, hormonal, and environmental influences,[4][5][6] and do not view it as a choice.[4][5][7] Although no single theory on the cause of sexual orientation has yet gained widespread support, scientists favor biologically-based theories.[4] There is considerably more evidence supporting nonsocial, biological causes of sexual orientation than social ones, especially for males.[8][9][10] There is no substantive evidence which suggests parenting or early childhood experiences play a role with regard to sexual orientation.[11] While some people believe that homosexual activity is unnatural,[12] scientific research shows that homosexuality is a normal and natural variation in human sexuality and is not in and of itself a source of negative psychological effects.[2][13] There is insufficient evidence to support the use of psychological interventions to change sexual orientation.[14][15]
+The most common terms for homosexual people are lesbian for females and gay for males, but gay also commonly refers to both homosexual females and males. The percentage of people who are gay or lesbian and the proportion of people who are in same-sex romantic relationships or have had same-sex sexual experiences are difficult for researchers to estimate reliably for a variety of reasons, including many gay and lesbian people not openly identifying as such due to prejudice or discrimination such as homophobia and heterosexism.[16] Homosexual behavior has also been documented in many non-human animal species.[22]
+Many gay and lesbian people are in committed same-sex relationships, though only in the 2010s have census forms and political conditions facilitated their visibility and enumeration.[23] These relationships are equivalent to heterosexual relationships in essential psychological respects.[3] Homosexual relationships and acts have been admired, as well as condemned, throughout recorded history, depending on the form they took and the culture in which they occurred.[24] Since the end of the 19th century, there has been a global movement towards freedom and equality for gay people, including the introduction of anti-bullying legislation to protect gay children at school, legislation ensuring non-discrimination, equal ability to serve in the military, equal access to health care, equal ability to adopt and parent, and the establishment of marriage equality.
+The word homosexual is a Greek and Latin hybrid, with the first element derived from Greek ὁμός homos, "same" (not related to the Latin homo, "man", as in Homo sapiens), thus connoting sexual acts and affections between members of the same sex, including lesbianism.[25][26] The first known appearance of homosexual in print is found in an 1869 German pamphlet by the Austrian-born novelist Karl-Maria Kertbeny, published anonymously,[27] arguing against a Prussian anti-sodomy law.[27][28] In 1886, the psychiatrist Richard von Krafft-Ebing used the terms homosexual and heterosexual in his book Psychopathia Sexualis. Krafft-Ebing's book was so popular among both laymen and doctors that the terms heterosexual and homosexual became the most widely accepted terms for sexual orientation.[29][30] As such, the current use of the term has its roots in the broader 19th-century tradition of personality taxonomy.
+Many modern style guides in the U.S. recommend against using homosexual as a noun, instead using gay man or lesbian.[31] Similarly, some recommend completely avoiding usage of homosexual as it has a negative, clinical history and because the word only refers to one's sexual behavior (as opposed to romantic feelings) and thus it has a negative connotation.[31] Gay and lesbian are the most common alternatives. The first letters are frequently combined to create the initialism LGBT (sometimes written as GLBT), in which B and T refer to bisexual and transgender people.
+Gay especially refers to male homosexuality,[32] but may be used in a broader sense to refer to all LGBT people. In the context of sexuality, lesbian refers only to female homosexuality. The word lesbian is derived from the name of the Greek island Lesbos, where the poet Sappho wrote largely about her emotional relationships with young women.[33][34]
+Although early writers also used the adjective homosexual to refer to any single-sex context (such as an all-girls school), today the term is used exclusively in reference to sexual attraction, activity, and orientation. The term homosocial is now used to describe single-sex contexts that are not specifically sexual. There is also a word referring to same-sex love, homophilia.
+Some synonyms for same-sex attraction or sexual activity include men who have sex with men or MSM (used in the medical community when specifically discussing sexual activity) and homoerotic (referring to works of art).[35][36] Pejorative terms in English include queer, faggot, fairy, poof, and homo.[37][38][39][40] Beginning in the 1990s, some of these have been reclaimed as positive words by gay men and lesbians, as in the usage of queer studies, queer theory, and even the popular American television program Queer Eye for the Straight Guy.[41] The word homo occurs in many other languages without the pejorative connotations it has in English.[42] As with ethnic slurs and racial slurs, the use of these terms can still be highly offensive. The range of acceptable use for these terms depends on the context and speaker.[43] Conversely, gay, a word originally embraced by homosexual men and women as a positive, affirmative term (as in gay liberation and gay rights),[44] has come into widespread pejorative use among young people.[45]
+The American LGBT rights organization GLAAD advises the media to avoid using the term homosexual to describe gay people or same-sex relationships as the term is "frequently used by anti-gay extremists to denigrate gay people, couples and relationships".[46]
+Some scholars argue that the term "homosexuality" is problematic when applied to ancient cultures since, for example, neither Greeks or Romans possessed any one word covering the same semantic range as the modern concept of "homosexuality".[47][48] Furthermore, there were diverse sexual practices that varied in acceptance depending on time and place.[47] Other scholars argue that there are significant continuities between ancient and modern homosexuality.[49][50]
+In a detailed compilation of historical and ethnographic materials of preindustrial cultures, "strong disapproval of homosexuality was reported for 41% of 42 cultures; it was accepted or ignored by 21%, and 12% reported no such concept. Of 70 ethnographies, 59% reported homosexuality absent or rare in frequency and 41% reported it present or not uncommon."[51]
+In cultures influenced by Abrahamic religions, the law and the church established sodomy as a transgression against divine law or a crime against nature. The condemnation of anal sex between males, however, predates Christian belief. It was frequent in ancient Greece; "unnatural" can be traced back to Plato.[52]
+Many historical figures, including Socrates, Lord Byron, Edward II, and Hadrian,[53] have had terms such as gay or bisexual applied to them. Some scholars, such as Michel Foucault, have regarded this as risking the anachronistic introduction of a contemporary construction of sexuality foreign to their times,[54] though other scholars challenge this.[55][50][49]
+In social science, there has been a dispute between "essentialist" and "constructionist" views of homosexuality. The debate divides those who believe that terms such as "gay" and "straight" refer to objective, culturally invariant properties of persons from those who believe that the experiences they name are artifacts of unique cultural and social processes. "Essentialists" typically believe that sexual preferences are determined by biological forces, while "constructionists" assume that sexual desires are learned.[56] The philosopher of science Michael Ruse has stated that the social constructionist approach, which is influenced by Foucault, is based on a selective reading of the historical record that confuses the existence of homosexual people with the way in which they are labelled or treated.[57]
+The first record of a possible homosexual couple in history is commonly regarded as Khnumhotep and Niankhkhnum, an ancient Egyptian male couple, who lived around 2400 BCE. The pair are portrayed in a nose-kissing position, the most intimate pose in Egyptian art, surrounded by what appear to be their heirs. The anthropologists Stephen Murray and Will Roscoe reported that women in Lesotho engaged in socially sanctioned "long term, erotic relationships" called motsoalle.[58] The anthropologist E. E. Evans-Pritchard also recorded that male Azande warriors in the northern Congo routinely took on young male lovers between the ages of twelve and twenty, who helped with household tasks and participated in intercrural sex with their older husbands.[59]
+As is true of many other non-Western cultures, it is difficult to determine the extent to which Western notions of sexual orientation and gender identity apply to Pre-Columbian cultures. Evidence of homoerotic sexual acts and transvestism has been found in many pre-conquest civilizations in Latin America, such as the Aztecs, Mayas, Quechuas, Moches, Zapotecs, the Incas, and the Tupinambá of Brazil.[60][61][62]
+The Spanish conquerors were horrified to discover sodomy openly practiced among native peoples, and attempted to crush it out by subjecting the berdaches (as the Spanish called them) under their rule to severe penalties, including public execution, burning and being torn to pieces by dogs.[63] The Spanish conquerors talked extensively of sodomy among the natives to depict them as savages and hence justify their conquest and forceful conversion to Christianity. As a result of the growing influence and power of the conquerors, many native cultures started condemning homosexual acts themselves.[citation needed]
+Among some of the indigenous peoples of the Americas in North America prior to European colonization, a relatively common form of same-sex sexuality centered around the figure of the Two-Spirit individual (the term itself was coined only in 1990). Typically, this individual was recognized early in life, given a choice by the parents to follow the path and, if the child accepted the role, raised in the appropriate manner, learning the customs of the gender it had chosen. Two-Spirit individuals were commonly shamans and were revered as having powers beyond those of ordinary shamans. Their sexual life was with the ordinary tribe members of the same sex.[citation needed]
+During the colonial times following the European invasion, homosexuality was prosecuted by the Inquisition, some times leading to death sentences on the charges of sodomy, and the practices became clandestine. Many homosexual individuals went into heterosexual marriages to keep appearances, and many turned to the clergy to escape public scrutiny of their lack of interest in the opposite sex.[citation needed]
+In 1986, the Supreme Court of the United States ruled in Bowers v. Hardwick that a state could criminalize sodomy, but, in 2003, overturned itself in Lawrence v. Texas and thereby legalized homosexual activity throughout the United States of America.
+Same-sex marriage in the United States expanded from one state in 2004 to all fifty states in 2015, through various state court rulings, state legislation, direct popular votes (referendums and initiatives), and federal court rulings.
+In East Asia, same-sex love has been referred to since the earliest recorded history.
+Homosexuality in China, known as the passions of the cut peach and various other euphemisms, has been recorded since approximately 600 BCE. Homosexuality was mentioned in many famous works of Chinese literature. The instances of same-sex affection and sexual interactions described in the classical novel Dream of the Red Chamber seem as familiar to observers in the present as do equivalent stories of romances between heterosexual people during the same period.  Confucianism, being primarily a social and political philosophy, focused little on sexuality, whether homosexual or heterosexual. Ming Dynasty literature, such as Bian Er Chai (弁而釵/弁而钗), portray homosexual relationships between men as more enjoyable and more "harmonious" than heterosexual relationships.[64] Writings from the Liu Song Dynasty by Wang Shunu claimed that homosexuality was as common as heterosexuality in the late 3rd century.[65]
+Opposition to homosexuality in China originates in the medieval Tang Dynasty (618–907), attributed to the rising influence of Christian and Islamic values,[66] but did not become fully established until the Westernization efforts of the late Qing Dynasty and the Republic of China.[67]
+The Laws of Manu mentions a "third sex", members of which may engage in nontraditional gender expression and homosexual activities.[68]
+The earliest Western documents (in the form of literary works, art objects, and mythographic materials) concerning same-sex relationships are derived from ancient Greece.
+In regard to male homosexuality, such documents depict an at times complex understanding in which relationships with women and relationships with adolescent boys could be a part of a normal man's love life. Same-sex relationships were a social institution variously constructed over time and from one city to another. The formal practice, an erotic yet often restrained relationship between a free adult male and a free adolescent, was valued for its pedagogic benefits and as a means of population control, though occasionally blamed for causing disorder. Plato praised its benefits in his early writings[69] but in his late works proposed its prohibition.[70] Aristotle, in the Politics, dismissed Plato's ideas about abolishing homosexuality (2.4); he explains that barbarians like the Celts accorded it a special honor (2.6.6), while the Cretans used it to regulate the population (2.7.5).[71]
+Little is known of female homosexuality in antiquity. Sappho, born on the island of Lesbos, was included by later Greeks in the canonical list of nine lyric poets. The adjectives deriving from her name and place of birth (Sapphic and Lesbian) came to be applied to female homosexuality beginning in the 19th century.[72][73] Sappho's poetry centers on passion and love for various personages and both genders. The narrators of many of her poems speak of infatuations and love (sometimes requited, sometimes not) for various females, but descriptions of physical acts between women are few and subject to debate.[74][75]
+In Ancient Rome, the young male body remained a focus of male sexual attention, but relationships were between older free men and slaves or freed youths who took the receptive role in sex. The Hellenophile emperor Hadrian is renowned for his relationship with Antinous, but the Christian emperor Theodosius I decreed a law on 6 August 390, condemning passive males to be burned at the stake. Notwithstanding these regulations taxes on brothels with boys available for homosexual sex continued to be collected until the end of the reign of Anastasius I in 518. Justinian, towards the end of his reign, expanded the proscription to the active partner as well (in 558), warning that such conduct can lead to the destruction of cities through the "wrath of God".
+During the Renaissance, wealthy cities in northern Italy—Florence and Venice in particular—were renowned for their widespread practice of same-sex love, engaged in by a considerable part of the male population and constructed along the classical pattern of Greece and Rome.[76][77] But even as many of the male population were engaging in same-sex relationships, the authorities, under the aegis of the Officers of the Night court, were prosecuting, fining, and imprisoning a good portion of that population.
+From the second half of the 13th century, death was the punishment for male homosexuality in most of Europe.[78]
+The relationships of socially prominent figures, such as King James I and the Duke of Buckingham, served to highlight the issue, including in anonymously authored street pamphlets: "The world is chang'd I know not how, For men Kiss Men, not Women now;...Of J. the First and Buckingham: He, true it is, his Wives Embraces fled, To slabber his lov'd Ganimede" (Mundus Foppensis, or The Fop Display'd, 1691).
+Love Letters Between a Certain Late Nobleman and the Famous Mr. Wilson was published in 1723 in England, and was presumed by some modern scholars to be a novel. The 1749 edition of John Cleland's popular novel Fanny Hill includes a homosexual scene, but this was removed in its 1750 edition. Also in 1749, the earliest extended and serious defense of homosexuality in English, Ancient and Modern Pederasty Investigated and Exemplified, written by Thomas Cannon, was published, but was suppressed almost immediately. It includes the passage, "Unnatural Desire is a Contradiction in Terms; downright Nonsense. Desire is an amatory Impulse of the inmost human Parts."[79] Around 1785 Jeremy Bentham wrote another defense, but this was not published until 1978.[80] Executions for sodomy continued in the Netherlands until 1803, and in England until 1835, James Pratt and John Smith being the last Englishmen to be so hanged.
+Between 1864 and 1880 Karl Heinrich Ulrichs published a series of twelve tracts, which he collectively titled Research on the Riddle of Man-Manly Love. In 1867, he became the first self-proclaimed homosexual person to speak out publicly in defense of homosexuality when he pleaded at the Congress of German Jurists in Munich for a resolution urging the repeal of anti-homosexual laws.[16] Sexual Inversion by Havelock Ellis, published in 1896, challenged theories that homosexuality was abnormal, as well as stereotypes, and insisted on the ubiquity of homosexuality and its association with intellectual and artistic achievement.[81]
+Although medical texts like these (written partly in Latin to obscure the sexual details) were not widely read by the general public, they did lead to the rise of Magnus Hirschfeld's Scientific-Humanitarian Committee, which campaigned from 1897 to 1933 against anti-sodomy laws in Germany, as well as a much more informal, unpublicized movement among British intellectuals and writers, led by such figures as Edward Carpenter and John Addington Symonds. Beginning in 1894 with Homogenic Love, Socialist activist and poet Edward Carpenter wrote a string of pro-homosexual articles and pamphlets, and "came out" in 1916 in his book My Days and Dreams. In 1900, Elisar von Kupffer published an anthology of homosexual literature from antiquity to his own time, Lieblingminne und Freundesliebe in der Weltliteratur.
+There are a handful of accounts by Arab travelers to Europe during the mid-1800s. Two of these travelers, Rifa'ah al-Tahtawi and Muhammad as-Saffar, show their surprise that the French sometimes deliberately mistranslated love poetry about a young boy, instead referring to a young female, to maintain their social norms and morals.[82]
+Israel is considered the most tolerant country in the Middle East and Asia to homosexuals,[83] with Tel Aviv being named "the gay capital of the Middle East"[84] and considered one of the most gay friendly cities in the world.[85] The annual Pride Parade in support of homosexuality takes place in Tel Aviv.[86]
+On the other hand, many governments in the Middle East often ignore, deny the existence of, or criminalize homosexuality. Homosexuality is illegal in almost all Muslim countries.[87] Same-sex intercourse officially carries the death penalty in several Muslim nations: Saudi Arabia, Iran, Mauritania, northern Nigeria, Sudan, and Yemen.[88]  Iranian President Mahmoud Ahmadinejad, during his 2007 speech at Columbia University, asserted that there were no gay people in Iran. However, the probable reason is that they keep their sexuality a secret for fear of government sanction or rejection by their families.[89]
+In ancient Sumer, a set of priests known as gala worked in the temples of the goddess Inanna, where they performed elegies and lamentations.[91]:285 Gala took female names, spoke in the eme-sal dialect, which was traditionally reserved for women, and appear to have engaged in homosexual intercourse.[92] The Sumerian sign for gala was a ligature of the signs for "penis" and "anus".[92] One Sumerian proverb reads: "When the gala wiped off his ass [he said], 'I must not arouse that which belongs to my mistress [i.e., Inanna].'"[92] In later Mesopotamian cultures, kurgarrū and assinnu were servants of the goddess Ishtar (Inanna's East Semitic equivalent), who dressed in female clothing and performed war dances in Ishtar's temples.[92] Several Akkadian proverbs seem to suggest that they may have also engaged in homosexual intercourse.[92]
+In ancient Assyria, homosexuality was present and common; it was also not prohibited, condemned, nor looked upon as immoral or disordered. Some religious texts contain prayers for divine blessings on homosexual relationships.[93][94] The Almanac of Incantations contained prayers favoring on an equal basis the love of a man for a woman, of a woman for a man, and of a man for man.[95]
+Some scholars argue that there are examples of homosexual love in ancient literature, like in the Mesopotamian Epic of Gilgamesh as well as in the Biblical story of David and Jonathan. In the Epic of Gilgamesh, the relationship between the main protagonist Gilgamesh and the character Enkidu has been seen by some to be homosexual in nature.[96][97][98][99] Similarly, David's love for Jonathan is "greater than the love of women."[100]
+In some societies of Melanesia, especially in Papua New Guinea, same-sex relationships were an integral part of the culture until the middle of the 1900s. The Etoro and Marind-anim for example, viewed heterosexuality as unclean and celebrated homosexuality instead. In some traditional Melanesian cultures a prepubertal boy would be paired with an older adolescent who would become his mentor and who would "inseminate" him (orally, anally, or topically, depending on the tribe) over a number of years in order for the younger to also reach puberty. Many Melanesian societies, however, have become hostile towards same-sex relationships since the introduction of Christianity by European missionaries.[101]
+The American Psychological Association, the American Psychiatric Association, and the National Association of Social Workers identify sexual orientation as "not merely a personal characteristic that can be defined in isolation. Rather, one's sexual orientation defines the universe of persons with whom one is likely to find the satisfying and fulfilling relationships":[3]
+Sexual orientation is commonly discussed as a characteristic of the individual, like biological sex, gender identity, or age. This perspective is incomplete because sexual orientation is always defined in relational terms and necessarily involves relationships with other individuals. Sexual acts and romantic attractions are categorized as homosexual or heterosexual according to the biological sex of the individuals involved in them, relative to each other. Indeed, it is by acting—or desiring to act—with another person that individuals express their heterosexuality, homosexuality, or bisexuality. This includes actions as simple as holding hands with or kissing another person. Thus, sexual orientation is integrally linked to the intimate personal relationships that human beings form with others to meet their deeply felt needs for love, attachment, and intimacy. In addition to sexual behavior, these bonds encompass nonsexual physical affection between partners, shared goals and values, mutual support, and ongoing commitment.[3]
+The Kinsey scale, also called the Heterosexual-Homosexual Rating Scale,[102] attempts to describe a person's sexual history or episodes of his or her sexual activity at a given time. It uses a scale from 0, meaning exclusively heterosexual, to 6, meaning exclusively homosexual. In both the Male and Female volumes of the Kinsey Reports, an additional grade, listed as "X", has been interpreted by scholars to indicate asexuality.[103]
+Often, sexual orientation and sexual identity are not distinguished, which can impact accurately assessing sexual identity and whether or not sexual orientation is able to change; sexual orientation identity can change throughout an individual's life, and may or may not align with biological sex, sexual behavior or actual sexual orientation.[104][105][106] Sexual orientation is stable and unlikely to change for the vast majority of people, but some research indicates that some people may experience change in their sexual orientation, and this is more likely for women than for men.[107] The American Psychological Association distinguishes between sexual orientation (an innate attraction) and sexual orientation identity (which may change at any point in a person's life).[108]
+People with a homosexual orientation can express their sexuality in a variety of ways, and may or may not express it in their behaviors.[2] Many have sexual relationships predominantly with people of their own sex, though some have sexual relationships with those of the opposite sex, bisexual relationships, or none at all (celibacy).[2] Studies have found same-sex and opposite-sex couples to be equivalent to each other in measures of satisfaction and commitment in relationships, that age and sex are more reliable than sexual orientation as a predictor of satisfaction and commitment to a relationship, and that people who are heterosexual or homosexual share comparable expectations and ideals with regard to romantic relationships.[109][110][111]
+Coming out (of the closet) is a phrase referring to one's disclosure of their sexual orientation or gender identity, and is described and experienced variously as a psychological process or journey.[113] Generally, coming out is described in three phases. The first phase is that of "knowing oneself", and the realization emerges that one is open to same-sex relations.[114] This is often described as an internal coming out. The second phase involves one's decision to come out to others, e.g. family, friends, or colleagues. The third phase more generally involves living openly as an LGBT person.[115] In the United States today, people often come out during high school or college age. At this age, they may not trust or ask for help from others, especially when their orientation is not accepted in society. Sometimes their own families are not even informed.
+According to Rosario, Schrimshaw, Hunter, Braun (2006), "the development of a lesbian, gay, or bisexual (LGB) sexual identity is a complex and often difficult process. Unlike members of other minority groups (e.g., ethnic and racial minorities), most LGB individuals are not raised in a community of similar others from whom they learn about their identity and who reinforce and support that identity. Rather, LGB individuals are often raised in communities that are either ignorant of or openly hostile toward homosexuality."[105]
+Outing is the practice of publicly revealing the sexual orientation of a closeted person.[116] Notable politicians, celebrities, military service people, and clergy members have been outed, with motives ranging from malice to political or moral beliefs. Many commentators oppose the practice altogether,[117] while some encourage outing public figures who use their positions of influence to harm other gay people.[118]
+Lesbians often experience their sexuality differently from gay men, and have different understandings about etiology from those derived from studies focused mostly on men.[citation needed]
+In a U.S.-based 1970s mail survey by Shere Hite, lesbians self-reported their reasons for being lesbian. This is the only major piece of research into female sexuality that has looked at how women understand being homosexual since Kinsey in 1953.[citation needed] The research yielded information about women's general understanding of lesbian relationships and their sexual orientation. Women gave various reasons for preferring sexual relations with women to sexual relations with men, including finding women more sensitive to other people's needs.[119]
+Since Hite carried out her study she has acknowledged that some women may have chosen the political identity of a lesbian. Julie Bindel, a UK journalist, reaffirmed that "political lesbianism continues to make intrinsic sense because it reinforces the idea that sexuality is a choice, and we are not destined to a particular fate because of our chromosomes." as recently as 2009.[120]
+Reliable data as to the size of the gay and lesbian population are of value in informing public policy.[123] For example, demographics would help in calculating the costs and benefits of domestic partnership benefits, of the impact of legalizing gay adoption, and of the impact of the U.S. military's Don't Ask Don't Tell policy.[123] Further, knowledge of the size of the "gay and lesbian population holds promise for helping social scientists understand a wide array of important questions—questions about the general nature of labor market choices, accumulation of human capital, specialization within households, discrimination, and decisions about geographic location."[123]
+Measuring the prevalence of homosexuality presents difficulties. It is necessary to consider the measuring criteria that are used, the cutoff point and the time span taken to define a sexual orientation.[16] Many people, despite having same-sex attractions, may be reluctant to identify themselves as gay or bisexual. The research must measure some characteristic that may or may not be defining of sexual orientation. The number of people with same-sex desires may be larger than the number of people who act on those desires, which in turn may be larger than the number of people who self-identify as gay, lesbian, or bisexual.[123]
+In 1948 and 1953, Alfred Kinsey reported that nearly 46% of the male subjects had "reacted" sexually to persons of both sexes in the course of their adult lives, and 37% had had at least one homosexual experience.[124][125] Kinsey's methodology was criticized by John Tukey for using convenience samples and not random samples.[126][127]
+A later study tried to eliminate the sample bias, but still reached similar conclusions.[128] Simon LeVay cites these Kinsey results as an example of the caution needed to interpret demographic studies, as they may give quite differing numbers depending on what criteria are used to conduct them, in spite of using sound scientific methods.[16]
+According to major studies, 2% to 11% of people have had some form of same-sex sexual contact within their lifetime;[129][130][131][132][133][134][135][136][137][138] this percentage rises to 16–21% when either or both same-sex attraction and behavior are reported.[138] In a 2006 study, 20% of respondents anonymously reported some homosexual feelings, although only 2–3% identified themselves as homosexual.[139] A 1992 study reported that 6.1% of males in Britain have had a homosexual experience, while in France the number was reported at 4.1%.[140]
+In the United States, according to a report by The Williams Institute in April 2011, 3.5% or approximately 9 million of the adult population identify as lesbian, gay, or bisexual.[141][142] According to the 2000 United States Census, there were about 601,209 same-sex unmarried partner households.[143]
+A 2013 study by the CDC, in which over 34,000 Americans were interviewed, puts the percentage of self-identifying lesbians and gay men at 1.6%, and of bisexuals at 0.7%.[144]
+According to a 2008 poll, 13% of Britons have had some form of same-sex sexual contact while only 6% of Britons identify themselves as either homosexual or bisexual.[145] In contrast, a survey by the UK Office for National Statistics (ONS) in 2010 found that 95% of Britons identified as heterosexual, 1.5% of Britons identified themselves as homosexual or bisexual, and the last 3.5% gave more vague answers such as "don't know", "other", or did not respond to the question.[125][146]
+In October 2012, Gallup started conducting annual surveys to study the demographics of LGBT people, determining that 3.4% (±1%) of adults identified as LGBT in the United States.[147] It was the nation's largest poll on the issue at the time.[148][149] In 2017, the percentage was estimated to have risen to 4.5% of adults, with the increase largely driven by Millennials. The poll attributes the rise to greater willingness of younger people to reveal their sexual identity.[150]
+The American Psychological Association, the American Psychiatric Association, and the National Association of Social Workers state:
+In 1952, when the American Psychiatric Association published its first Diagnostic and Statistical Manual of Mental Disorders, homosexuality was included as a disorder. Almost immediately, however, that classification began to be subjected to critical scrutiny in research funded by the National Institute of Mental Health. That study and subsequent research consistently failed to produce any empirical or scientific basis for regarding homosexuality as a disorder or abnormality, rather than a normal and healthy sexual orientation. As results from such research accumulated, professionals in medicine, mental health, and the behavioral and social sciences reached the conclusion that it was inaccurate to classify homosexuality as a mental disorder and that the DSM classification reflected untested assumptions based on once-prevalent social norms and clinical impressions from unrepresentative samples comprising patients seeking therapy and individuals whose conduct brought them into the criminal justice system.
+In recognition of the scientific evidence,[151] the American Psychiatric Association removed homosexuality from the DSM in 1973, stating that "homosexuality per se implies no impairment in judgment, stability, reliability, or general social or vocational capabilities." After thoroughly reviewing the scientific data, the American Psychological Association adopted the same position in 1975, and urged all mental health professionals "to take the lead in removing the stigma of mental illness that has long been associated with homosexual orientations." The National Association of Social Workers has adopted a similar policy.
+Thus, mental health professionals and researchers have long recognized that being homosexual poses no inherent obstacle to leading a happy, healthy, and productive life, and that the vast majority of gay and lesbian people function well in the full array of social institutions and interpersonal relationships.[3]
+The consensus of research and clinical literature demonstrates that same-sex sexual and romantic attractions, feelings, and behaviors are normal and positive variations of human sexuality.[152] There is now a large body of research evidence that indicates that being gay, lesbian or bisexual is compatible with normal mental health and social adjustment.[11] The World Health Organization's ICD-9 (1977) listed homosexuality as a mental illness; it was removed from the ICD-10, endorsed by the Forty-third World Health Assembly on 17 May 1990.[153][154][155] Like the DSM-II, the ICD-10 added ego-dystonic sexual orientation to the list, which refers to people who want to change their gender identities or sexual orientation because of a psychological or behavioral disorder (F66.1). The Chinese Society of Psychiatry removed homosexuality from its Chinese Classification of Mental Disorders in 2001 after five years of study by the association.[156] According to the Royal College of Psychiatrists "This unfortunate history demonstrates how marginalisation of a group of people who have a particular personality feature (in this case homosexuality) can lead to harmful medical practice and a basis for discrimination in society.[11] There is now a large body of research evidence that indicates that being gay, lesbian or bisexual is compatible with normal mental health and social adjustment. However, the experiences of discrimination in society and possible rejection by friends, families and others, such as employers, means that some LGB people experience a greater than expected prevalence of mental health difficulties and substance misuse problems. Although there have been claims by conservative political groups in the USA that this higher prevalence of mental health difficulties is confirmation that homosexuality is itself a mental disorder, there is no evidence whatever to substantiate such a claim."[157]
+Most lesbian, gay, and bisexual people who seek psychotherapy do so for the same reasons as heterosexual people (stress, relationship difficulties, difficulty adjusting to social or work situations, etc.); their sexual orientation may be of primary, incidental, or no importance to their issues and treatment. Whatever the issue, there is a high risk for anti-gay bias in psychotherapy with lesbian, gay, and bisexual clients.[158] Psychological research in this area has been relevant to counteracting prejudicial ("homophobic") attitudes and actions, and to the LGBT rights movement generally.[159]
+The appropriate application of affirmative psychotherapy is based on the following scientific facts:[152]
+Although scientists favor biological models for the cause of sexual orientation,[4] they do not believe that the development of sexual orientation is the result of any one factor. They generally believe that it is determined by a complex interplay of biological and environmental factors, and is shaped at an early age.[2][5][6] There is considerably more evidence supporting nonsocial, biological causes of sexual orientation than social ones, especially for males.[8] There is no substantive evidence which suggests parenting or early childhood experiences play a role with regard to sexual orientation.[11] Scientists do not believe that sexual orientation is a choice.[4][5][7]
+The American Academy of Pediatrics stated in Pediatrics in 2004:
+There is no scientific evidence that abnormal parenting, sexual abuse, or other adverse life events influence sexual orientation. Current knowledge suggests that sexual orientation is usually established during early childhood.[4][160]
+The American Psychological Association, American Psychiatric Association, and National Association of Social Workers stated in 2006:
+Currently, there is no scientific consensus about the specific factors that cause an individual to become heterosexual, homosexual, or bisexual—including possible biological, psychological, or social effects of the parents' sexual orientation. However, the available evidence indicates that the vast majority of lesbian and gay adults were raised by heterosexual parents and the vast majority of children raised by lesbian and gay parents eventually grow up to be heterosexual.[2]
+Despite numerous attempts, no "gay gene" has been identified. However, there is substantial evidence for a genetic basis of homosexuality, especially in males, based on twin studies; some association with regions of Chromosome 8, the Xq28 locus on the X chromosome, and other sites across many chromosomes.[161]
+Sanders et al. 2015
+Sanders et al. 2015
+Starting in the 2010s, potential epigenetic factors have become a topic of increased attention in genetic research on sexual orientation. A study presented at the ASHG 2015 Annual Meeting found that the methylation pattern in nine regions of the genome appeared very closely linked to sexual orientation, with a resulting algorithm using the methylation pattern to predict the sexual orientation of a control group with almost 70% accuracy.[164][165]
+Research into the causes of homosexuality plays a role in political and social debates and also raises concerns about genetic profiling and prenatal testing.[166][167]
+Since homosexuality tends to lower reproductive success, and since there is considerable evidence that human sexual orientation is genetically influenced, it is unclear how it is maintained in the population at a relatively high frequency.[168] There are many possible explanations, such as genes predisposing to homosexuality also conferring advantage in heterosexuals, a kin selection effect, social prestige, and more.[169] A 2009 study also suggested a significant increase in fecundity in the females related to the homosexual people from the maternal line (but not in those related from the paternal one).[170]
+There are no studies of adequate scientific rigor that conclude that sexual orientation change efforts work to change a person's sexual orientation. Those efforts have been controversial due to tensions between the values held by some faith-based organizations, on the one hand, and those held by LGBT rights organizations and professional and scientific organizations and other faith-based organizations, on the other.[14] The longstanding consensus of the behavioral and social sciences and the health and mental health professions is that homosexuality per se is a normal and positive variation of human sexual orientation, and therefore not a mental disorder.[14] The American Psychological Association says that "most people experience little or no sense of choice about their sexual orientation".[171]
+Some individuals and groups have promoted the idea of homosexuality as symptomatic of developmental defects or spiritual and moral failings and have argued that sexual orientation change efforts, including psychotherapy and religious efforts, could alter homosexual feelings and behaviors. Many of these individuals and groups appeared to be embedded within the larger context of conservative religious political movements that have supported the stigmatization of homosexuality on political or religious grounds.[14]
+No major mental health professional organization has sanctioned efforts to change sexual orientation and virtually all of them have adopted policy statements cautioning the profession and the public about treatments that purport to change sexual orientation. These include the American Psychiatric Association, American Psychological Association, American Counseling Association, National Association of Social Workers in the US,[172] the Royal College of Psychiatrists,[173] and the Australian Psychological Society.[174] The American Psychological Association and the Royal College of Psychiatrists expressed concerns that the positions espoused by NARTH are not supported by the science and create an environment in which prejudice and discrimination can flourish.[173][175]
+The American Psychological Association states that "sexual orientation is not a choice that can be changed at will, and that sexual orientation is most likely the result of a complex interaction of environmental, cognitive and biological factors...is shaped at an early age...[and evidence suggests] biological, including genetic or inborn hormonal factors, play a significant role in a person's sexuality."[5] They say that "sexual orientation identity—not sexual orientation—appears to change via psychotherapy, support groups, and life events."[14] The American Psychiatric Association says "individuals maybe become aware at different points in their lives that they are heterosexual, gay, lesbian, or bisexual" and "opposes any psychiatric treatment, such as 'reparative' or 'conversion' therapy, which is based upon the assumption that homosexuality per se is a mental disorder, or based upon a prior assumption that the patient should change his/her homosexual orientation". They do, however, encourage gay affirmative psychotherapy.[176] Similarly, the American *Psychological* Association[177] is doubtful about the effectiveness and side-effect profile of sexual orientation change efforts, including conversion therapy.
+The American Psychological Association "encourages mental health professionals to avoid misrepresenting the efficacy of sexual orientation change efforts by promoting or promising change in sexual orientation when providing assistance to individuals distressed by their own or others' sexual orientation and concludes that the benefits reported by participants in sexual orientation change efforts can be gained through approaches that do not attempt to change sexual orientation".[14]
+Scientific research has been generally consistent in showing that lesbian and gay parents are as fit and capable as heterosexual parents, and their children are as psychologically healthy and well-adjusted as children reared by heterosexual parents.[178][179][180] According to scientific literature reviews, there is no evidence to the contrary.[3][181][182][183][184]
+A 2001 review suggested that the children with lesbian or gay parents appear less traditionally gender-typed and are more likely to be open to homoerotic relationships, partly due to genetic (80% of the children being raised by same-sex couples in the US are not adopted and most are the result of previous heterosexual marriages.[185]) and family socialization processes (children grow up in relatively more tolerant school, neighborhood, and social contexts, which are less heterosexist), even though majority of children raised by same-sex couples identify as heterosexual.[186] A 2005 review by Charlotte J. Patterson for the American Psychological Association found that the available data did not suggest higher rates of homosexuality among the children of lesbian or gay parents.[187]
+The terms "men who have sex with men" (MSM) and "women who have sex with women" (WSW) refer to people who engage in sexual activity with others of the same sex regardless of how they identify themselves—as many choose not to accept social identities as lesbian, gay and bisexual.[188][189][190][191][192] These terms are often used in medical literature and social research to describe such groups for study, without needing to consider the issues of sexual self-identity. The terms are seen as problematic by some, however, because they "obscure social dimensions of sexuality; undermine the self-labeling of lesbian, gay, and bisexual people; and do not sufficiently describe variations in sexual behavior".[193]
+In contrast to its benefits, sexual behavior can be a disease vector. Safe sex is a relevant harm reduction philosophy.[194] Many countries currently prohibit men who have sex with men from donating blood; the policy of the United States Food and Drug Administration states that "they are, as a group, at increased risk for HIV, hepatitis B and certain other infections that can be transmitted by transfusion."[195]
+These safer sex recommendations are agreed upon by public health officials for women who have sex with women to avoid sexually transmitted infections (STIs):
+These safer sex recommendations are agreed upon by public health officials for men who have sex with men to avoid sexually transmitted infections:
+When it was first described in medical literature, homosexuality was often approached from a view that sought to find an inherent psychopathology as its root cause. Much literature on mental health and homosexual patients centered on their depression, substance abuse, and suicide. Although these issues exist among people who are non-heterosexual, discussion about their causes shifted after homosexuality was removed from the Diagnostic and Statistical Manual (DSM) in 1973. Instead, social ostracism, legal discrimination, internalization of negative stereotypes, and limited support structures indicate factors homosexual people face in Western societies that often adversely affect their mental health.[199]
+Stigma, prejudice, and discrimination stemming from negative societal attitudes toward homosexuality lead to a higher prevalence of mental health disorders among lesbians, gay men, and bisexuals compared to their heterosexual peers.[200] Evidence indicates that the liberalization of these attitudes over the 1990s through the 2010s is associated with a decrease in such mental health risks among younger LGBT people.[201]
+Gay and lesbian youth bear an increased risk of suicide, substance abuse, school problems, and isolation because of a "hostile and condemning environment, verbal and physical abuse, rejection and isolation from family and peers".[202] Further, LGBT youths are more likely to report psychological and physical abuse by parents or caretakers, and more sexual abuse. Suggested reasons for this disparity are that (1) LGBT youths may be specifically targeted on the basis of their perceived sexual orientation or gender non-conforming appearance, and (2) that "risk factors associated with sexual minority status, including discrimination, invisibility, and rejection by family members...may lead to an increase in behaviors that are associated with risk for victimization, such as substance abuse, sex with multiple partners, or running away from home as a teenager."[203] A 2008 study showed a correlation between the degree of rejecting behavior by parents of LGB adolescents and negative health problems in the teenagers studied:
+Higher rates of family rejection were significantly associated with poorer health outcomes. On the basis of odds ratios, lesbian, gay, and bisexual young adults who reported higher levels of family rejection during adolescence were 8.4 times more likely to report having attempted suicide, 5.9 times more likely to report high levels of depression, 3.4 times more likely to use illegal drugs, and 3.4 times more likely to report having engaged in unprotected sexual intercourse compared with peers from families that reported no or low levels of family rejection.[204]
+Crisis centers in larger cities and information sites on the Internet have arisen to help youth and adults.[205] The Trevor Project, a suicide prevention helpline for gay youth, was established following the 1998 airing on HBO of the Academy Award winning short film Trevor.[206]
+Most nations do not prohibit consensual sex between unrelated persons above the local age of consent. Some jurisdictions further recognize identical rights, protections, and privileges for the family structures of same-sex couples, including marriage. Some countries and jurisdictions mandate that all individuals restrict themselves to heterosexual activity and disallow homosexual activity via sodomy laws. Offenders can face the death penalty in Islamic countries and jurisdictions ruled by sharia. There are, however, often significant differences between official policy and real-world enforcement.
+Although homosexual acts were decriminalized in some parts of the Western world, such as Poland in 1932, Denmark in 1933, Sweden in 1944, and the England and Wales in 1967, it was not until the mid-1970s that the gay community first began to achieve limited civil rights in some developed countries. A turning point was reached in 1973 when the American Psychiatric Association, which previously listed homosexuality in the DSM-I in 1952, removed homosexuality in the DSM-II, in recognition of scientific evidence.[3] In 1977, Quebec became the first state-level jurisdiction in the world to prohibit discrimination on the grounds of sexual orientation. During the 1980s and 1990s, most developed countries enacted laws decriminalizing homosexual behavior and prohibiting discrimination against lesbian and gay people in employment, housing, and services. On the other hand, many countries today in the Middle East and Africa, as well as several countries in Asia, the Caribbean and the South Pacific, outlaw homosexuality. In 2013, the Supreme Court of India upheld Section 377 of the Indian Penal Code,[207] but in 2018 overturned itself and legalized homosexual activity throughout India.[208] The Section 377, a British colonial-era law which criminalizes homosexual activity, remains in effect in more than 40 former British colonies.[209][210]
+10 countries or jurisdictions, all of which are Islamic and ruled by sharia, impose the death penalty for homosexuality.
+In the European Union, discrimination of any type based on sexual orientation or gender identity is illegal under the Charter of Fundamental Rights of the European Union.[220]
+Since the 1960s, many LGBT people in the West, particularly those in major metropolitan areas, have developed a so-called gay culture. To many,[who?] gay culture is exemplified by the gay pride movement, with annual parades and displays of rainbow flags. Yet not all LGBT people choose to participate in "queer culture", and many gay men and women specifically decline to do so. To some[who?] it seems to be a frivolous display, perpetuating gay stereotypes.
+With the outbreak of AIDS in the early 1980s, many LGBT groups and individuals organized campaigns to promote efforts in AIDS education, prevention, research, patient support, and community outreach, as well as to demand government support for these programs.
+The death toll wrought by the AIDS epidemic at first seemed to slow the progress of the gay rights movement, but in time it galvanized some parts of the LGBT community into community service and political action, and challenged the heterosexual community to respond compassionately. Major American motion pictures from this period that dramatized the response of individuals and communities to the AIDS crisis include An Early Frost (1985), Longtime Companion (1990), And the Band Played On (1993), Philadelphia (1993), and Common Threads: Stories from the Quilt (1989).
+Publicly gay politicians have attained numerous government posts, even in countries that had sodomy laws in their recent past. Examples include Guido Westerwelle, Germany's Vice-Chancellor; Peter Mandelson, a British Labour Party cabinet minister and Per-Kristian Foss, formerly Norwegian Minister of Finance.
+LGBT movements are opposed by a variety of individuals and organizations. Some social conservatives believe that all sexual relationships with people other than an opposite-sex spouse undermine the traditional family[221] and that children should be reared in homes with both a father and a mother.[222][223] Some argue that gay rights may conflict with individuals' freedom of speech,[224][225] religious freedoms in the workplace,[226][227] the ability to run churches,[228] charitable organizations[229][230] and other religious organizations[231] in accordance with one's religious views, and that the acceptance of homosexual relationships by religious organizations might be forced through threatening to remove the tax-exempt status of churches whose views do not align with those of the government.[232][233][234][235] Some critics charge that political correctness has led to the association of sex between males and HIV being downplayed.[236]
+Policies and attitudes toward gay and lesbian military personnel vary widely around the world. Some countries allow gay men, lesbians, and bisexual people to serve openly and have granted them the same rights and privileges as their heterosexual counterparts. Many countries neither ban nor support LGB service members. A few countries continue to ban homosexual personnel outright.
+Most Western military forces have removed policies excluding sexual minority members. Of the 26 countries that participate militarily in NATO, more than 20 permit openly gay, lesbian and bisexual people to serve. Of the permanent members of the United Nations Security Council, three (United Kingdom, France and United States) do so. The other two generally do not: China bans gay and lesbian people outright, Russia excludes all gay and lesbian people during peacetime but allows some gay men to serve in wartime (see below). Israel is the only country in the Middle East region that allows openly LGB people to serve in the military.
+While the question of homosexuality in the military has been highly politicized in the United States, it is not necessarily so in many countries. Generally speaking, sexuality in these cultures is considered a more personal aspect of one's identity than it is in the United States.
+According to the American Psychological Association, empirical evidence fails to show that sexual orientation is germane to any aspect of military effectiveness including unit cohesion, morale, recruitment and retention.[237] Sexual orientation is irrelevant to task cohesion, the only type of cohesion that critically predicts the team's military readiness and success.[238]
+Societal acceptance of non-heterosexual orientations such as homosexuality is lowest in Asian, African and Eastern European countries,[239][240] and is highest in Western Europe, Australia, and the Americas. Western society has become increasingly accepting of homosexuality since the 1990s. In 2017, Professor Amy Adamczyk contended that these cross-national differences in acceptance can be largely explained by three factors: the relative strength of democratic institutions, the level of economic development, and the religious context of the places where people live.[241]
+In 2006, the American Psychological Association, American Psychiatric Association and National Association of Social Workers stated in an amicus brief presented to the Supreme Court of California: "Gay men and lesbians form stable, committed relationships that are equivalent to heterosexual relationships in essential respects. The institution of marriage offers social, psychological, and health benefits that are denied to same-sex couples. By denying same-sex couples the right to marry, the state reinforces and perpetuates the stigma historically associated with homosexuality. Homosexuality remains stigmatized, and this stigma has negative consequences. California's prohibition on marriage for same-sex couples reflects and reinforces this stigma". They concluded: "There is no scientific basis for distinguishing between same-sex couples and heterosexual couples with respect to the legal rights, obligations, benefits, and burdens conferred by civil marriage."[3]
+Though the relationship between homosexuality and religion is complex, current authoritative bodies and doctrines of the world's largest religions view homosexual behaviour negatively.[citation needed] This can range from quietly discouraging homosexual activity, to explicitly forbidding same-sex sexual practices among adherents and actively opposing social acceptance of homosexuality. Some teach that homosexual desire itself is sinful,[242] others state that only the sexual act is a sin,[243] others are completely accepting of gays and lesbians,[244] while some encourage homosexuality.[245] Some claim that homosexuality can be overcome through religious faith and practice. On the other hand, voices exist within many of these religions that view homosexuality more positively, and liberal religious denominations may bless same-sex marriages. Some view same-sex love and sexuality as sacred, and a mythology of same-sex love can be found around the world.[246]
+Gay bullying can be the verbal or physical abuse against a person who is perceived by the aggressor to be  lesbian, gay,  bisexual or transgender, including persons who are actually heterosexual or of non-specific or unknown sexual orientation.  In the US, teenage students heard anti-gay slurs such as "homo", "faggot" and "sissy" about 26 times a day on average, or once every 14 minutes, according to a 1998 study by Mental Health America (formerly National Mental Health Association).[247]
+In many cultures, homosexual people are frequently subject to prejudice and discrimination. A 2011 Dutch study concluded that 49% of Holland's youth and 58% of youth foreign to the country reject homosexuality.[248]  Similar to other minority groups they can also be subject to stereotyping. These attitudes tend to be due to forms of homophobia and heterosexism (negative attitudes, bias, and discrimination in favor of opposite-sex sexuality and relationships). Heterosexism can include the presumption that everyone is heterosexual or that opposite-sex attractions and relationships are the norm and therefore superior. Homophobia is a fear of, aversion to, or discrimination against homosexual people. It manifests in different forms, and a number of different types have been postulated, among which are internalized homophobia, social homophobia, emotional homophobia, rationalized homophobia, and others.[249] Similar is lesbophobia (specifically targeting lesbians) and biphobia (against bisexual people). When such attitudes manifest as crimes they are often called hate crimes and gay bashing.
+Negative stereotypes characterize LGB people as less romantically stable, more promiscuous and more likely to abuse children, but there is no scientific basis to such assertions. Gay men and lesbians form stable, committed relationships that are equivalent to heterosexual relationships in essential respects.[3] Sexual orientation does not affect the likelihood that people will abuse children.[250][251][252] Claims that there is scientific evidence to support an association between being gay and being a pedophile are based on misuses of those terms and misrepresentation of the actual evidence.[251]
+In the United States, the FBI reported that 20.4% of hate crimes reported to law enforcement in 2011 were based on sexual orientation bias. 56.7% of these crimes were based on bias against homosexual men. 11.1% were based on bias against homosexual women. 29.6% were based on anti-homosexual bias without regard to gender.[253] The 1998 murder of Matthew Shepard, a gay student, is a notorious such incident in the U.S. LGBT people, especially lesbians, may become the victims of "corrective rape", a violent crime with the supposed aim of making them heterosexual. In certain parts of the world, LGBT people are also at risk of "honor killings" perpetrated by their families or relatives.[254][255][256]
+In Morocco, a constitutional monarchy following Islamic laws, homosexual acts are a punishable offence. With a population hostile towards LGBT people, the country has witnessed public demonstrations against homosexuals, public denunciations of presumed homosexual individuals, as well as violent intrusions in private homes. The community in the country is exposed to additional risk of prejudice, social rejection and violence, with a greater impossibility of obtaining protection even from the police.[257]
+In 2017, Abderrahim El Habachi became one of the many homosexual Moroccans to flee the country for fear, and sought shelter in the United Kingdom. In August 2019, he accused the UK Home Office of not taking LGBT migrants seriously. He highlighted that they are being put through tough times because the Home Office don't believe their sexual orientation or in the struggles they face in their countries.[258]
+Homosexual and bisexual behaviors occur in a number of other animal species. Such behaviors include sexual activity, courtship, affection, pair bonding, and parenting,[20] and are widespread; a 1999 review by researcher Bruce Bagemihl shows that homosexual behavior has been documented in about 500 species, ranging from primates to gut worms.[20][21] Animal sexual behavior takes many different forms, even within the same species. The motivations for and implications of these behaviors have yet to be fully understood, since most species have yet to be fully studied.[260] According to Bagemihl, "the animal kingdom [does] it with much greater sexual diversity—including homosexual, bisexual and nonreproductive sex—than the scientific community and society at large have previously been willing to accept".[261]
+A review paper by N. W. Bailey and Marlene Zuk looking into studies of same-sex sexual behaviour in animals challenges the view that such behaviour lowers reproductive success, citing several hypotheses about how same-sex sexual behavior might be adaptive; these hypotheses vary greatly among different species. Bailey and Zuk also suggest future research needs to look into evolutionary consequences of same-sex sexual behaviour, rather than only looking into origins of such behaviour.[262]
+Category:LGBT culture

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+Homosexuality is romantic attraction, sexual attraction, or sexual behavior between members of the same sex or gender.[1]  As a sexual orientation, homosexuality is "an enduring pattern of emotional, romantic, and/or sexual attractions" to people of the same sex. It "also refers to a person's sense of identity based on those attractions, related behaviors, and membership in a community of others who share those attractions."[2][3]
+Along with bisexuality and heterosexuality, homosexuality is one of the three main categories of sexual orientation within the heterosexual–homosexual continuum.[2] Scientists do not know the exact cause of sexual orientation, but they theorize that it is caused by a complex interplay of genetic, hormonal, and environmental influences,[4][5][6] and do not view it as a choice.[4][5][7] Although no single theory on the cause of sexual orientation has yet gained widespread support, scientists favor biologically-based theories.[4] There is considerably more evidence supporting nonsocial, biological causes of sexual orientation than social ones, especially for males.[8][9][10] There is no substantive evidence which suggests parenting or early childhood experiences play a role with regard to sexual orientation.[11] While some people believe that homosexual activity is unnatural,[12] scientific research shows that homosexuality is a normal and natural variation in human sexuality and is not in and of itself a source of negative psychological effects.[2][13] There is insufficient evidence to support the use of psychological interventions to change sexual orientation.[14][15]
+The most common terms for homosexual people are lesbian for females and gay for males, but gay also commonly refers to both homosexual females and males. The percentage of people who are gay or lesbian and the proportion of people who are in same-sex romantic relationships or have had same-sex sexual experiences are difficult for researchers to estimate reliably for a variety of reasons, including many gay and lesbian people not openly identifying as such due to prejudice or discrimination such as homophobia and heterosexism.[16] Homosexual behavior has also been documented in many non-human animal species.[22]
+Many gay and lesbian people are in committed same-sex relationships, though only in the 2010s have census forms and political conditions facilitated their visibility and enumeration.[23] These relationships are equivalent to heterosexual relationships in essential psychological respects.[3] Homosexual relationships and acts have been admired, as well as condemned, throughout recorded history, depending on the form they took and the culture in which they occurred.[24] Since the end of the 19th century, there has been a global movement towards freedom and equality for gay people, including the introduction of anti-bullying legislation to protect gay children at school, legislation ensuring non-discrimination, equal ability to serve in the military, equal access to health care, equal ability to adopt and parent, and the establishment of marriage equality.
+The word homosexual is a Greek and Latin hybrid, with the first element derived from Greek ὁμός homos, "same" (not related to the Latin homo, "man", as in Homo sapiens), thus connoting sexual acts and affections between members of the same sex, including lesbianism.[25][26] The first known appearance of homosexual in print is found in an 1869 German pamphlet by the Austrian-born novelist Karl-Maria Kertbeny, published anonymously,[27] arguing against a Prussian anti-sodomy law.[27][28] In 1886, the psychiatrist Richard von Krafft-Ebing used the terms homosexual and heterosexual in his book Psychopathia Sexualis. Krafft-Ebing's book was so popular among both laymen and doctors that the terms heterosexual and homosexual became the most widely accepted terms for sexual orientation.[29][30] As such, the current use of the term has its roots in the broader 19th-century tradition of personality taxonomy.
+Many modern style guides in the U.S. recommend against using homosexual as a noun, instead using gay man or lesbian.[31] Similarly, some recommend completely avoiding usage of homosexual as it has a negative, clinical history and because the word only refers to one's sexual behavior (as opposed to romantic feelings) and thus it has a negative connotation.[31] Gay and lesbian are the most common alternatives. The first letters are frequently combined to create the initialism LGBT (sometimes written as GLBT), in which B and T refer to bisexual and transgender people.
+Gay especially refers to male homosexuality,[32] but may be used in a broader sense to refer to all LGBT people. In the context of sexuality, lesbian refers only to female homosexuality. The word lesbian is derived from the name of the Greek island Lesbos, where the poet Sappho wrote largely about her emotional relationships with young women.[33][34]
+Although early writers also used the adjective homosexual to refer to any single-sex context (such as an all-girls school), today the term is used exclusively in reference to sexual attraction, activity, and orientation. The term homosocial is now used to describe single-sex contexts that are not specifically sexual. There is also a word referring to same-sex love, homophilia.
+Some synonyms for same-sex attraction or sexual activity include men who have sex with men or MSM (used in the medical community when specifically discussing sexual activity) and homoerotic (referring to works of art).[35][36] Pejorative terms in English include queer, faggot, fairy, poof, and homo.[37][38][39][40] Beginning in the 1990s, some of these have been reclaimed as positive words by gay men and lesbians, as in the usage of queer studies, queer theory, and even the popular American television program Queer Eye for the Straight Guy.[41] The word homo occurs in many other languages without the pejorative connotations it has in English.[42] As with ethnic slurs and racial slurs, the use of these terms can still be highly offensive. The range of acceptable use for these terms depends on the context and speaker.[43] Conversely, gay, a word originally embraced by homosexual men and women as a positive, affirmative term (as in gay liberation and gay rights),[44] has come into widespread pejorative use among young people.[45]
+The American LGBT rights organization GLAAD advises the media to avoid using the term homosexual to describe gay people or same-sex relationships as the term is "frequently used by anti-gay extremists to denigrate gay people, couples and relationships".[46]
+Some scholars argue that the term "homosexuality" is problematic when applied to ancient cultures since, for example, neither Greeks or Romans possessed any one word covering the same semantic range as the modern concept of "homosexuality".[47][48] Furthermore, there were diverse sexual practices that varied in acceptance depending on time and place.[47] Other scholars argue that there are significant continuities between ancient and modern homosexuality.[49][50]
+In a detailed compilation of historical and ethnographic materials of preindustrial cultures, "strong disapproval of homosexuality was reported for 41% of 42 cultures; it was accepted or ignored by 21%, and 12% reported no such concept. Of 70 ethnographies, 59% reported homosexuality absent or rare in frequency and 41% reported it present or not uncommon."[51]
+In cultures influenced by Abrahamic religions, the law and the church established sodomy as a transgression against divine law or a crime against nature. The condemnation of anal sex between males, however, predates Christian belief. It was frequent in ancient Greece; "unnatural" can be traced back to Plato.[52]
+Many historical figures, including Socrates, Lord Byron, Edward II, and Hadrian,[53] have had terms such as gay or bisexual applied to them. Some scholars, such as Michel Foucault, have regarded this as risking the anachronistic introduction of a contemporary construction of sexuality foreign to their times,[54] though other scholars challenge this.[55][50][49]
+In social science, there has been a dispute between "essentialist" and "constructionist" views of homosexuality. The debate divides those who believe that terms such as "gay" and "straight" refer to objective, culturally invariant properties of persons from those who believe that the experiences they name are artifacts of unique cultural and social processes. "Essentialists" typically believe that sexual preferences are determined by biological forces, while "constructionists" assume that sexual desires are learned.[56] The philosopher of science Michael Ruse has stated that the social constructionist approach, which is influenced by Foucault, is based on a selective reading of the historical record that confuses the existence of homosexual people with the way in which they are labelled or treated.[57]
+The first record of a possible homosexual couple in history is commonly regarded as Khnumhotep and Niankhkhnum, an ancient Egyptian male couple, who lived around 2400 BCE. The pair are portrayed in a nose-kissing position, the most intimate pose in Egyptian art, surrounded by what appear to be their heirs. The anthropologists Stephen Murray and Will Roscoe reported that women in Lesotho engaged in socially sanctioned "long term, erotic relationships" called motsoalle.[58] The anthropologist E. E. Evans-Pritchard also recorded that male Azande warriors in the northern Congo routinely took on young male lovers between the ages of twelve and twenty, who helped with household tasks and participated in intercrural sex with their older husbands.[59]
+As is true of many other non-Western cultures, it is difficult to determine the extent to which Western notions of sexual orientation and gender identity apply to Pre-Columbian cultures. Evidence of homoerotic sexual acts and transvestism has been found in many pre-conquest civilizations in Latin America, such as the Aztecs, Mayas, Quechuas, Moches, Zapotecs, the Incas, and the Tupinambá of Brazil.[60][61][62]
+The Spanish conquerors were horrified to discover sodomy openly practiced among native peoples, and attempted to crush it out by subjecting the berdaches (as the Spanish called them) under their rule to severe penalties, including public execution, burning and being torn to pieces by dogs.[63] The Spanish conquerors talked extensively of sodomy among the natives to depict them as savages and hence justify their conquest and forceful conversion to Christianity. As a result of the growing influence and power of the conquerors, many native cultures started condemning homosexual acts themselves.[citation needed]
+Among some of the indigenous peoples of the Americas in North America prior to European colonization, a relatively common form of same-sex sexuality centered around the figure of the Two-Spirit individual (the term itself was coined only in 1990). Typically, this individual was recognized early in life, given a choice by the parents to follow the path and, if the child accepted the role, raised in the appropriate manner, learning the customs of the gender it had chosen. Two-Spirit individuals were commonly shamans and were revered as having powers beyond those of ordinary shamans. Their sexual life was with the ordinary tribe members of the same sex.[citation needed]
+During the colonial times following the European invasion, homosexuality was prosecuted by the Inquisition, some times leading to death sentences on the charges of sodomy, and the practices became clandestine. Many homosexual individuals went into heterosexual marriages to keep appearances, and many turned to the clergy to escape public scrutiny of their lack of interest in the opposite sex.[citation needed]
+In 1986, the Supreme Court of the United States ruled in Bowers v. Hardwick that a state could criminalize sodomy, but, in 2003, overturned itself in Lawrence v. Texas and thereby legalized homosexual activity throughout the United States of America.
+Same-sex marriage in the United States expanded from one state in 2004 to all fifty states in 2015, through various state court rulings, state legislation, direct popular votes (referendums and initiatives), and federal court rulings.
+In East Asia, same-sex love has been referred to since the earliest recorded history.
+Homosexuality in China, known as the passions of the cut peach and various other euphemisms, has been recorded since approximately 600 BCE. Homosexuality was mentioned in many famous works of Chinese literature. The instances of same-sex affection and sexual interactions described in the classical novel Dream of the Red Chamber seem as familiar to observers in the present as do equivalent stories of romances between heterosexual people during the same period.  Confucianism, being primarily a social and political philosophy, focused little on sexuality, whether homosexual or heterosexual. Ming Dynasty literature, such as Bian Er Chai (弁而釵/弁而钗), portray homosexual relationships between men as more enjoyable and more "harmonious" than heterosexual relationships.[64] Writings from the Liu Song Dynasty by Wang Shunu claimed that homosexuality was as common as heterosexuality in the late 3rd century.[65]
+Opposition to homosexuality in China originates in the medieval Tang Dynasty (618–907), attributed to the rising influence of Christian and Islamic values,[66] but did not become fully established until the Westernization efforts of the late Qing Dynasty and the Republic of China.[67]
+The Laws of Manu mentions a "third sex", members of which may engage in nontraditional gender expression and homosexual activities.[68]
+The earliest Western documents (in the form of literary works, art objects, and mythographic materials) concerning same-sex relationships are derived from ancient Greece.
+In regard to male homosexuality, such documents depict an at times complex understanding in which relationships with women and relationships with adolescent boys could be a part of a normal man's love life. Same-sex relationships were a social institution variously constructed over time and from one city to another. The formal practice, an erotic yet often restrained relationship between a free adult male and a free adolescent, was valued for its pedagogic benefits and as a means of population control, though occasionally blamed for causing disorder. Plato praised its benefits in his early writings[69] but in his late works proposed its prohibition.[70] Aristotle, in the Politics, dismissed Plato's ideas about abolishing homosexuality (2.4); he explains that barbarians like the Celts accorded it a special honor (2.6.6), while the Cretans used it to regulate the population (2.7.5).[71]
+Little is known of female homosexuality in antiquity. Sappho, born on the island of Lesbos, was included by later Greeks in the canonical list of nine lyric poets. The adjectives deriving from her name and place of birth (Sapphic and Lesbian) came to be applied to female homosexuality beginning in the 19th century.[72][73] Sappho's poetry centers on passion and love for various personages and both genders. The narrators of many of her poems speak of infatuations and love (sometimes requited, sometimes not) for various females, but descriptions of physical acts between women are few and subject to debate.[74][75]
+In Ancient Rome, the young male body remained a focus of male sexual attention, but relationships were between older free men and slaves or freed youths who took the receptive role in sex. The Hellenophile emperor Hadrian is renowned for his relationship with Antinous, but the Christian emperor Theodosius I decreed a law on 6 August 390, condemning passive males to be burned at the stake. Notwithstanding these regulations taxes on brothels with boys available for homosexual sex continued to be collected until the end of the reign of Anastasius I in 518. Justinian, towards the end of his reign, expanded the proscription to the active partner as well (in 558), warning that such conduct can lead to the destruction of cities through the "wrath of God".
+During the Renaissance, wealthy cities in northern Italy—Florence and Venice in particular—were renowned for their widespread practice of same-sex love, engaged in by a considerable part of the male population and constructed along the classical pattern of Greece and Rome.[76][77] But even as many of the male population were engaging in same-sex relationships, the authorities, under the aegis of the Officers of the Night court, were prosecuting, fining, and imprisoning a good portion of that population.
+From the second half of the 13th century, death was the punishment for male homosexuality in most of Europe.[78]
+The relationships of socially prominent figures, such as King James I and the Duke of Buckingham, served to highlight the issue, including in anonymously authored street pamphlets: "The world is chang'd I know not how, For men Kiss Men, not Women now;...Of J. the First and Buckingham: He, true it is, his Wives Embraces fled, To slabber his lov'd Ganimede" (Mundus Foppensis, or The Fop Display'd, 1691).
+Love Letters Between a Certain Late Nobleman and the Famous Mr. Wilson was published in 1723 in England, and was presumed by some modern scholars to be a novel. The 1749 edition of John Cleland's popular novel Fanny Hill includes a homosexual scene, but this was removed in its 1750 edition. Also in 1749, the earliest extended and serious defense of homosexuality in English, Ancient and Modern Pederasty Investigated and Exemplified, written by Thomas Cannon, was published, but was suppressed almost immediately. It includes the passage, "Unnatural Desire is a Contradiction in Terms; downright Nonsense. Desire is an amatory Impulse of the inmost human Parts."[79] Around 1785 Jeremy Bentham wrote another defense, but this was not published until 1978.[80] Executions for sodomy continued in the Netherlands until 1803, and in England until 1835, James Pratt and John Smith being the last Englishmen to be so hanged.
+Between 1864 and 1880 Karl Heinrich Ulrichs published a series of twelve tracts, which he collectively titled Research on the Riddle of Man-Manly Love. In 1867, he became the first self-proclaimed homosexual person to speak out publicly in defense of homosexuality when he pleaded at the Congress of German Jurists in Munich for a resolution urging the repeal of anti-homosexual laws.[16] Sexual Inversion by Havelock Ellis, published in 1896, challenged theories that homosexuality was abnormal, as well as stereotypes, and insisted on the ubiquity of homosexuality and its association with intellectual and artistic achievement.[81]
+Although medical texts like these (written partly in Latin to obscure the sexual details) were not widely read by the general public, they did lead to the rise of Magnus Hirschfeld's Scientific-Humanitarian Committee, which campaigned from 1897 to 1933 against anti-sodomy laws in Germany, as well as a much more informal, unpublicized movement among British intellectuals and writers, led by such figures as Edward Carpenter and John Addington Symonds. Beginning in 1894 with Homogenic Love, Socialist activist and poet Edward Carpenter wrote a string of pro-homosexual articles and pamphlets, and "came out" in 1916 in his book My Days and Dreams. In 1900, Elisar von Kupffer published an anthology of homosexual literature from antiquity to his own time, Lieblingminne und Freundesliebe in der Weltliteratur.
+There are a handful of accounts by Arab travelers to Europe during the mid-1800s. Two of these travelers, Rifa'ah al-Tahtawi and Muhammad as-Saffar, show their surprise that the French sometimes deliberately mistranslated love poetry about a young boy, instead referring to a young female, to maintain their social norms and morals.[82]
+Israel is considered the most tolerant country in the Middle East and Asia to homosexuals,[83] with Tel Aviv being named "the gay capital of the Middle East"[84] and considered one of the most gay friendly cities in the world.[85] The annual Pride Parade in support of homosexuality takes place in Tel Aviv.[86]
+On the other hand, many governments in the Middle East often ignore, deny the existence of, or criminalize homosexuality. Homosexuality is illegal in almost all Muslim countries.[87] Same-sex intercourse officially carries the death penalty in several Muslim nations: Saudi Arabia, Iran, Mauritania, northern Nigeria, Sudan, and Yemen.[88]  Iranian President Mahmoud Ahmadinejad, during his 2007 speech at Columbia University, asserted that there were no gay people in Iran. However, the probable reason is that they keep their sexuality a secret for fear of government sanction or rejection by their families.[89]
+In ancient Sumer, a set of priests known as gala worked in the temples of the goddess Inanna, where they performed elegies and lamentations.[91]:285 Gala took female names, spoke in the eme-sal dialect, which was traditionally reserved for women, and appear to have engaged in homosexual intercourse.[92] The Sumerian sign for gala was a ligature of the signs for "penis" and "anus".[92] One Sumerian proverb reads: "When the gala wiped off his ass [he said], 'I must not arouse that which belongs to my mistress [i.e., Inanna].'"[92] In later Mesopotamian cultures, kurgarrū and assinnu were servants of the goddess Ishtar (Inanna's East Semitic equivalent), who dressed in female clothing and performed war dances in Ishtar's temples.[92] Several Akkadian proverbs seem to suggest that they may have also engaged in homosexual intercourse.[92]
+In ancient Assyria, homosexuality was present and common; it was also not prohibited, condemned, nor looked upon as immoral or disordered. Some religious texts contain prayers for divine blessings on homosexual relationships.[93][94] The Almanac of Incantations contained prayers favoring on an equal basis the love of a man for a woman, of a woman for a man, and of a man for man.[95]
+Some scholars argue that there are examples of homosexual love in ancient literature, like in the Mesopotamian Epic of Gilgamesh as well as in the Biblical story of David and Jonathan. In the Epic of Gilgamesh, the relationship between the main protagonist Gilgamesh and the character Enkidu has been seen by some to be homosexual in nature.[96][97][98][99] Similarly, David's love for Jonathan is "greater than the love of women."[100]
+In some societies of Melanesia, especially in Papua New Guinea, same-sex relationships were an integral part of the culture until the middle of the 1900s. The Etoro and Marind-anim for example, viewed heterosexuality as unclean and celebrated homosexuality instead. In some traditional Melanesian cultures a prepubertal boy would be paired with an older adolescent who would become his mentor and who would "inseminate" him (orally, anally, or topically, depending on the tribe) over a number of years in order for the younger to also reach puberty. Many Melanesian societies, however, have become hostile towards same-sex relationships since the introduction of Christianity by European missionaries.[101]
+The American Psychological Association, the American Psychiatric Association, and the National Association of Social Workers identify sexual orientation as "not merely a personal characteristic that can be defined in isolation. Rather, one's sexual orientation defines the universe of persons with whom one is likely to find the satisfying and fulfilling relationships":[3]
+Sexual orientation is commonly discussed as a characteristic of the individual, like biological sex, gender identity, or age. This perspective is incomplete because sexual orientation is always defined in relational terms and necessarily involves relationships with other individuals. Sexual acts and romantic attractions are categorized as homosexual or heterosexual according to the biological sex of the individuals involved in them, relative to each other. Indeed, it is by acting—or desiring to act—with another person that individuals express their heterosexuality, homosexuality, or bisexuality. This includes actions as simple as holding hands with or kissing another person. Thus, sexual orientation is integrally linked to the intimate personal relationships that human beings form with others to meet their deeply felt needs for love, attachment, and intimacy. In addition to sexual behavior, these bonds encompass nonsexual physical affection between partners, shared goals and values, mutual support, and ongoing commitment.[3]
+The Kinsey scale, also called the Heterosexual-Homosexual Rating Scale,[102] attempts to describe a person's sexual history or episodes of his or her sexual activity at a given time. It uses a scale from 0, meaning exclusively heterosexual, to 6, meaning exclusively homosexual. In both the Male and Female volumes of the Kinsey Reports, an additional grade, listed as "X", has been interpreted by scholars to indicate asexuality.[103]
+Often, sexual orientation and sexual identity are not distinguished, which can impact accurately assessing sexual identity and whether or not sexual orientation is able to change; sexual orientation identity can change throughout an individual's life, and may or may not align with biological sex, sexual behavior or actual sexual orientation.[104][105][106] Sexual orientation is stable and unlikely to change for the vast majority of people, but some research indicates that some people may experience change in their sexual orientation, and this is more likely for women than for men.[107] The American Psychological Association distinguishes between sexual orientation (an innate attraction) and sexual orientation identity (which may change at any point in a person's life).[108]
+People with a homosexual orientation can express their sexuality in a variety of ways, and may or may not express it in their behaviors.[2] Many have sexual relationships predominantly with people of their own sex, though some have sexual relationships with those of the opposite sex, bisexual relationships, or none at all (celibacy).[2] Studies have found same-sex and opposite-sex couples to be equivalent to each other in measures of satisfaction and commitment in relationships, that age and sex are more reliable than sexual orientation as a predictor of satisfaction and commitment to a relationship, and that people who are heterosexual or homosexual share comparable expectations and ideals with regard to romantic relationships.[109][110][111]
+Coming out (of the closet) is a phrase referring to one's disclosure of their sexual orientation or gender identity, and is described and experienced variously as a psychological process or journey.[113] Generally, coming out is described in three phases. The first phase is that of "knowing oneself", and the realization emerges that one is open to same-sex relations.[114] This is often described as an internal coming out. The second phase involves one's decision to come out to others, e.g. family, friends, or colleagues. The third phase more generally involves living openly as an LGBT person.[115] In the United States today, people often come out during high school or college age. At this age, they may not trust or ask for help from others, especially when their orientation is not accepted in society. Sometimes their own families are not even informed.
+According to Rosario, Schrimshaw, Hunter, Braun (2006), "the development of a lesbian, gay, or bisexual (LGB) sexual identity is a complex and often difficult process. Unlike members of other minority groups (e.g., ethnic and racial minorities), most LGB individuals are not raised in a community of similar others from whom they learn about their identity and who reinforce and support that identity. Rather, LGB individuals are often raised in communities that are either ignorant of or openly hostile toward homosexuality."[105]
+Outing is the practice of publicly revealing the sexual orientation of a closeted person.[116] Notable politicians, celebrities, military service people, and clergy members have been outed, with motives ranging from malice to political or moral beliefs. Many commentators oppose the practice altogether,[117] while some encourage outing public figures who use their positions of influence to harm other gay people.[118]
+Lesbians often experience their sexuality differently from gay men, and have different understandings about etiology from those derived from studies focused mostly on men.[citation needed]
+In a U.S.-based 1970s mail survey by Shere Hite, lesbians self-reported their reasons for being lesbian. This is the only major piece of research into female sexuality that has looked at how women understand being homosexual since Kinsey in 1953.[citation needed] The research yielded information about women's general understanding of lesbian relationships and their sexual orientation. Women gave various reasons for preferring sexual relations with women to sexual relations with men, including finding women more sensitive to other people's needs.[119]
+Since Hite carried out her study she has acknowledged that some women may have chosen the political identity of a lesbian. Julie Bindel, a UK journalist, reaffirmed that "political lesbianism continues to make intrinsic sense because it reinforces the idea that sexuality is a choice, and we are not destined to a particular fate because of our chromosomes." as recently as 2009.[120]
+Reliable data as to the size of the gay and lesbian population are of value in informing public policy.[123] For example, demographics would help in calculating the costs and benefits of domestic partnership benefits, of the impact of legalizing gay adoption, and of the impact of the U.S. military's Don't Ask Don't Tell policy.[123] Further, knowledge of the size of the "gay and lesbian population holds promise for helping social scientists understand a wide array of important questions—questions about the general nature of labor market choices, accumulation of human capital, specialization within households, discrimination, and decisions about geographic location."[123]
+Measuring the prevalence of homosexuality presents difficulties. It is necessary to consider the measuring criteria that are used, the cutoff point and the time span taken to define a sexual orientation.[16] Many people, despite having same-sex attractions, may be reluctant to identify themselves as gay or bisexual. The research must measure some characteristic that may or may not be defining of sexual orientation. The number of people with same-sex desires may be larger than the number of people who act on those desires, which in turn may be larger than the number of people who self-identify as gay, lesbian, or bisexual.[123]
+In 1948 and 1953, Alfred Kinsey reported that nearly 46% of the male subjects had "reacted" sexually to persons of both sexes in the course of their adult lives, and 37% had had at least one homosexual experience.[124][125] Kinsey's methodology was criticized by John Tukey for using convenience samples and not random samples.[126][127]
+A later study tried to eliminate the sample bias, but still reached similar conclusions.[128] Simon LeVay cites these Kinsey results as an example of the caution needed to interpret demographic studies, as they may give quite differing numbers depending on what criteria are used to conduct them, in spite of using sound scientific methods.[16]
+According to major studies, 2% to 11% of people have had some form of same-sex sexual contact within their lifetime;[129][130][131][132][133][134][135][136][137][138] this percentage rises to 16–21% when either or both same-sex attraction and behavior are reported.[138] In a 2006 study, 20% of respondents anonymously reported some homosexual feelings, although only 2–3% identified themselves as homosexual.[139] A 1992 study reported that 6.1% of males in Britain have had a homosexual experience, while in France the number was reported at 4.1%.[140]
+In the United States, according to a report by The Williams Institute in April 2011, 3.5% or approximately 9 million of the adult population identify as lesbian, gay, or bisexual.[141][142] According to the 2000 United States Census, there were about 601,209 same-sex unmarried partner households.[143]
+A 2013 study by the CDC, in which over 34,000 Americans were interviewed, puts the percentage of self-identifying lesbians and gay men at 1.6%, and of bisexuals at 0.7%.[144]
+According to a 2008 poll, 13% of Britons have had some form of same-sex sexual contact while only 6% of Britons identify themselves as either homosexual or bisexual.[145] In contrast, a survey by the UK Office for National Statistics (ONS) in 2010 found that 95% of Britons identified as heterosexual, 1.5% of Britons identified themselves as homosexual or bisexual, and the last 3.5% gave more vague answers such as "don't know", "other", or did not respond to the question.[125][146]
+In October 2012, Gallup started conducting annual surveys to study the demographics of LGBT people, determining that 3.4% (±1%) of adults identified as LGBT in the United States.[147] It was the nation's largest poll on the issue at the time.[148][149] In 2017, the percentage was estimated to have risen to 4.5% of adults, with the increase largely driven by Millennials. The poll attributes the rise to greater willingness of younger people to reveal their sexual identity.[150]
+The American Psychological Association, the American Psychiatric Association, and the National Association of Social Workers state:
+In 1952, when the American Psychiatric Association published its first Diagnostic and Statistical Manual of Mental Disorders, homosexuality was included as a disorder. Almost immediately, however, that classification began to be subjected to critical scrutiny in research funded by the National Institute of Mental Health. That study and subsequent research consistently failed to produce any empirical or scientific basis for regarding homosexuality as a disorder or abnormality, rather than a normal and healthy sexual orientation. As results from such research accumulated, professionals in medicine, mental health, and the behavioral and social sciences reached the conclusion that it was inaccurate to classify homosexuality as a mental disorder and that the DSM classification reflected untested assumptions based on once-prevalent social norms and clinical impressions from unrepresentative samples comprising patients seeking therapy and individuals whose conduct brought them into the criminal justice system.
+In recognition of the scientific evidence,[151] the American Psychiatric Association removed homosexuality from the DSM in 1973, stating that "homosexuality per se implies no impairment in judgment, stability, reliability, or general social or vocational capabilities." After thoroughly reviewing the scientific data, the American Psychological Association adopted the same position in 1975, and urged all mental health professionals "to take the lead in removing the stigma of mental illness that has long been associated with homosexual orientations." The National Association of Social Workers has adopted a similar policy.
+Thus, mental health professionals and researchers have long recognized that being homosexual poses no inherent obstacle to leading a happy, healthy, and productive life, and that the vast majority of gay and lesbian people function well in the full array of social institutions and interpersonal relationships.[3]
+The consensus of research and clinical literature demonstrates that same-sex sexual and romantic attractions, feelings, and behaviors are normal and positive variations of human sexuality.[152] There is now a large body of research evidence that indicates that being gay, lesbian or bisexual is compatible with normal mental health and social adjustment.[11] The World Health Organization's ICD-9 (1977) listed homosexuality as a mental illness; it was removed from the ICD-10, endorsed by the Forty-third World Health Assembly on 17 May 1990.[153][154][155] Like the DSM-II, the ICD-10 added ego-dystonic sexual orientation to the list, which refers to people who want to change their gender identities or sexual orientation because of a psychological or behavioral disorder (F66.1). The Chinese Society of Psychiatry removed homosexuality from its Chinese Classification of Mental Disorders in 2001 after five years of study by the association.[156] According to the Royal College of Psychiatrists "This unfortunate history demonstrates how marginalisation of a group of people who have a particular personality feature (in this case homosexuality) can lead to harmful medical practice and a basis for discrimination in society.[11] There is now a large body of research evidence that indicates that being gay, lesbian or bisexual is compatible with normal mental health and social adjustment. However, the experiences of discrimination in society and possible rejection by friends, families and others, such as employers, means that some LGB people experience a greater than expected prevalence of mental health difficulties and substance misuse problems. Although there have been claims by conservative political groups in the USA that this higher prevalence of mental health difficulties is confirmation that homosexuality is itself a mental disorder, there is no evidence whatever to substantiate such a claim."[157]
+Most lesbian, gay, and bisexual people who seek psychotherapy do so for the same reasons as heterosexual people (stress, relationship difficulties, difficulty adjusting to social or work situations, etc.); their sexual orientation may be of primary, incidental, or no importance to their issues and treatment. Whatever the issue, there is a high risk for anti-gay bias in psychotherapy with lesbian, gay, and bisexual clients.[158] Psychological research in this area has been relevant to counteracting prejudicial ("homophobic") attitudes and actions, and to the LGBT rights movement generally.[159]
+The appropriate application of affirmative psychotherapy is based on the following scientific facts:[152]
+Although scientists favor biological models for the cause of sexual orientation,[4] they do not believe that the development of sexual orientation is the result of any one factor. They generally believe that it is determined by a complex interplay of biological and environmental factors, and is shaped at an early age.[2][5][6] There is considerably more evidence supporting nonsocial, biological causes of sexual orientation than social ones, especially for males.[8] There is no substantive evidence which suggests parenting or early childhood experiences play a role with regard to sexual orientation.[11] Scientists do not believe that sexual orientation is a choice.[4][5][7]
+The American Academy of Pediatrics stated in Pediatrics in 2004:
+There is no scientific evidence that abnormal parenting, sexual abuse, or other adverse life events influence sexual orientation. Current knowledge suggests that sexual orientation is usually established during early childhood.[4][160]
+The American Psychological Association, American Psychiatric Association, and National Association of Social Workers stated in 2006:
+Currently, there is no scientific consensus about the specific factors that cause an individual to become heterosexual, homosexual, or bisexual—including possible biological, psychological, or social effects of the parents' sexual orientation. However, the available evidence indicates that the vast majority of lesbian and gay adults were raised by heterosexual parents and the vast majority of children raised by lesbian and gay parents eventually grow up to be heterosexual.[2]
+Despite numerous attempts, no "gay gene" has been identified. However, there is substantial evidence for a genetic basis of homosexuality, especially in males, based on twin studies; some association with regions of Chromosome 8, the Xq28 locus on the X chromosome, and other sites across many chromosomes.[161]
+Sanders et al. 2015
+Sanders et al. 2015
+Starting in the 2010s, potential epigenetic factors have become a topic of increased attention in genetic research on sexual orientation. A study presented at the ASHG 2015 Annual Meeting found that the methylation pattern in nine regions of the genome appeared very closely linked to sexual orientation, with a resulting algorithm using the methylation pattern to predict the sexual orientation of a control group with almost 70% accuracy.[164][165]
+Research into the causes of homosexuality plays a role in political and social debates and also raises concerns about genetic profiling and prenatal testing.[166][167]
+Since homosexuality tends to lower reproductive success, and since there is considerable evidence that human sexual orientation is genetically influenced, it is unclear how it is maintained in the population at a relatively high frequency.[168] There are many possible explanations, such as genes predisposing to homosexuality also conferring advantage in heterosexuals, a kin selection effect, social prestige, and more.[169] A 2009 study also suggested a significant increase in fecundity in the females related to the homosexual people from the maternal line (but not in those related from the paternal one).[170]
+There are no studies of adequate scientific rigor that conclude that sexual orientation change efforts work to change a person's sexual orientation. Those efforts have been controversial due to tensions between the values held by some faith-based organizations, on the one hand, and those held by LGBT rights organizations and professional and scientific organizations and other faith-based organizations, on the other.[14] The longstanding consensus of the behavioral and social sciences and the health and mental health professions is that homosexuality per se is a normal and positive variation of human sexual orientation, and therefore not a mental disorder.[14] The American Psychological Association says that "most people experience little or no sense of choice about their sexual orientation".[171]
+Some individuals and groups have promoted the idea of homosexuality as symptomatic of developmental defects or spiritual and moral failings and have argued that sexual orientation change efforts, including psychotherapy and religious efforts, could alter homosexual feelings and behaviors. Many of these individuals and groups appeared to be embedded within the larger context of conservative religious political movements that have supported the stigmatization of homosexuality on political or religious grounds.[14]
+No major mental health professional organization has sanctioned efforts to change sexual orientation and virtually all of them have adopted policy statements cautioning the profession and the public about treatments that purport to change sexual orientation. These include the American Psychiatric Association, American Psychological Association, American Counseling Association, National Association of Social Workers in the US,[172] the Royal College of Psychiatrists,[173] and the Australian Psychological Society.[174] The American Psychological Association and the Royal College of Psychiatrists expressed concerns that the positions espoused by NARTH are not supported by the science and create an environment in which prejudice and discrimination can flourish.[173][175]
+The American Psychological Association states that "sexual orientation is not a choice that can be changed at will, and that sexual orientation is most likely the result of a complex interaction of environmental, cognitive and biological factors...is shaped at an early age...[and evidence suggests] biological, including genetic or inborn hormonal factors, play a significant role in a person's sexuality."[5] They say that "sexual orientation identity—not sexual orientation—appears to change via psychotherapy, support groups, and life events."[14] The American Psychiatric Association says "individuals maybe become aware at different points in their lives that they are heterosexual, gay, lesbian, or bisexual" and "opposes any psychiatric treatment, such as 'reparative' or 'conversion' therapy, which is based upon the assumption that homosexuality per se is a mental disorder, or based upon a prior assumption that the patient should change his/her homosexual orientation". They do, however, encourage gay affirmative psychotherapy.[176] Similarly, the American *Psychological* Association[177] is doubtful about the effectiveness and side-effect profile of sexual orientation change efforts, including conversion therapy.
+The American Psychological Association "encourages mental health professionals to avoid misrepresenting the efficacy of sexual orientation change efforts by promoting or promising change in sexual orientation when providing assistance to individuals distressed by their own or others' sexual orientation and concludes that the benefits reported by participants in sexual orientation change efforts can be gained through approaches that do not attempt to change sexual orientation".[14]
+Scientific research has been generally consistent in showing that lesbian and gay parents are as fit and capable as heterosexual parents, and their children are as psychologically healthy and well-adjusted as children reared by heterosexual parents.[178][179][180] According to scientific literature reviews, there is no evidence to the contrary.[3][181][182][183][184]
+A 2001 review suggested that the children with lesbian or gay parents appear less traditionally gender-typed and are more likely to be open to homoerotic relationships, partly due to genetic (80% of the children being raised by same-sex couples in the US are not adopted and most are the result of previous heterosexual marriages.[185]) and family socialization processes (children grow up in relatively more tolerant school, neighborhood, and social contexts, which are less heterosexist), even though majority of children raised by same-sex couples identify as heterosexual.[186] A 2005 review by Charlotte J. Patterson for the American Psychological Association found that the available data did not suggest higher rates of homosexuality among the children of lesbian or gay parents.[187]
+The terms "men who have sex with men" (MSM) and "women who have sex with women" (WSW) refer to people who engage in sexual activity with others of the same sex regardless of how they identify themselves—as many choose not to accept social identities as lesbian, gay and bisexual.[188][189][190][191][192] These terms are often used in medical literature and social research to describe such groups for study, without needing to consider the issues of sexual self-identity. The terms are seen as problematic by some, however, because they "obscure social dimensions of sexuality; undermine the self-labeling of lesbian, gay, and bisexual people; and do not sufficiently describe variations in sexual behavior".[193]
+In contrast to its benefits, sexual behavior can be a disease vector. Safe sex is a relevant harm reduction philosophy.[194] Many countries currently prohibit men who have sex with men from donating blood; the policy of the United States Food and Drug Administration states that "they are, as a group, at increased risk for HIV, hepatitis B and certain other infections that can be transmitted by transfusion."[195]
+These safer sex recommendations are agreed upon by public health officials for women who have sex with women to avoid sexually transmitted infections (STIs):
+These safer sex recommendations are agreed upon by public health officials for men who have sex with men to avoid sexually transmitted infections:
+When it was first described in medical literature, homosexuality was often approached from a view that sought to find an inherent psychopathology as its root cause. Much literature on mental health and homosexual patients centered on their depression, substance abuse, and suicide. Although these issues exist among people who are non-heterosexual, discussion about their causes shifted after homosexuality was removed from the Diagnostic and Statistical Manual (DSM) in 1973. Instead, social ostracism, legal discrimination, internalization of negative stereotypes, and limited support structures indicate factors homosexual people face in Western societies that often adversely affect their mental health.[199]
+Stigma, prejudice, and discrimination stemming from negative societal attitudes toward homosexuality lead to a higher prevalence of mental health disorders among lesbians, gay men, and bisexuals compared to their heterosexual peers.[200] Evidence indicates that the liberalization of these attitudes over the 1990s through the 2010s is associated with a decrease in such mental health risks among younger LGBT people.[201]
+Gay and lesbian youth bear an increased risk of suicide, substance abuse, school problems, and isolation because of a "hostile and condemning environment, verbal and physical abuse, rejection and isolation from family and peers".[202] Further, LGBT youths are more likely to report psychological and physical abuse by parents or caretakers, and more sexual abuse. Suggested reasons for this disparity are that (1) LGBT youths may be specifically targeted on the basis of their perceived sexual orientation or gender non-conforming appearance, and (2) that "risk factors associated with sexual minority status, including discrimination, invisibility, and rejection by family members...may lead to an increase in behaviors that are associated with risk for victimization, such as substance abuse, sex with multiple partners, or running away from home as a teenager."[203] A 2008 study showed a correlation between the degree of rejecting behavior by parents of LGB adolescents and negative health problems in the teenagers studied:
+Higher rates of family rejection were significantly associated with poorer health outcomes. On the basis of odds ratios, lesbian, gay, and bisexual young adults who reported higher levels of family rejection during adolescence were 8.4 times more likely to report having attempted suicide, 5.9 times more likely to report high levels of depression, 3.4 times more likely to use illegal drugs, and 3.4 times more likely to report having engaged in unprotected sexual intercourse compared with peers from families that reported no or low levels of family rejection.[204]
+Crisis centers in larger cities and information sites on the Internet have arisen to help youth and adults.[205] The Trevor Project, a suicide prevention helpline for gay youth, was established following the 1998 airing on HBO of the Academy Award winning short film Trevor.[206]
+Most nations do not prohibit consensual sex between unrelated persons above the local age of consent. Some jurisdictions further recognize identical rights, protections, and privileges for the family structures of same-sex couples, including marriage. Some countries and jurisdictions mandate that all individuals restrict themselves to heterosexual activity and disallow homosexual activity via sodomy laws. Offenders can face the death penalty in Islamic countries and jurisdictions ruled by sharia. There are, however, often significant differences between official policy and real-world enforcement.
+Although homosexual acts were decriminalized in some parts of the Western world, such as Poland in 1932, Denmark in 1933, Sweden in 1944, and the England and Wales in 1967, it was not until the mid-1970s that the gay community first began to achieve limited civil rights in some developed countries. A turning point was reached in 1973 when the American Psychiatric Association, which previously listed homosexuality in the DSM-I in 1952, removed homosexuality in the DSM-II, in recognition of scientific evidence.[3] In 1977, Quebec became the first state-level jurisdiction in the world to prohibit discrimination on the grounds of sexual orientation. During the 1980s and 1990s, most developed countries enacted laws decriminalizing homosexual behavior and prohibiting discrimination against lesbian and gay people in employment, housing, and services. On the other hand, many countries today in the Middle East and Africa, as well as several countries in Asia, the Caribbean and the South Pacific, outlaw homosexuality. In 2013, the Supreme Court of India upheld Section 377 of the Indian Penal Code,[207] but in 2018 overturned itself and legalized homosexual activity throughout India.[208] The Section 377, a British colonial-era law which criminalizes homosexual activity, remains in effect in more than 40 former British colonies.[209][210]
+10 countries or jurisdictions, all of which are Islamic and ruled by sharia, impose the death penalty for homosexuality.
+In the European Union, discrimination of any type based on sexual orientation or gender identity is illegal under the Charter of Fundamental Rights of the European Union.[220]
+Since the 1960s, many LGBT people in the West, particularly those in major metropolitan areas, have developed a so-called gay culture. To many,[who?] gay culture is exemplified by the gay pride movement, with annual parades and displays of rainbow flags. Yet not all LGBT people choose to participate in "queer culture", and many gay men and women specifically decline to do so. To some[who?] it seems to be a frivolous display, perpetuating gay stereotypes.
+With the outbreak of AIDS in the early 1980s, many LGBT groups and individuals organized campaigns to promote efforts in AIDS education, prevention, research, patient support, and community outreach, as well as to demand government support for these programs.
+The death toll wrought by the AIDS epidemic at first seemed to slow the progress of the gay rights movement, but in time it galvanized some parts of the LGBT community into community service and political action, and challenged the heterosexual community to respond compassionately. Major American motion pictures from this period that dramatized the response of individuals and communities to the AIDS crisis include An Early Frost (1985), Longtime Companion (1990), And the Band Played On (1993), Philadelphia (1993), and Common Threads: Stories from the Quilt (1989).
+Publicly gay politicians have attained numerous government posts, even in countries that had sodomy laws in their recent past. Examples include Guido Westerwelle, Germany's Vice-Chancellor; Peter Mandelson, a British Labour Party cabinet minister and Per-Kristian Foss, formerly Norwegian Minister of Finance.
+LGBT movements are opposed by a variety of individuals and organizations. Some social conservatives believe that all sexual relationships with people other than an opposite-sex spouse undermine the traditional family[221] and that children should be reared in homes with both a father and a mother.[222][223] Some argue that gay rights may conflict with individuals' freedom of speech,[224][225] religious freedoms in the workplace,[226][227] the ability to run churches,[228] charitable organizations[229][230] and other religious organizations[231] in accordance with one's religious views, and that the acceptance of homosexual relationships by religious organizations might be forced through threatening to remove the tax-exempt status of churches whose views do not align with those of the government.[232][233][234][235] Some critics charge that political correctness has led to the association of sex between males and HIV being downplayed.[236]
+Policies and attitudes toward gay and lesbian military personnel vary widely around the world. Some countries allow gay men, lesbians, and bisexual people to serve openly and have granted them the same rights and privileges as their heterosexual counterparts. Many countries neither ban nor support LGB service members. A few countries continue to ban homosexual personnel outright.
+Most Western military forces have removed policies excluding sexual minority members. Of the 26 countries that participate militarily in NATO, more than 20 permit openly gay, lesbian and bisexual people to serve. Of the permanent members of the United Nations Security Council, three (United Kingdom, France and United States) do so. The other two generally do not: China bans gay and lesbian people outright, Russia excludes all gay and lesbian people during peacetime but allows some gay men to serve in wartime (see below). Israel is the only country in the Middle East region that allows openly LGB people to serve in the military.
+While the question of homosexuality in the military has been highly politicized in the United States, it is not necessarily so in many countries. Generally speaking, sexuality in these cultures is considered a more personal aspect of one's identity than it is in the United States.
+According to the American Psychological Association, empirical evidence fails to show that sexual orientation is germane to any aspect of military effectiveness including unit cohesion, morale, recruitment and retention.[237] Sexual orientation is irrelevant to task cohesion, the only type of cohesion that critically predicts the team's military readiness and success.[238]
+Societal acceptance of non-heterosexual orientations such as homosexuality is lowest in Asian, African and Eastern European countries,[239][240] and is highest in Western Europe, Australia, and the Americas. Western society has become increasingly accepting of homosexuality since the 1990s. In 2017, Professor Amy Adamczyk contended that these cross-national differences in acceptance can be largely explained by three factors: the relative strength of democratic institutions, the level of economic development, and the religious context of the places where people live.[241]
+In 2006, the American Psychological Association, American Psychiatric Association and National Association of Social Workers stated in an amicus brief presented to the Supreme Court of California: "Gay men and lesbians form stable, committed relationships that are equivalent to heterosexual relationships in essential respects. The institution of marriage offers social, psychological, and health benefits that are denied to same-sex couples. By denying same-sex couples the right to marry, the state reinforces and perpetuates the stigma historically associated with homosexuality. Homosexuality remains stigmatized, and this stigma has negative consequences. California's prohibition on marriage for same-sex couples reflects and reinforces this stigma". They concluded: "There is no scientific basis for distinguishing between same-sex couples and heterosexual couples with respect to the legal rights, obligations, benefits, and burdens conferred by civil marriage."[3]
+Though the relationship between homosexuality and religion is complex, current authoritative bodies and doctrines of the world's largest religions view homosexual behaviour negatively.[citation needed] This can range from quietly discouraging homosexual activity, to explicitly forbidding same-sex sexual practices among adherents and actively opposing social acceptance of homosexuality. Some teach that homosexual desire itself is sinful,[242] others state that only the sexual act is a sin,[243] others are completely accepting of gays and lesbians,[244] while some encourage homosexuality.[245] Some claim that homosexuality can be overcome through religious faith and practice. On the other hand, voices exist within many of these religions that view homosexuality more positively, and liberal religious denominations may bless same-sex marriages. Some view same-sex love and sexuality as sacred, and a mythology of same-sex love can be found around the world.[246]
+Gay bullying can be the verbal or physical abuse against a person who is perceived by the aggressor to be  lesbian, gay,  bisexual or transgender, including persons who are actually heterosexual or of non-specific or unknown sexual orientation.  In the US, teenage students heard anti-gay slurs such as "homo", "faggot" and "sissy" about 26 times a day on average, or once every 14 minutes, according to a 1998 study by Mental Health America (formerly National Mental Health Association).[247]
+In many cultures, homosexual people are frequently subject to prejudice and discrimination. A 2011 Dutch study concluded that 49% of Holland's youth and 58% of youth foreign to the country reject homosexuality.[248]  Similar to other minority groups they can also be subject to stereotyping. These attitudes tend to be due to forms of homophobia and heterosexism (negative attitudes, bias, and discrimination in favor of opposite-sex sexuality and relationships). Heterosexism can include the presumption that everyone is heterosexual or that opposite-sex attractions and relationships are the norm and therefore superior. Homophobia is a fear of, aversion to, or discrimination against homosexual people. It manifests in different forms, and a number of different types have been postulated, among which are internalized homophobia, social homophobia, emotional homophobia, rationalized homophobia, and others.[249] Similar is lesbophobia (specifically targeting lesbians) and biphobia (against bisexual people). When such attitudes manifest as crimes they are often called hate crimes and gay bashing.
+Negative stereotypes characterize LGB people as less romantically stable, more promiscuous and more likely to abuse children, but there is no scientific basis to such assertions. Gay men and lesbians form stable, committed relationships that are equivalent to heterosexual relationships in essential respects.[3] Sexual orientation does not affect the likelihood that people will abuse children.[250][251][252] Claims that there is scientific evidence to support an association between being gay and being a pedophile are based on misuses of those terms and misrepresentation of the actual evidence.[251]
+In the United States, the FBI reported that 20.4% of hate crimes reported to law enforcement in 2011 were based on sexual orientation bias. 56.7% of these crimes were based on bias against homosexual men. 11.1% were based on bias against homosexual women. 29.6% were based on anti-homosexual bias without regard to gender.[253] The 1998 murder of Matthew Shepard, a gay student, is a notorious such incident in the U.S. LGBT people, especially lesbians, may become the victims of "corrective rape", a violent crime with the supposed aim of making them heterosexual. In certain parts of the world, LGBT people are also at risk of "honor killings" perpetrated by their families or relatives.[254][255][256]
+In Morocco, a constitutional monarchy following Islamic laws, homosexual acts are a punishable offence. With a population hostile towards LGBT people, the country has witnessed public demonstrations against homosexuals, public denunciations of presumed homosexual individuals, as well as violent intrusions in private homes. The community in the country is exposed to additional risk of prejudice, social rejection and violence, with a greater impossibility of obtaining protection even from the police.[257]
+In 2017, Abderrahim El Habachi became one of the many homosexual Moroccans to flee the country for fear, and sought shelter in the United Kingdom. In August 2019, he accused the UK Home Office of not taking LGBT migrants seriously. He highlighted that they are being put through tough times because the Home Office don't believe their sexual orientation or in the struggles they face in their countries.[258]
+Homosexual and bisexual behaviors occur in a number of other animal species. Such behaviors include sexual activity, courtship, affection, pair bonding, and parenting,[20] and are widespread; a 1999 review by researcher Bruce Bagemihl shows that homosexual behavior has been documented in about 500 species, ranging from primates to gut worms.[20][21] Animal sexual behavior takes many different forms, even within the same species. The motivations for and implications of these behaviors have yet to be fully understood, since most species have yet to be fully studied.[260] According to Bagemihl, "the animal kingdom [does] it with much greater sexual diversity—including homosexual, bisexual and nonreproductive sex—than the scientific community and society at large have previously been willing to accept".[261]
+A review paper by N. W. Bailey and Marlene Zuk looking into studies of same-sex sexual behaviour in animals challenges the view that such behaviour lowers reproductive success, citing several hypotheses about how same-sex sexual behavior might be adaptive; these hypotheses vary greatly among different species. Bailey and Zuk also suggest future research needs to look into evolutionary consequences of same-sex sexual behaviour, rather than only looking into origins of such behaviour.[262]
+Category:LGBT culture

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+The Aegean Sea[a] is an elongated embayment of the Mediterranean Sea located between the Greek and Anatolian peninsulas. The sea has an area of some 215,000 square kilometres.[2] In the north, the Aegean is connected to the Marmara Sea and the Black Sea by the straits of the Dardanelles and Bosphorus. The Aegean Islands are located within the sea and some bound it on its southern periphery, including Crete and Rhodes. The sea reaches a maximum depth of 3,544 meters, to the east of Crete.
+The Aegean Islands can be divided into several island groups, including Dodecanese, the Cyclades, the Sporades, the Saronic islands and the North Aegean Islands, as well as Crete and its surrounding islands. The Dodecanese, located to the southeast, includes the islands of Rhodes, Kos, and Patmos; the islands of Delos and Naxos are within the Cyclades to the south of the sea. Lesbos is part of the North Aegean Islands. Euboea, the second largest island in Greece, is located in the Aegean, despite being administered as part of Central Greece. Nine out of twelve of the Administrative regions of Greece border the sea, along with the Turkish provinces of Edirne, Canakkale, Balıkesir, Izmir, Aydın and Muğla to the east of the sea. Various Turkish islands in the sea are Imbros, Tenedos, Cunda Island, and the Foça Islands.
+The Aegean Sea has been historically important, especially in regards to the civilization of Ancient Greece, who inhabited the area around the coast of the Aegean and the Aegean islands. The Aegean islands facilitated contact between the people of the area and between Europe and Asia. Along with the Greeks, Thracians lived among the northern coast. The Romans conquered the area under the Roman Empire, and later the Byzantine Empire held it against advances by the First Bulgarian Empire. The Fourth Crusade weakened Byzantine control of the area, and it was eventually conquered by the Ottoman Empire, with the exception of Crete, which was a Venetian colony until 1669. The Greek War of Independence allowed a Greek state on the coast of the Aegean from 1829 onwards. The Ottoman Empire held a presence over the sea for over 500 years, until it was replaced by modern Turkey.
+The rocks making up the floor of the Aegean are mainly limestone, though often greatly altered by volcanic activity that has convulsed the region in relatively recent geologic times. Of particular interest are the richly coloured sediments in the region of the islands of Santorini and Milos, in the south Aegean.[2] Notable cities on the Aegean coastline include Athens, Thessaloniki, Volos, Kavala and Heraklion in Greece, and İzmir and Bodrum in Turkey.
+A number of issues concerning sovereignty within the Aegean Sea are disputed between Greece and Turkey. The Aegean dispute has had a large effect on Greek-Turkish relations since the 1970s. Issues include the delimitation of territorial waters, national airspace, exclusive economic zones and flight information regions.[3]
+Late Latin authors referred the name Aegaeus to Aegeus, whom they said to have jumped into that sea (instead of jumping from the Athenian acropolis, as told by some Greek authors). He was the father of Theseus, the mythical king and founder-hero of Athens. Aegeus had told Theseus to put up white sails when returning if he was successful in killing the Minotaur. When Theseus returned, he forgot these instructions, and Aegeus thinking his son to have died then drowned himself in the sea.[4]
+The sea was known in Latin as Aegaeum mare under the control of the Roman Empire. The Venetians, who ruled many Greek islands in the High and Late Middle Ages, popularized the name Archipelago (Greek: αρχιπέλαγος, meaning "main sea" or "chief sea"), a name that held on in many European countries until the early modern period. In the South Slavic languages, the Aegean is called White Sea (Bulgarian: Бяло море/Byalo more; Macedonian: Belo more/Бело море; Serbo-Croatian: Belo more/Бело море).[5] The Turkish name for the sea is Ege Denizi, derived from the Greek name.
+The Aegean Sea is an elongated embayment of the Mediterranean Sea, and covers about 214,000 square kilometres (83,000 sq mi) in area, measuring about 670 kilometres (420 mi) longitudinally and 390 kilometres (240 mi) latitudinal. The sea's maximum depth is 3,543 metres (11,624 ft), located at a point east of Crete. The Aegean Islands are found within its waters, with the following islands delimiting the sea on the south, generally from west to east: Kythera, Antikythera, Crete, Kasos, Karpathos and Rhodes. The Anatolian peninsula marks the eastern boundary of the sea, while the Greek mainland marks the west. Several seas are contained within the Aegean Sea; the Thracian Sea is a section of the Aegean located to the north, the Icarian Sea to the east, the Myrtoan Sea to the west, while the Sea of Crete is the southern section.
+The Greek regions that border the sea, in alphabetical order, are Attica, Central Greece, Central Macedonia, Crete, Eastern Macedonia and Thrace, North Aegean, Peloponnese, South Aegean, and Thessaly. The historical region of Macedonia also borders the sea, to the north.
+The Aegean Islands, which almost all belong to Greece, can be divided into seven groups:
+Many of the Aegean islands or island chains, are geographically extensions of the mountains on the mainland. One chain extends across the sea to Chios, another extends across Euboea to Samos, and a third extends across the Peloponnese and Crete to Rhodes, dividing the Aegean from the Mediterranean.
+The bays and gulfs of the Aegean beginning at the South and moving clockwise include on Crete, the Mirabello, Almyros, Souda and Chania bays or gulfs, on the mainland the Myrtoan Sea to the west with the Argolic Gulf, the Saronic Gulf northwestward, the Petalies Gulf which connects with the South Euboic Sea, the Pagasetic Gulf which connects with the North Euboic Sea, the Thermian Gulf northwestward, the Chalkidiki Peninsula including the Cassandra and the Singitic Gulfs, northward the Strymonian Gulf and the Gulf of Kavala and the rest are in Turkey; Saros Gulf, Edremit Gulf, Dikili Gulf, Gulf of Çandarlı, Gulf of İzmir, Gulf of Kuşadası, Gulf of Gökova, Güllük Gulf.
+The Aegean sea is connected to the Sea of Marmara by the Dardanelles, also known from Classical Antiquity as the Hellespont. The Dardanelles are located to the northeast of the sea. It ultimately connects with the Black Sea through the Bosphoros strait, upon which lies the city of Istanbul. The Dardanelles and the Bosphoros are known as the Turkish Straits.
+According to the International Hydrographic Organization, the limits of the Aegean Sea as follows:[8]
+Aegean surface water circulates in a counterclockwise gyre, with hypersaline Mediterranean water moving northward along the west coast of Turkey, before being displaced by less dense Black Sea outflow. The dense Mediterranean water sinks below the Black Sea inflow to a depth of 23–30 metres (75–98 ft), then flows through the Dardanelles Strait and into the Sea of Marmara at velocities of 5–15 cm/s (2–6 in/s). The Black Sea outflow moves westward along the northern Aegean Sea, then flows southwards along the east coast of Greece.[9]
+The physical oceanography of the Aegean Sea is controlled mainly by the regional climate, the fresh water discharge from major rivers draining southeastern Europe, and the seasonal variations in the Black Sea surface water outflow through the Dardanelles Strait.
+Analysis[10] of the Aegean during 1991 and 1992 revealed three distinct water masses:
+The climate of the Aegean Sea largely reflects the climate of Greece and Western Turkey, which is to say, predominately Mediterranean. According to the Köppen climate classification, most of the Aegean is classified as Hot-summer Mediterranean (Csa), with hotter and drier summers along with milder and wetter winters. However, high temperatures during summers are generally not quite as high as those in arid or semiarid climates due to the presence of a large body of water. This is most predominant in the west and east coasts of the Aegean, and within the Aegean islands. In the north of the Aegean Sea, the climate is instead classified as Cold semi-arid (BSk), which feature cooler summers than Hot-summer Mediterranean climates.
+The Etesian winds are a dominant weather influence in the Aegean Basin.
+The below table lists climate conditions of some major Aegean cities:
+Numerous Greek and Turkish settlements are located along their mainland coast, as well as on towns on the Aegean islands. The largest cities are Athens and Thessaloniki in Greece and İzmir in Turkey. The most populated of the Aegean islands is Crete, followed by Euboea and Rhodes.
+Athens
+Thessaloniki
+İzmir
+Heraklion
+Greece has established several marine protected areas along its coasts. According to the Network of Managers of Marine Protected Areas in the Mediterranean (MedPAN), four Greek MPAs are participating in the Network. These include Alonnisos Marine Park, while the Missolonghi–Aitoliko Lagoons and the island of Zakynthosare not on the Aegean.[13]
+The current coastline dates back to about 4000 BC. Before that time, at the peak of the last ice age (about 18,000 years ago) sea levels everywhere were 130 metres lower, and there were large well-watered coastal plains instead of much of the northern Aegean. When they were first occupied, the present-day islands including Milos with its important obsidian production were probably still connected to the mainland. The present coastal arrangement appeared around 9,000 years ago, with post-ice age sea levels continuing to rise for another 3,000 years after that.[14]
+The subsequent Bronze Age civilizations of Greece and the Aegean Sea have given rise to the general term Aegean civilization. In ancient times, the sea was the birthplace of two ancient civilizations – the Minoans of Crete and the Myceneans of the Peloponnese.[15]
+The Minoan civilization was a Bronze Age civilization on the island of Crete and other Aegean islands, flourishing from around 2700 to 1450 BC before a period of decline, finally ending at around 1100 BC. It represented the first advanced civilization in Europe, leaving behind massive building complexes, tools, stunning artwork, writing systems, and a massive network of trade.[16] The Minoan period saw extensive trade between Crete, Aegean, and Mediterranean settlements, particularly the Near East. The most notable Minoan palace is that of Knossos, followed by that of Phaistos.
+After the decline of the Minoan civilization, the Mycenaean Greeks arose, becoming the first advanced civilization in mainland Greece, which lasted from approximately 1600 to 1100 BC. It is believed that the site of Mycenae, which sits close to the Aegean coast, was the center of Mycenaean civilization. The Mycenaeans introduced several innovations in the fields of engineering, architecture and military infrastructure, while trade over vast areas of the Mediterranean, including the Aegean, was essential for the Mycenaean economy. Their syllabic script, the Linear B, offers the first written records of the Greek language and their religion already included several deities that can also be found in the Olympic Pantheon. Mycenaean Greece was dominated by a warrior elite society and consisted of a network of palace-centered states that developed rigid hierarchical, political, social and economic systems. At the head of this society was the king, known as wanax.
+The civilization of Mycenaean Greeks perished with the collapse of Bronze Age culture in the eastern Mediterranean, to be followed by the so-called Greek Dark Ages. It is undetermined what cause the collapse of the Mycenaeans. During the Greek Dark Ages, writing in the Linear B script ceased, vital trade links were lost, and towns and villages were abandoned.
+The Archaic period followed the Greek Dark Ages in the 8th century BC. Greece became divided into small self-governing communities, and adopted the Phoenician alphabet, modifying it to create the Greek alphabet. By the 6th century BC several cities had emerged as dominant in Greek affairs: Athens, Sparta, Corinth, and Thebes, of which Athens, Sparta, and Corinth were closest to the Aegean Sea. Each of them had brought the surrounding rural areas and smaller towns under their control, and Athens and Corinth had become major maritime and mercantile powers as well. In the 8th and 7th centuries BC many Greeks emigrated to form colonies in Magna Graecia (Southern Italy and Sicily), Asia Minor and further afield.
+The Aegean Sea would later come to be under the control, albeit briefly, of the Kingdom of Macedonia. Philip II and his son Alexander the Great led a series of conquests that led not only to the unification of the Greek mainland and the control of the Aegean Sea under his rule, but also the destruction of the Achaemenid Empire. After Alexander the Great's death, his empire was divided among his generals. Cassander became king of the Hellenistic kingdom of Macedon, which held territory along the western coast of the Aegean, roughly corresponding to modern-day Greece. The Kingdom of Lysimachus had control over the sea's eastern coast. Greece had entered the Hellenistic period.
+The Macedonian Wars were a series of conflicts fought by the Roman Republic and its Greek allies in the eastern Mediterranean against several different major Greek kingdoms. They resulted in Roman control or influence over the eastern Mediterranean basin, including the Aegean, in addition to their hegemony in the western Mediterranean after the Punic Wars. During Roman rule, the land around the Aegean Sea fell under the provinces of Achaea, Macedonia, Thracia, Asia and Creta et Cyrenica (island of Crete)
+The Fall of the Western Roman Empire allowed its successor state, the Byzantine Empire, to continue Roman control over the Aegean Sea. However, their territory would later be threatened by the Early Muslim conquests initiated by Muhammad in the 7th century. Although the Rashidun Caliphate did not manage to obtain land along the cost of the Aegean sea, its conquest of the Eastern Anatolian peninsula as well as Egypt, the Levant, and North Africa left the Byzantine Empire weakened. The Umayyad Caliphate expanded the territorial gains of the Rashidun Caliphate, conquering much of North Africa, and threatened the Byzantine Empire's control of Western Anatolia, where it meets the Aegean Sea.
+During the 820s, Crete was conquered by a group of Berbers Andalusians exiles led by Abu Hafs Umar al-Iqritishi, and it became an independent Islamic state. The Byzantine Empire launched a campaign that took most of the island back in 842 and 843 under Theoktistos, but the reconquest was not completed and was soon reversed. Later attempts by the Byzantine Empire to recover the island were without success. For the approximately 135 years of its existence, the emirate of Crete was one of the major foes of Byzantium. Crete commanded the sea lanes of the Eastern Mediterranean and functioned as a forward base and haven for Muslim corsair fleets that ravaged the Byzantine-controlled shores of the Aegean Sea. Crete returned to Byzantine rule under Nikephoros Phokas, who launched a huge campaign against the Emirate of Crete in 960 to 961.
+Meanwhile, the Bulgarian Empire threatened Byzantine control of Northern Greece and the Aegean coast to the south. Under Presian I and his successor Boris I, the Bulgarian Empire managed to obtain a small portion of the northern Aegean coast. Simeon I of Bulgaria led Bulgaria to its greatest territorial expansion, and managed to conqueror much of the northern and western coasts of the Aegean. The Byzantines later regained control. The Second Bulgarian Empire achieved similar success along, again, the northern and western coasts, under Ivan Asen II of Bulgaria.
+The Seljuq Turks, under the Seljuk Empire, invaded the Byzantine Empire in 1068, from which they annexed almost all of Anatolia, including the east coast of the Aegean Sea, during the reign of Alp Arslan, the second Sultan of the Seljuk Empire. After the death of his successor, Malik Shah I, the empire was divided, and Malik Shah was succeeded in Anatolia by Kilij Arslan I, who founded the Sultanate of Rum. The Byzantines yet again recaptured the eastern coast of the Aegean.
+After Constantinople was occupied by Western European and Venetian forces during the Fourth Crusade, the area around the Aegean sea was fragmented into multiple entities, including the Latin Empire, the Kingdom of Thessalonica, the Empire of Nicaea, the Principality of Achaea, and the Duchy of Athens. The Venetians created the maritime state of the Duchy of the Archipelago, which included all the Cyclades except Mykonos and Tinos. The Empire of Nicaea, a Byzantine rump state, managed to effect the Recapture of Constantinople from the Latins in 1261 and defeat Epirus. Byzantine successes were not to last; the Ottomans would conquer the area around the Aegean coast, but before their expansion the Byzantine Empire had already been weakened from internal conflict. By the late 14th century the Byzantine Empire had lost all control of the coast of the Aegean Sea and could exercise power around their capital, Constantinople. The Ottoman Empire then gained control of all the Aegean coast with the exception of Crete, which was a Venetian colony until 1669.
+The Greek War of Independence allowed a Greek state on the coast of the Aegean from 1829 onward. The Ottoman Empire held a presence over the sea for over 500 years until its dissolution following World War I, when it was replaced by modern Turkey. During the war, Greece gained control over the area around the northern coast of the Aegean. By the 1930s, Greece and Turkey had about resumed their present-day borders.
+In the Italo-Turkish War of 1912, Italy captured the Dodecanese islands, and had occupied them since, reneging on the 1919 Venizelos–Tittoni agreement to cede them to Greece. The Greco-Italian War took place from October 1940 to April 1941 as part of the Balkans Campaign of World War II. The Italian war aim was to establish a Greek puppet state, which would permit the Italian annexation of the Sporades and the Cyclades islands in the Aegean Sea, to be administered as a part of the Italian Aegean Islands. The German invasion resulted in the Axis occupation of Greece. The German troops evacuated Athens on 12 October 1944, and by the end of the month, they had withdrawn from mainland Greece. Greece was then liberated by Allied troops.
+Many of the islands in the Aegean have safe harbours and bays. In ancient times, navigation through the sea was easier than travelling across the rough terrain of the Greek mainland, and to some extent, the coastal areas of Anatolia. Many of the islands are volcanic, and marble and iron are mined on other islands. The larger islands have some fertile valleys and plains.
+Of the main islands in the Aegean Sea, two belong to Turkey – Bozcaada (Tenedos) and Gökçeada (Imbros); the rest belong to Greece. Between the two countries, there are political disputes over several aspects of political control over the Aegean space, including the size of territorial waters, air control and the delimitation of economic rights to the continental shelf. These issues are known as the Aegean dispute.
+Multiple ports are located along the Greek and Turkish coasts of the Aegean Sea. The port of Piraeus in Athens is the chief port in Greece, the largest passenger port in Europe[17][18] and the third largest in the world,[19] servicing about 20 million passengers annually. With a throughput of 1.4 million TEUs, Piraeus is placed among the top ten ports in container traffic in Europe and the top container port in the Eastern Mediterranean.[20] Piraeus is also the commercial hub of Greek shipping. Piraeus bi-annually acts as the focus for a major shipping convention, known as Posidonia, which attracts maritime industry professionals from all over the world. Piraeus is currently Greece's third-busiest port in terms of tons of goods transported, behind Aghioi Theodoroi and Thessaloniki.[21] The central port serves ferry routes to almost every island in the eastern portion of Greece, the island of Crete, the Cyclades, the Dodecanese, and much of the northern and the eastern Aegean Sea, while the western part of the port is used for cargo services.
+As of 2007, the Port of Thessaloniki was the second-largest container port in Greece after the port of Piraeus, making it one of the busiest ports in Greece. In 2007, the Port of Thessaloniki handled 14,373,245 tonnes of cargo and 222,824 TEU's. Paloukia, on the island of Salamis, is a major passenger port.
+Fish are Greece's second largest agricultural export, and Greece has Europe's largest fishing fleet.[22]  Fish captured include sardines, mackerel, grouper, grey mullets, sea bass, and seabream. There is a considerable difference between fish catches between the pelagic and demersal zones;[23] with respect to pelagic fisheries, the catches from the northern, central and southern Aegean area groupings are dominated, respectively, by anchovy, horse mackerels, and boops. For demersal fisheries, the catches from the northern and southern Aegean area groupings are dominated by grey mullets and pickerel (Spicara smaris) respectively.
+The industry has been impacted by the Great Recession.[clarification needed] Overfishing and habitat destruction is also a concern, threatening grouper, and seabream populations, resulting in perhaps a 50% decline of fish catch.[24] To address these concerns, Greek fishermen have been offered a compensation by the government. Although some species are defined as protected or threatened under EU legislation, several illegal species such as the molluscs Pinna nobilis, Charonia tritonis and Lithophaga lithophaga, can be bought in restaurants and fish markets around Greece.[25]
+The Aegean islands within the Aegean Sea are significant tourist destinations. Tourism to the Aegean islands contributes a significant portion of tourism in Greece, especially since the second half of the 20th century.[26] A total of five UNESCO World Heritage sites are located the Aegean Islands; these include the Monastery of Saint John the Theologian and the Cave of the Apocalypse on Patmos,[27] the Pythagoreion and Heraion of Samos in Samos,[28] the Nea Moni of Chios,[29] the island of Delos,[30] and the Medieval City of Rhodes.[31]
+Greece is one of the most visited countries in Europe and the world with over 33 million visitors in 2018,[32] and the tourism industry around a quarter of Greece's Gross Domestic Product.[33] The islands of Santorini, Crete, Lesbos, Delos, and Mykonos are common tourist destinations. An estimated 2 million tourists visit Santorini annually.[34] However, concerns relating to overtourism have arisen in recent years, such as issues of inadequate infrastructure and overcrowding.[35] Alongside Greece, Turkey has also been successful in developing resort areas and attracting large number of tourists,[36] contributing to tourism in Turkey. The phrase "Blue Cruise" refers to recreational voyages along the Turkish Riviera, including across the Aegean.[37] The ancient city of Troy, a World Heritage Site, is on the Turkish coast of the Aegean.[38]
+Greece and Turkey both take part in the Blue Flag beach certification programme of the Foundation for Environmental Education. The certification is awarded for beaches and marinas meeting strict quality standards including environmental protection, water quality, safety and services criteria.[39] As of 2015, the Blue Flag has been awarded to 395 beaches and 9 marinas in Greece. Southern Aegean beaches on the Turkish coast include Muğla, with 102 beaches awarded with the blue flag, along with İzmir and Aydın, who have 49 and 30 beaches awarded respectively.[40]

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+A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name snail is also used for most of the members of the molluscan class Gastropoda that have a coiled shell that is large enough for the animal to retract completely into. When the word "snail" is used in this most general sense, it includes not just land snails but also numerous species of sea snails and freshwater snails. Gastropods that naturally lack a shell, or have only an internal shell, are mostly called slugs, and land snails that have only a very small shell (that they cannot retract into) are often called semi-slugs.
+Snails have considerable human relevance, including as food items, as pests, and as vectors of disease, and their shells are used as decorative objects and are incorporated into jewelry.[1] The snail has also had some cultural significance, tending to be associated with patience. The snail has also been used as a metaphor: someone who is not moving fast enough is "slow as a snail." The snail is the same or similar shape as the Cochlea.[2]
+Snails that respire using a lung belong to the group Pulmonata. As traditionally defined, the Pulmonata were found to be polyphyletic in a molecular study per Jörger et al., dating from 2010.[3] But snails with gills also form a polyphyletic group; in other words, snails with lungs and snails with gills form a number of taxonomic groups that are not necessarily more closely related to each other than they are related to some other groups.
+Both snails that have lungs and snails that have gills have diversified so widely over geological time that a few species with gills can be found on land and numerous species with lungs can be found in freshwater. Even a few marine species have lungs.
+Snails can be found in a very wide range of environments, including ditches, deserts, and the abyssal depths of the sea. Although land snails may be more familiar to laymen, marine snails constitute the majority of snail species, and have much greater diversity and a greater biomass. Numerous kinds of snail can also be found in fresh water.
+Most snails have thousands of microscopic tooth-like structures located on a banded ribbon-like tongue called a radula. The radula works like a file, ripping food into small pieces. Many snails are herbivorous, eating plants or rasping algae from surfaces with their radulae, though a few land species and many marine species are omnivores or predatory carnivores. Snails cannot absorb colored pigments when eating paper or cardboard so their feces are also colored.[4]
+Several species of the genus Achatina and related genera are known as giant African land snails; some grow to 15 in (38 cm) from snout to tail, and weigh 1 kg (2 lb).[5]  The largest living species of sea snail is Syrinx aruanus; its shell can measure up to 90 cm (35 in) in length, and the whole animal with the shell can weigh up to 18 kg (40 lb).  Recently, the smallest land snails, Angustopila dominikae,  have  been discovered in China, and measure 0.86mm long.[6]
+The snail Lymnaea makes decisions by using only two types of neurons: one deciding whether the snail is hungry, and the other deciding whether there is food in the vicinity.[1]
+The largest known land gastropod is the African giant snail Achatina achatina, the largest recorded specimen of which measured 39.3 centimetres (15.5 in) from snout to tail when fully extended, with a shell length of 27.3 cm (10.7 in) in December 1978. It weighed exactly 900 g (2 lb). Named Gee Geronimo, this snail was owned by Christopher Hudson (1955–79) of Hove, East Sussex, UK, and was collected in Sierra Leone in June 1976.[2]
+Gastropods that lack a conspicuous shell are commonly called slugs rather than snails.[3] Some species of slug have a maroon-brown shell, some have only an internal vestige that serves mainly as a calcium lactate repository, and others have some to no shell at all. Other than that there is little morphological difference between slugs and snails. There are however important differences in habitats and behavior.
+A shell-less animal is much more maneuverable and compressible, so even quite large land slugs can take advantage of habitats or retreats with very little space, retreats that would be inaccessible to a similar-sized snail. Slugs squeeze themselves into confined spaces such as under loose bark on trees or under stone slabs, logs or wooden boards lying on the ground. In such retreats they are in less danger from either predators or desiccation. Those are often suitable places for laying their eggs.
+Slugs as a group are far from monophyletic; scientifically speaking "slug" is a term of convenience with little taxonomic significance. The reduction or loss of the shell has evolved many times independently within several very different lineages of gastropods. The various taxa of land and sea gastropods with slug morphology occur within numerous higher taxonomic groups of shelled species; such independent slug taxa are not in general closely related to one another.
+Land snails are known as an agricultural and garden pest but some species are an edible delicacy and occasionally household pets.
+There are a variety of snail-control measures that gardeners and farmers use in an attempt to reduce damage to valuable plants. Traditional pesticides are still used, as are many less toxic control options such as concentrated garlic or wormwood solutions. Copper metal is also a snail repellent, and thus a copper band around the trunk of a tree will prevent snails from climbing up and reaching the foliage and fruit. A layer of a dry, finely ground, and scratchy substance such as diatomaceous earth can also deter snails.[4]
+The decollate snail (Rumina decollata) will capture and eat garden snails, and because of this it has sometimes been introduced as a biological pest control agent. However, this is not without problems, as the decollate snail is just as likely to attack and devour other gastropods that may represent a valuable part of the native fauna of the region.
+In French cuisine, edible snails are served for instance in Escargot à la Bourguignonne. The practice of rearing snails for food is known as heliciculture. For purposes of cultivation, the snails are kept in a dark place in a wired cage with dry straw or dry wood. Coppiced wine-grape vines are often used for this purpose. During the rainy period, the snails come out of hibernation and release most of their mucus onto the dry wood/straw. The snails are then prepared for cooking. Their texture when cooked is slightly chewy and tender.
+As well as being relished as gourmet food, several species of land snails provide an easily harvested source of protein to many people in poor communities around the world. Many land snails are valuable because they can feed on a wide range of agricultural wastes, such as shed leaves in banana plantations. In some countries, giant African land snails are produced commercially for food.
+Land snails, freshwater snails and sea snails are all eaten in many  countries. In certain parts of the world, snails are fried as for example, in Indonesia, they are fried as satay, a dish known as sate kakul. The eggs of certain snail species are eaten in a fashion similar to the way caviar is eaten.[citation needed]
+In Bulgaria, snails are traditionally cooked in an oven with rice or fried in a pan with vegetable oil and red paprika powder. Before they are used for those dishes, however, they are thoroughly boiled in hot water (for up to 90 minutes) and manually extracted from their shells. The two species most commonly used for food in the country are Helix lucorum and Helix pomatia.[citation needed]
+Snails and slug species that are not normally eaten in certain areas have occasionally been used as famine food in historical times. A history of Scotland written in the 1800s recounts a description of various snails and their use as food items in times of plague.[5]
+Skin creams derived from Cornu aspersum snails are sold for use on wrinkles, scars, dry skin, and acne. A research study suggested that secretions produced under stress by Cornu aspersa might facilitate regeneration of wounded tissue.[6]
+Because of its slowness, the snail has traditionally been seen as a symbol of laziness. In Christian culture, it has been used as a symbol of the deadly sin of sloth.[7][8] Psalms 58:8 uses snail slime as a metaphorical punishment. In Mayan mythology, the snail is associated with sexual desire, being personified by the god Uayeb.[9]
+Snails were widely noted and used in divination.[7] The Greek poet Hesiod wrote that snails signified the time to harvest by climbing the stalks, while the Aztec moon god Tecciztecatl bore a snail shell on his back. This symbolised rebirth; the snail's penchant for appearing and disappearing was analogised with the moon.[10]
+Professor Ronald Chase of McGill University in Montreal has suggested the ancient myth of Cupid's arrows might be based on early observations of the love dart behavior of the land snail species Cornu aspersum.[11]
+In contemporary speech, the expression "a snail's pace" is often used to describe a slow, inefficient process. The phrase "snail mail" is used to mean regular postal service delivery of paper messages as opposed to the delivery of email, which can be virtually instantaneous.
+Keong Emas (Javanese and Indonesian for Golden Snail) is a popular Javanese folklore about a princess magically transformed and contained in a golden snail shell. The folklore is a part of popular Javanese Panji cycle telling the stories about the prince Panji Asmoro Bangun (also known as Raden Inu Kertapati) and his consort, princess Dewi Sekartaji (also known as Dewi Chandra Kirana).
+Certain varieties of snails, notably the family Muricidae, produce a secretion that is a color-fast natural dye. The ancient Tyrian purple was made in this way as were other purple and blue dyes.[12][13][14] The extreme expense of extracting this secretion is sufficient quantities limited its use to the very wealthy. It is such dyes as these that led to certain shades of purple and blue being associated with royalty and wealth.[15]
+Throughout history, snails have been kept as pets. There are many famous snails such as Lefty (Born Jeremy) and within fiction, Gary and Brian the snail .[16]