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lean on them. Absent in bonobos. 3. sit and turn back Solicit grooming by approaching another, then turning back to potential groomer. See also Solicit grooming. Also in bonobos. See video. sit behind another Change position to groom back of companion. Also in bonobos. See video. S sit face to face Shift posture to sit en face, in social grooming or seeking to receive food. Also in bonobos. 2. See video. sit on In social play, one player sits on other player. Cf. step on in which smaller playmate rides on larger one lying supine. Cf. also trample. Category 38. See video. sit prone Slouch or bend forward, extend arms, and put palms on ground, often when being groomed. Also in bonobos. See video. sit sideways 171 sit

sideways Shift posture to sit beside another, to solicit grooming. See also solicit grooming. Also in bonobos. See video. sitting hunch See hunch and sit. slap Plooij SLA. Goodall: A hitting down gesture when the palmar surface of the hand is brought into contact with the objective. Done with one or both hands. Context includes aggression, charging display, drumming, social play, solicitation of courtship and play, etc. Bonobos slap in aggression, charging display, drumming and social play. See video. slap belly See slap self. slap branch Whiten et al. Branch slap. Slap branch with palmar surface of one hand for getting attention. Few adult males of M group do in courtship display. Customary at Bossou, Ta and Budongo. Branch slap described for Lomako bonobos may differ from chimpanzee

pattern. Cf. rap. slap buttress or tree trunk Strike tree with hand or alternate hands during charging display. Goodalls Drum. See also drum and slap wall. Included in Kanos Drum for bonobos. See video. slap ground Strike substrate with hand or alternate hands during pant-hooting, threatening, charging display, fighting, and play. Slap ground and pant-hoot in response to pant-hoot calls from distance. Recorded from Beni. Bonobos slap ground while staring at opponent, so context more narrow. See video. See also video enter hole. slap in invitation Nishida Invitation slap. Repeatedly strike rock or ground with one or both hands, to entice youngster into play. Elements include: slap ground, play face, stand quadrupedal. Adolescent male, Cadmus, repeatedly slapped hand of adult female lying on ground with back of hand

softly while showing play face. Absent in bonobos of Wamba. 5. See video. slap other Done during play, play invitation, charging display or fighting. Juvenile male, Xmas, slapped back of adult male while grooming him. Pikas Slap for bonobos. See video. S 172 smack lip slap self Nishida et al. Slap belly. Slap own belly, chest, or thigh. Five-yearold juvenile male, Cadmus, struck belly with right hand to make drum-like sounds, while hanging from branch by left arm, even when alone. Solo play or response to approach of human observer. Other juveniles, Xmas and Michio, later began regularly to slap belly or chest in intimidation display and solicitation of play, while hanging in tree, standing bipedal or walking bipedal on ground. Male infant, Caesar, slapped belly and thigh

in self play. Absent in bonobos of Wamba. See videos. slap-stamp Goodall: Slapping with hands and stamping with feet together. Plooijs Slap stamp. Element of charging display. Absent in bonobos of Wamba. 5. See video. slap wall Slap wall of metal house with one or both hands or alternate hands, while standing bipedal or quadrupedal. Done regularly by M group adult males at Mahale and by Kasakela males at Gombe. See also slap buttress or tree trunk and drum. See video. See also video slap buttress or tree trunk. sleep Lie or sit immobile with closed eyes, unconscious. See video. slide down boulder Youngsters occasionally slide supine or prone down large slanted boulder. 8. See video. slide down vertically Goodall Fireman slide: A chimp may slide down a

vertical or diagonal pole or tree in an upright or crouched position. All four limbs are in contact with the pole or tree. Kanos Vertical slide. See video. S sloth position See hang in sloth position. sloth walk See walk in sloth position. smack lip Goodall Lip smack: Occurs during grooming. Mouth slightly opened and closed rhythmically. Plooijs LIP. Also at Ta. See also clack teeth and sputter. Similar in bonobos, but unknown if homologous to smack lip. 3. See video. smell 173 smell See sniff. snatch Take suddenly and forcibly object such as food from another, against resistance. Adult males snatch food, especially meat, from others of any age-sex class, mothers from immature offspring, and juvenile sons from mothers. Juvenile may snatch branch from playmate. In bonobos,

mostly done by juveniles. See video. sneer Goodall: The upper lip is retracted, sometimes on one side more than the other, to expose part of the upper teeth. In the wild the context is associated with fear of a human or a human associated situation. Only a few chimps sneer. Plooijs Sneer. At Mahale few individuals sneer. Cf. grin. 9. See video. sneeze Goodall: Chimps sneeze in the same way as humans. See also probe nasal passage. Kanos Sneeze for bonobos. See sickness. See video. sniff Goodall Sniff. Put nose close to something, such as substrate, vegetation, tree trunk, fruit, food wadge, feces, urine etc. apparently to obtain information about which chimpanzee or what kind of animal was there. Sniff ground after left behind by companions, or when

arriving at place where strange chimpanzees were before. Males sniffed more frequently than females in sexual and social situations, while females did so more often during feeding and self-checking. Also reported from Ta. Bonobos sniffing predominantly by juveniles. See video. sniff finger Plooij SFI. Goodall: One finger, usually index finger, is carefully smelled after being used to investigate some object. The most usual context is during inspection of the genital area. of a female. See inspect genitals. Juvenile bonobos do so. See video. sniff fruit Hold and sniff large hard-shelled fruits such as Saba comorensis and Voacanga lutescens to judge ripeness. 3. See video. sniff genital See inspect genitals. sniff mouth Sit or stand quadrupedally very close to another individual and sniff its mouth to obtain odor of

food being eaten. Occurs especially for food new to S 174 solicit copulation immature individual. From behavior pattern only, difficult to discriminate from peer when peerer does not sniff. 7. See video. sniff with tool Goodall Investigation probe: A stick or twig is used by a chimp for touching some object. See also probe. Absent in bonobos of Wamba. 5. See video. sniffle Audible sound of runny nose. See eat nasal mucus and sickness. snub Rebuff by inaction. Apparently deliberate refusal to look at conspecific who approaches closely to seek contact. At Mahale, alpha male, Ntologi, did not look at beta male, Nsaba, even if Nsaba continuously followed Ntologi apparently seeking to groom him. Apparently signals intimidation that alpha male does not wish to interact with beta. Cf.

ignore. 8. See video. social play See play socially. social scratch See scratch socially. soil-eating See eat termite soil. solicit companion When about to depart, adult encourages continued proximity from another resting nearby or about to leave in another direction by look back, look through thighs, touch, pull, shake branch, etc. Cf. depart together and herd. 3. See video. S solicit copulation Courtship. Gesture and posture directed to estrous female by male or vice versa that normally leads to copulation. As male behavior toward estrous female, includes clip leaf, bend shrub, pull through in courtship, swagger bipedal, hop quadrupedal on spot, hunch and sit, shake branch, open thighs, extend hand, palm downward, erect penis, bristle, gaze repeatedly, club ground, approach, etc. Female solicits by gaze, glance, clip leaf,

approach, follow, dance bipedal and crouch. Clip leaf, bend shrub, dance bipedal, etc. unique to Mahale. Goodalls Courtship includes display brachiation. In bonobos of Wamba, hunch bipedal, shake branch, rock, open thighs, and hunch and sit are courtship displays. See video. See also videos aggress in sexual frustration and copulate dorso-ventral. solicit grooming 175 solicit grooming van Hooff Groom-present. Goodalls Present for grooming: The rump, back, or bowed head are the most usual body parts presented to a chosen grooming partner at the start of a session. Scratch self, raise arm, sit and turn back, sit sideways, raise leg, raise arm to hold branch, raise arm slowly, lie with back to another, etc. also used. Alter position of limbs, head or body-part to facilitate continuation of grooming. Self-scratching

most important of four patterns of solicitation for grooming in bonobos of Wamba. See video. solicit grooming turn Signal requesting partner to take turn in social grooming: stop grooming and release partners body, stop grooming and scratch or groom self, stop grooming and present back or other body-part to partner. Bonobos show similar behavior. See video. solicit play Invite another to play. Includes: approach with play face, flee, extend hand, hit lightly, hit and run, hold object in mouth, lie supine, lie prone, slap in invitation, stamp in invitation, somersault, play face, play walk, throw branch, tickle, tilt head, watch, and many others. Includes Goodalls Try play: Unsuccessful attempt to initiate play with another and Hayakis Paraplay. Bonobos invite play by approach with play face, hit lightly or

pull the hand of partner. 2. See videos. solicit play with object Invite play by holding plant part in mouth, hand, foot, or groin pocket. Customary at Gombe, Mahale, Kibale, Budongo, and Ta. Present at Bossou, Assirik, and Goualougo. Also, includes club ground, flail long object, throw, etc. See video. solicit play with object in mouth Whiten et al. Play start: Invite play by holding stem or other object in mouth. Probably chimpanzee universal. See video. solicit reassurance contact Aggressed-against or frightened infant runs to mother, whimpers with extended hand or throws temper tantrum. Aggressed-against adult seeks contact with third party or with aggressor by extending hand, etc. Goodall mentioned but did not name pattern. Kanos Request for appeasement. Aggressed-against bonobo female pesters aggressor female and persistently peers

at her, so that aggressor invites victim to G-G rub. See video. solicit riding Mother or allomother invites infant to ride on back or belly by extend hand, hoo, look back, raise arm with elbow bent, touch etc. S 17 spit juice Cf. depart together, put dorsal, put ventral and scoop infant. 3. See video. solicit support Request support from third party when threatened or attacked. Includes: search for conspecific, approach, extend arm, scream, touch, mount, thrust, kiss, embrace, press teeth against back, etc. At Wamba, bonobos solicit only mother for support. See video. solitary play See play solo. solo play See play solo. somersault Goodall: Roll head over heels. Plooijs Somersault. Usually repeated many times. Accompanies social play such as circle and wrestle. Also used as solicitation

of play. Mature and immature chimpanzees do. At Mahale, 34 year old infants often do in solo play when large group travels long distance. See also headstand. Absent in bonobos of Wamba, but seems to occur in captivity. See video. somersault, backward Somersault backwards. Context of occurrence is similar to somersault, forward and somersault, side. 5. See video. somersault, backward with dry leaves Collect dry leaves on belly and somersault backward with them; while rolling, leaves scatter little by little. Juvenile males, such as Xmas and Primus, show in solo play. See video. somersault, forward Usual somersault in forward direction. Often used as solicitation of play and during play. Circle quadrupedal usually accompanied by this type of somersault. Also done as solo play. See video. S somersault, side

Somersault sideways. Used as solicitation of play and during social play, but also done as solo play. 5. See video. spar At start of fight, both parties stand bipedally and flail one or both arms in hitting movements, with or without screaming. Also in play. Bonobo juveniles hit each other in play. spit juice Forcefully expel juice of fruit such as unripe Saba comorensis, buds such as Milicia excelsa, or young leaves such as Brachystegia spiciformis. 7. See video. spit seed 177 spit seed Spit large seeds such as Garcinia huillensis, Voacanga lutescens, Syzigium guineense, etc. at site where eaten. Also in bonobos. See video. spit water Infant or juvenile occasionally spits water during drinking session. Solo play?. See video. splash water Solo play: Youngster splashes water with

palm or fingers of hand, foot, leg, or mouth. Sprinkle water onto playmate. See also play with water. Cf. stir water. 8. See video. See also video play with water. sponge See leaf-sponge. sponge push-pull See push and pull sponge. spring Infant rushes and jumps onto mothers buttocks to catch up with her rapid movement. 3. See video. sputter Nishida et al.: Sound uttered by many chimpanzees of Ngogo but only few of Mahale when grooming another and finding something interesting. Usually, done in same context as clack teeth and smack lip. While sputtering they do not show rhythmic lip movements. Sounds as if chimpanzees force air through lips, or force saliva through teeth. Present at Bossou. 9. See video. squash ectoparasite on arm Whiten et al. Index-hit:

Squash ectoparasite on arm. Customary only at Ta. See squash ectoparasite on palm. Absent at Mahale. squash ectoparasite on palm Nakamura and Nishida Squash ectoparasite on palm. Customary only at Bossou. Cf. squash ectoparasite on arm. Absent at Mahale. squash leaf Whiten et al. Leaf-squash: Squash ectoparasite on leaf. Mahale chimpanzees take louse from body, place it on leaf, fold leaf, and squash it with thumb and fingers. Habitual at Gombe. See also groom leaf. 5. See video. squat Hunt et al.: The body weight is borne solely by the feet/foot, both hip and knee are strongly flexed. Also in bonobos. Cf. sit. S 178 stamp other squat bob See bob. squeal in copulation High-pitched, scream-like sound accompanied by grin, emitted by estrous female at final stage of

copulation. Goodalls Copulation scream. Bonobos emit similar call. See video. squeeze Compress contents of mouth by pushing chin upwards. Hand or arms pressed against chin to facilitate wadging. See also wadge. Also in bonobos. See video. staccato call Newborn emits call in response to mothers sudden movement, approach by others, noises in surroundings, etc. Plooijs Staccato. See video. stalk Approach stealthily to prey or other chimpanzees, especially neighbors. Absent in bonobos. stamp Goodall: Forceful downward kick of one foot, or alternate feet. In charging displays, The chimp stamps his feet one after the other as he runs along. One foot is often stamped harder than the other. A standing chimp may stamp alternate feet.A sitting chimp may stamp several times with one foot. van Hooffs Stamp. Plooijs Stamp.

Done also during social play, in solicitation of play and courtship. Cf. kick. Bonobos stamp while running, but not while sitting. Pikas Stomp for captive bonobos. 2. See video. S stamp bipedal Stamp sole on ground, buttress, tree trunk, wall of house, or another while upright. Component of charging display. Adolescent males stamp with alternate feet on ground in courtship. Bonobos of Wamba stamp bipedal in courtship, but not in charging display. 3. See video. stamp in invitation Nishida Invitation stamp. Mainly youngsters, lightly stamp on ground while watching potential playmate, inviting social play or alloparental care. Bipedal or quadrupedal. Absent in bonobos of Wamba. 7. See video. stamp other Goodall: Stamping on a victim is a major component of withinspecies fighting, and can result in severe bruising

and contusions. Also stamp softly on playmate repeatedly in social play. Also in bonobos. 2. See video. stamp quadrupedal 179 stamp quadrupedal Stamp quadrupedal on ground, rock, buttress, tree trunk, wall of house, another, or carcass of animal. Component of charging display, etc. Juveniles stamp on rock or lying adult in play. Absent in bonobos? Category 3. See video. See also video pant-hoot. stamp trot van Hooff: The animal walks or trots along while stamping heavily with its hind feet on the ground. Often this is done in a rhythmical manner in that one foot is placed down gently and the other with force. The head is mostly kept rather low and tucked back between the shoulders which are pulled upwards. The head may make low amplitude vertical

rocking movements. Kanos Stamping run and Pikas Stomp for bonobos. See video. stamp water Juvenile female, Jiddah, while sitting, repeatedly stamped on small pool of water, as part of water play in stream. Adolescent male, Cadmus, bent branches onto water and stamped repeatedly on them as solo play. 9. See video. stand bipedal Plooij STB. Stand upright. Stance assumed when watching distant object or taking food from tall shrubs such as Ficus urceolaris. Mother may drop infant by almost standing bipedal. Cf. walk bipedal. Bipedal posture occurs in courtship, mixed with other elements, such as shake branch, thrust, leap, and stamp. Common in bonobos. See video. stand quadrupedal Stop walking and scan environment or wait for another, etc. Plooijs STQ. Common in bonobos. See video. stand quadrupedal heel

up While waiting for companion, or monitoring environment, stand still with heel up and toes on ground. Kanos Waiting posture for bonobos. See video. stand with head down, bottom up Nishida: Head down hips up: Postural pattern with tickling by mouth. The mother or caretaker stands on all fours, with elbows flexed, and putting its mouth against the infants body on the ground. The open mouth may be pressed against any part of the infants body, but especially its belly, back, head, neck, foot or arms. Similar to Fore-limb crouch. Absent in bonobos?. See video. stare fixedly Goodall Fixed stare: The chimp stares intently at the individual with whom it is interacting. At Mahale, groomee often stares fixedly at S 180 stepping-stick groomer during intense social grooming. Also

in threat, predation, and monitoring. Cf. glance, look, peer, and watch. Kanos Stare for bonobos, who stare mutually during mating. See video. stare fixedly with head down and bottom up Infant female, Acadia, stood quadrupedal with head lowered onto palmer side of flexed lower right arm, resting on ground, while raising buttocks, and stared at mother being groomed by adult male. Request for suckling? Mother finally suckled Acadia. Weanling, Acadia, approached mother, hesitant to make contact with her, and showed this idiosyncratic posture. See video. startle flee See flee after startle. startle flinch See parry. startle reaction See parry. steal Remove surreptitiously object, without owners awareness. See Goodalls Steal, which includes both snatch and steal. Also in bonobos. stem pull-through See pull through stem. step on Infant or

juvenile elicits play by stepping on reclining partner. Often response of younger partner to older partner who lies supine in playsoliciting posture. Goodalls Ride onto. 7. See video. S step over Step over another lying on ground, who shows no response to stepping individual. No obvious function. 7. See video. step up on leg Infant steps onto leg of mother in climbing up to ride on her back. 5. See video. stepping-stick Branch used as sandal to avoid sharp thorns when moving in kapok tree. Reported from Tenkere. Absent at Mahale. See also seat-stick. stingless bees 181 stingless bees Small, colonial, honey producing bees of genus Trigona. See eat insect. stir water Stir water vigorously with hand, index finger, large leaf, or foot in river. Solo play of

juveniles, adolescents and adults. See also play with water. See videos. See also video play with water. stomp See stamp. store Hold leaves in hand and start chewing only after collecting handful. Occurs when eating tough leaves of Ficus exasperata. Seen also at Gombe. 5. See video. strike down prey See knock with both arms or knock with one arm. strip leaf Sit and tear off one or few leaves in quick succession from shrub or small tree. Older adolescent male, Orion, does this in courtship. Cf. pull through in courtship in which pull-through movement against stem is performed. Habitual threat gesture at Gombe and Kibale, and present at Bossou. See video. strip leaves by pull-through See pull through. stroke Stroke back of another with extended fingers. Done

by adolescent male, Cadmus. Idiosyncratic? Stroking face of another with fingers and palm is rare and of unknown function. Cf. scratch socially, stroke type. 9. See video. struggle free When pinned down by stronger partner in play, or by group during gang attack, victim exerts maximum effort to get free by push, pull, kick, bite, etc. 3. stumble Lose footing, but rarely trip over stone or stick. See video. See also video slap other. subcutaneous tumor Tumor-like skin swelling of hemispheric shape, approximately size of ping-pong ball, often protruding from lower abdomen of chimpanzee. See video. S 182 support older subgroup See party. subordinate Lower-ranking of two individuals. Cf. dominant. See Fig. 11. suck Suck nipple. Plooijs Suck. Goodalls Suckle. Kanos Suckle for bonobos. See video. suck in

reassurance Goodall Reassurance suckle: A brief suck when the infant has been frightened or hurt. Absent in bonobos. See video. suck self Suck own nipples. Female who has lost newborn rarely sucks her nipples. Matsumoto-Odas Self suckle. 8. See video. suck thumb Infant sucks own thumb, but rarely sucks other digits. See videos. suck toe Infant sucks own toe like suck thumb. Adults and immatures suck toe to remove eggs of sand flea. See also remove sand flea. Category 2. See video. suckle Mother allows infant to suck from nipples. Goodalls Suckle. Kanos Suckle for bonobos. See video. supplant Nishida and Turner Displacement. Displace another from food patch by bark, approach, run, push, raise arm quickly, or shake branch. Adult males usually supplant adult females and other adult

males. Cf. vacate. In bonobos, adult females supplant adult males, who supplant other adult males. See video. S support Intervene in fight and side with either of conflicting parties. Rare in bonobos. See video. support dominant Intervene in fight and side with dominant party. de Waals Winner support. Cf. support subordinate. 3. See video. support older Intervene in fight and side with older rather than younger party. See video. support subordinate 183 support subordinate Intervene in fight and side with subordinate party. de Waals Loser support. Cf. support dominant. 3. See video. suspend When chimpanzee suspended above ground, dangle by hand object such as carcass. 3. See video. suspend and shake up and down When chimpanzee suspended above ground, dangle and shake object up and down by hand.

Solo object play. 8. See video. swagger bipedal Goodall Bipedal swagger: In an upright or semi-upright posture the chimp sways, often rhythmically, from one foot to the other. The animal may remain in one spot to swagger or move forward during the swagger. The arms are normally held out from the body, the shoulders hunched up. It is often used during aggression, greeting, and courtship. van Hooffs Sway walk. Plooijs Bipedal swagger. Nishidas Bipedal swagger. Often performed as separating intervention just before charging display. Similar behavior occurs in bonobos of Wamba. See video. swagger on knuckles Adult male, Pim, approaches estrous female in orthograde posture with knuckles on ground and penis erect. Idiosyncratic courtship display. 9. See video. swallow leaf Wrangham and Nishida Leaf-swallow. Ingest slowly and without

chewing, usually in early morning, leaves of certain species of herbs and trees. Leaves of Aneilema spp., Aspilia mossambicensis, Commelina spp., Ficus exasperata, Lippia pulicata and Trema orientalis swallowed at Mahale. Known from Ta, Kibale, Bossou, Gombe, and other localities. Species of swallowed leaves differ from place to place. Culture universal in chimpanzees? Reported for bonobos. See video. swallow seed Ingest seeds without chewing them. Includes tiny seeds such as fig and large seeds such as Pycnanthus angolensis. sway and move Climb high up, sway tree trunk back and forth using bodys weight, then catch and grasp terminal branch of another tree while original tree is bent over. Kanos Sway tree/branch for bonobos. Arboreal locomotion. Cf. swing and take. See video. S 184 swing forward and upward sway

woody vegetation Component of charging display. While standing bipedal, vigorously shake tree trunk of short tree/large horizontal branch/woody vine/ fallen trunk of tree/rock, back and forth with both hands, sometimes destroying the dead tree as result. Type of threat display and used also as courtship. Probably same as Goodalls Branch sway: The chimp vigorously sways a growing branch usually when standing on ones feet. During a charging display, a male may pause and sway branches. Plooijs Branch-sway. Absent in bonobos of Wamba. 5. See videos. swelling of sexual skin Female anogenital region begins to swell cyclically at about 7 years of age and shows maximal swelling from about 10 years of age. After adolescence, sexual skin shows tumescence, maximal tumescence and dutemescence over typical menstrual cycle of 35

days. Maximal tumescence lasts 710 days, and in last two days of tumescence ovulation occurs. Plooijs Swelling. See also adolescent swelling and flabby bottom. swing Hang from woody vine by arms and swing back and forth. Social and locomotor play, solo play, threat, and charging display. If performer changes from one support to another, becomes brachiate. Goodalls Swing is equivalent to brachiation. Kanos Swing for bonobos. See video. swing above Swing back and force above potential playmate while watching intently. Play solicitation, especially by infant female. 9. See video. S swing and grasp Swing back and forth and when reaching farthest point, leap to or grab terminal branch of neighboring tree and transfer to it. Used when food abundant in canopy. Arboreal locomotion. Cf. leap between trees which

lacks swinging. Also, cf. sway and move. See video. swing and kick Hang from woody vine, swing, and forcefully kick buttress of tree, taking advantage of momentum of swinging movement. Sometimes, swing and kick other on back. Absent in bonobos of Wamba. See video. See also video play solo. swing forward and upward Hang by hands, extend legs forward by swinging, rotate whole body, and at same time assume orthograde position. Similar to human gymnastics. Arboreal solo play. 9. See video. T tag Nishida. Alternately chase and flee. Includes run, play-pant, play face etc. Also in bonobos. Ta Study area of Pan troglodytes verus within Ta National Park, Cote dIvoire. Research organized by C. Boesch from 1979 to present. take Take food or tool with owners knowledge and

presumed consent. Modified from Goodalls Take. Tolerated scrounging equivalent to being shared. Cf. snatch, steal and share food. Plooijs Take away object. Also in bonobos. See videos. take dorsal See put dorsal. take finger in mouth Nishida, Goodall: Reach for the hand/foot of another and place part of hand/food or fingers/toes in its own mouth. Like push finger into mouth, reassurance behavior. Take finger in mouth much less frequent than push finger into mouth at Mahale. Absent in bonobos of Wamba. 5. See video. take ventral See put ventral. T tandem walk See follow in contact. tap heel Solicit copulation by hitting heel against ground/substrate. Courtship by males at Bossou. Shown by only one adolescent male, Orion, at Mahale. Idiosyncratic. See video. 185 18 threaten teach At Mahale,

teaching, if any, comprises only discouragement or negative reinforcement. Mother or allomother only rarely removes and discards leaf from mouth of infant, if leaf is not in diet of group. No systematic teaching. Teaching absent in bonobos of Wamba. 9. tear Remove leaf from branch or stalk with hand. Common method of harvesting foliage. See video. tease Dominant provokes subordinate by mild threats, such as throwing branch or fishing, without recourse to violence. Different from Adangs Tease, which is similar to harass. At Mahale, adolescent female with monkey tail tried to fish for juvenile male: She carried colobus tail up tree and dangled it from above. Whenever male jumped for it, she withdrew it, so that he could not reach it. After several attempts, male got tail, and

female grinned, suggesting that she was just trifling with male, not intending to give up tail. Adolescents and juveniles throw branches at other adolescents or juveniles, apparently to tease them. Cf. trifle with and harass. 8. See video. teeth-bare See grin. teeth-clack See clack teeth. temper tantrum See throw temper tantrum. Tenkere Study area of Pan troglodytes verus in Outamba-Kilimi National Park, northern Sierra Leone. Research done by R. Alp. termite-fish See fish for termites. T territory Area occupied, patrolled, and defended by unit group or community. Cf. core area. thigh support See transport with thigh support. threaten Intention movement or anticipatory gesture of aggression. Goodalls Threat: a repertoire of gestures to elicit submissive behavior in the throw 187 individual the gestures are directed toward The repertoire includes

tip head, flail arm, hit toward, throw at, shake branch, stamp, slap, charge, display, charging, hunch, stare fixedly, bark, cough bark, waa bark, wraa, compress lips and scream with grin-full-open. Kano lists for bonobos similar elements: stare, head-tip, wrist-shake, slap, branch-sway, arm-raise, threatening approach, and pseudo-charge. 2. See video. throw Plooij THO. Goodall: Objects, such as stones, rocks, branches, sticks, handful of grass, etc. may be thrown underarm or overarm Bonobos also throw, but underarm only. See video. throw at Goodall Throw at: When an object is aimed at a specific objective aimed throwing. Throw object directionally. Customary at Gombe, Mahale, Ta, Bossou, and Goualougo. Present at Assirik, Kibale and Budongo. Aimed throwing is divided into throw at animate object and throw at inanimate object. 3. throw at

animate object Aimed throwing at animal, conspecific, prey such as bushpig, or competitor such as baboon and human. Rarely throw stone or branch at conspecific as intimidation or play and rarely hit other. Habituated chimpanzees of Mahale and Ta throw branches at human for intimidation or play solicitation, but rarely hit. Throw at conspecific is customary at Goualougo. Throw at prey not recorded at Mahale. Absent in Wamba bonobos, but Lomako bonobos threw fruit at tortoise. 9. See video. throw at inanimate object Aimed throwing at inanimate objects. Adult males at Mahale throw heavy stick or rock against wall of metal house to make loud noise, or into water. 3. See video. throw branch Throw branch or stick in upright posture with arm. Element of charging display by

males and females, and courtship pattern to estrous female by juvenile or older infant males. Older juvenile male once threw 1-m stick from distance of 8 m while pant-grunting to adult male. Bonobos throw branch forward at end of branch-dragging display. See video. throw dry leaves In dry season, males of Mahale run, pick up pile of dry leaves from ground, and hurl them by hand. Occurs most frequently at climax or onset of T 188 thrust charging display, but also during pant-grunting. Juvenile male throws dry leaves as solo play while traveling. Not used in courtship. Absent in bonobos of Wamba. See video. throw fruit against rock At Gombe, throw hard-shelled fruit against rock or tree trunk to crack it open. Plooijs Banging. Absent at Mahale although

Strychnos available. throw sand Adult male, Fanana, throws sand with hand while running bipedally. Within M groups range, sandy ground is rare. Element of charging display and play. 9. See video. throw splash Adult males of M group, Ntologi, Kalunde, Nsaba, Fanana, and Masudi, lift and throw heavy rocks into stream often with two hands, which produces loud splash that intimidates others. Special case of throw stone or rock or throw at. See videos. throw stone or rock Hurl stones in charging display and also in play. Adult male, Alofu, simulated to hurl stone when his sex partner refused to follow him during herding. Bonobos of Wamba throw only branches because stones unavailable. See throw splash. 5. See video. T throw temper tantrum Goodall: The chimpanzee screams loudly,

and may leap up, fling arms above his/her head and then slap them onto the ground or beat the ground with his hands. He may hold himself to the ground on his face, hug a tree, or himself. He may rush off, tumbling over and over, still screaming. The screaming often results in glottal cramps Plooijs Throw temper tantrum. Typically done by infants during weaning conflict, when mother rejects sucking, food sharing, transport, grooming etc. Adults sometimes show in response to rejection by dominants. At Mahale, infant had tantrum while hanging and swinging by feet, as if it were about to fall head first from high above ground. In temper tantrums, infant not only protests, but also appears to monitor how much it can induce care from rejecting

mother. See monitor mother. Bonobos show similar behavior. See videos. thrust Plooij THU. Goodalls Pelvic thrust: Rhythmic back and forth movements of the pelvis Occurs during copulation and in reassurance mounting Two adult males embrace ventro-ventrally and bipedally in reassurance or reconciliation, and one thrusts against other. Adult females occasionally mount other females and thrust. Immatures thrust in play. May thrust on part of others body such as leg or on object. Adolescent male, Cadmus, thrust against colobus carcass. Bonobo males thrust bipedal 189 thrust like chimpanzees. See video. thrust bipedal Stand upright on feet and thrust in non-mating context. Courtship or reassurance shown by few adult and adolescent males. Intention movement of copulation likely as origin. See video. thrust in vacuum Thrust, but without any conspecific or

object. Infants or juveniles do as solo play and during play walk. 8. See video. thrust, misdirected Thrust not at rump, but on other body parts, such as leg, head, or shoulder, etc. Function is similar to usual thrust. 7. See video. thump See rap. tick Ectoparasites of chimpanzees of genus Ixodus, etc. tickle Plooij TIC. Goodall: The chimp puts one or both hands/feet on the body of the partner, usually between the neck and shoulder or in the groin, and makes tickling movements with the fingers. See also poke and pat. Bonobos may touch, push lightly, hold, or grasp infant or juvenile in social play, which may be equivalent to chimpanzee tickling. See video. tickle self with object Whiten et al. Self-tickle: Tickle self using objects. Habitual

at Gombe. Present at Goulougo. Absent at Mahale. tickle with hand or foot See tickle. tickle with mouth See mouth. tilt head Incline head to one side while standing quadruped, to solicit play, or to look above. 8. See video. tip head Goodall Head tip: a threatening gesture. Head is jerked very slightly backwards, at the same time chin is raised. The performer faces the individual being threatened and the gesture is usually accompanied by a cough bark and often an arm raise. Plooijs Tip. Also in bonobos. T 190 tool composite Tongo Study site of Pan troglodytes schwein-furthii within Virunga National Park, Democratic Republic of Congo. Research done from 1987 to 1990. tool Object extracted or detached from environment and used to obtain benefits or pleasure, in

aid or extension of manipulable organs such as limbs and mouth. See make tool and use tool. See Fig. 3. T Fig. 3 Tool: Tools used by K group to fish for Camponotus ants tool composite Two or more tool types used simultaneously and complementarily, to achieve single aim, e.g. hammer and anvil. Human examples are bow and arrow, mortar and pestle, etc. Absent at Mahale. tool set 191 tool set Two or more tool types used in obligate sequence to achieve single aim. Sugiyamas Tool composite. Includes push and pull sponge, pound pestle and leaf-sponge, hammer nut and pick out nutmeat, puncture and fish for termites, pound, lever open and dip fluid. At Mahale, two cases of using tool-set seen when immatures retrieved leaf-sponge from tree-hole with

stick. Cf. tool composite. 3. See videos. tool use See use tool. touch van Hooff Touch. Plooji TOU. Nishida Touch. Goodall: A chimpanzee reaches out with a hand and touches another, with fingers, by laying the whole palmar surface of the hand on the others body Touch signals reassurance to subordinate from dominant, and appeasement to dominant from subordinate. Mother touches infant to signal departure. At Mahale, estrous females approach and touch males on shoulder, etc., to solicit copulation. To touch head of another sometimes signals to groomee by groomer to change posture. Parts frequently touched are mouth, chin, head, hand, shoulder, and genitals. Mouth most often touched by adult male, Kalunde, to appease alpha male, Fanana. See also push finger into mouth, groom, inspect, pat and push.

Adult bonobos much less often touch each other, mostly done by infants and juveniles. See video. touch fruit Touching hard-shelled fruit important source of information about ripeness. Sometimes, only touching suffices, but may be sniffed and bitten. Type of inspect fruit. Absent in bonobos of Wamba. Category 3. See video. touch scrotum In mounting, mounted male often reaches back to touch genital area of mounter with hand, while screaming loudly. Adult male also touches scrotum of another for reassurance. While pant-grunting to adult male, adult female often reaches to touch scrotum. Kortlandts Caress anothers scrotum, for chimpanzees of Beni. 5. See video. See also video sniff finger. touch with foot Function is similar to touch with hand, but rare. Juvenile extends its leg and clandestinely touches any body-part

of newborn while grooming mothers back. 3. See video. T 192 transport bipedal touch with hand See touch. toy Object used for solo or social play. Object extracted or detached from environment used for play solicitation or as focal object during social play becomes tool. Comprises branch, fruit, leaf, flower, stone, sand, animal, piece of animal skin, piece of clothes, earthenware, cardboard, etc. See video. toy with See trifle with. tradition Behavior learned socially, shared by at least most members of at least one age or sex class, and transmitted from generation to generation. See also culture and fashion. trample Plooij Trample. Put foot or feet on body of reclining other. Sometimes accompanied by stamping. Rare pattern of social play. Cf. sit on and step on. 8. See

video. trampoline See hang and stamp. transfer Migrate permanently from one unit group to another. Females normally transfer from natal group to another during adolescence- at age 11 at Mahale but 1213 at Ta. No sexually mature males transfer at Mahale, thus, chimpanzees are patrilocal. Also for Gombe, Ta, Ngogo and the bonobos of Wamba, although female emigration varies across sites from 50 to 90%. Rarely, mother with juvenile offspring transfers. Adult male bonobos rarely tranfer. See emigrate and immigrate. Cf. visit. See video. T transport Nishida: Quadrupedally carry infant on belly or on back. Carry food between lips, in hand or foot, on back or head, in groin pocket or neck pocket, or drag food by hand. During play, juveniles often transport objects such as twig, fruit,

ant-nest etc. McGrews Play-start. See solicit play with object in mouth and ride. Bonobos also transport food similarly. 2. See video. transport bipedal When carrying much food or infant, walk on feet only. Bonobos carry sugar cane bipedally. See video. transport corpse of infant 193 transport corpse of infant Mother carries dead body of infant by hand, neck or groin pocket, or on back. May persist for 3 months, and corpse becomes mummy. Known from Bossou, Ndoki, Gombe, and Mahale. Also in bonobos. See video. See also video descend. transport food Transport of food occurs when: individual seeks to follow companion who has eaten and departed, seeks to eat in safe position in tree, and seeks to eat in shade. Bonobos also carry food. See video. See also

video transport in hand. transport in foot Carry grasped in foot object such as fruit, branch, stone, carcass, etc., usually in tree. Rare. 3. See video. transport in groin pocket Transport object clamped between thigh and abdomen. Goodalls Groin pocket. See also hold object in groin pocket. Kanos Groin pocket for bonobos. See video. transport in hand Common means of transport by hand of object such as fruit, branch, stone, carcass, etc. Stone carrying occurs during solo and social play, and occasionally before intimidation display or attack by stone throwing. See video. transport in mouth Carry object by mouth. Hold in mouth vines or twigs as materials of fishing tools and transport to fishing sites. Hold in mouth food such as leafy twigs, fruit during transport, or carcass.

Juvenile transports leafy branch in mouth to solicit play and during play. Mother, Xtina, held infants nape or back in mouth and transported for several months. Adolescent male, Darwin, also did so few times. Recorded at Bossou. Bonobos carry objects in mouth. See videos. transport in neck pocket Goodall Neck pocket, Plooij ONP. Transport object clamped between lower jaw and shoulder or chest. See also hold object in neck pocket. Rare in bonobos. See video. transport on back Carry dorsally object such as branch, carcass, piece of skin, etc. Infant carried on back from 3 months after birth. 3. See video. T 194 travel alone after childbirth transport on head or nape Plooij OOH. Carry piece of skin or carcass balanced or draped on head. Mother, Chausiku, regularly

carried newborn on head. Juvenile male, Michio, carried stone on nape in solo play. See also hold object on head. See video. See also video transport corpse of infant. transport on shoulder Carry large object such as carcass on shoulder. 3. See video. transport quadrupedal Transport object with hand, in mouth, neck pocket, on shoulder, or on head, while walking on four limbs. transport two offspring Mother simultaneously carries juvenile offspring on back and infant offspring on belly, or rarely carries both on back. Occurs in dangerous situations such as crossing large river. Bonobo mothers seem to carry two offspring more often than do chimpanzee mothers, as birth interval is shorter. 2. See video. transport with hand support Goodall Hand support. Plooijs Support. Mother supports infants back with

hand. Newborn or infant weekened by injury or illness typically carried this way, as gripping power of lower limbs is weak. Sometimes seen during ride dangling. Kanos Hand support for bonobos. 2. See video. transport with thigh support Goodall Thigh support: Method of supporting infant especially newborn, where the mother walks or runs with her thighs flexed, thus providing support for the infants back. Usually the mother moves with rounded back and takes short steps. Mother also carries newborn in flexed lower limbs when brachiating or hanging in tree. Kanos Thigh support in bonobos. T travel Go from one place to another. Goodalls Travel. Plooijs Travel. Include walk quadrupedal, walk bipedal, walk tripedal, wade, crutch, run, leap, somersault, climb, brachiate and descend. Kanos Travel in bonobos. See Susman

et al. for bonobo locomotor patterns. See videos. travel alone after childbirth Days or weeks after parturition, mother with newborn spends time alone, apart from other group members, probably avoiding harassment. At Wamba, travel and play 195 bonobo mother often accompanied by one or two males directly after giving birth. 3. travel and play Travel while playing socially or solo in locomotor-rotational play such as somersault and pirouette. During travel youngsters occasionally find something interesting such as hole in the ground, slope with full of dry leaves, animal carcass, etc. which elicits play. 3. See video. trifle with Tease for amusement captured small animal such as hyrax, squirrel, or leopard cub by hit, slap, flail, stamp, drag, drop, etc. Hirata et al.s Toy with. Victim sometimes killed in

process. Play face shown suggests that trifling sometimes fun. Cf. tease and abuse carcass. 3. See video. try play See solicit play. tug of war See pull object from opposing sides. tumble Lose balance and fall down slope. 3. See video. turn around Turn back to approaching individual. Goodalls Turn away: Mother may turn from her infant to prevent suckling or begging. Adults may also turn their backs on beggars. Kanos Turn back in bonobos. See video. turn away See turn around. turn face away Turn face away from partner who approaches to give kiss. 8. See video. turn face downward See lower head. turn face upward Incline head backward, while being groomed on face. Also in bonobos. 2. turn round See look back. T 19 twist turn

up lip Evert upper lip over muzzle, exposing upper gums and teeth for grooming. Cf. flip lip. 8. See video. twist When flesh difficult to detach by pulling from carcass possessed by another, wrench bone so that piece of carcass is detached. Mentioned in Boesch, but not in Goodall. 3. See video. T U Ugalla Study area of Pan troglodytes schweinfurthii in Tongwe East Forest Reserve, Tanzania. Research organized by J. Moore from 1985 to present. Also, Research conducted by T. Kano, H. Ogawa and G. Idani from 1991 to present. Another current research by A. Hernandez, A. Piel, and F. Stewart. Pioneer studies done by J. Itani and T. Kano in 19 19 7 and T. Nishida in 1975. unit group Social unit of chimpanzees, also called

community. Multi-male, multi-female group with group size of up to 150, occupying territory 10200 km2 in area. Adult females outnumber adult males. Shows fission and fusion grouping pattern. Females emigrate from natal unit group at age of about 11 years. Inter-group relationships are antagonistic. Unit group also main reproductive unit and extra-group paternity is rare at Gombe, Ta and Mahale. Bonobos have unit group of similar size, composition and male philopatry, but inter-group relations are peaceful and even fusion of two groups is indicated. See Fig. 37. uproot See pull out. upside-down suspension See hang upside-down by feet and hang upside-down by hands. urinate Excrete urine. See video. U urinate, prone Lies on belly and excretes urine. 8. See video. See also video defecate, prone. urinate quadrupedal Female

excretes urine in standing quadrupedal posture. Male does with adducted thighs, sometimes while walking. 3. See video. 197 198 use tool Fig. 37 Unit group: Procession of M group, more than 0 chimpanzees at one time urinate, sitting Urinate in sitting posture. 3. See video. U use tool Use detached object to change state or position of another object. At Mahale includes fish for carpenter ant, fish for termite, clip leaf, probe, drag branch, club, flail long object, throw stone or rock, solicit play with object in mouth, toy, probe nasal passage, squash leaf and make bed. Great local variation exists in tool-use. Tool-using repertoire of bonobos is limited: cover from rain, drag branch, throw branch and make bed at Wamba. 3. See video. See also video expel.

vacate Upon approach of more dominant individual, one of grooming party leaves partner and vacates position to dominant individual. Subordinate individual also may leave feeding patch when dominant approaches. Cf. supplant. 3. See video. ventral riding See ride ventral. ventro-dorsal copulation See copulate dorso-ventral. ventro-ventral copulation See copulate ventro-ventral. ventro-ventral embrace See embrace full. vertical climb See climb vertical. vertical leap See leap down. vertical slide See slide down vertically. vigilance Visual monitoring to collect information on possible threat of predation or of social conflict among conspecifics. See glance, look, monitor, and watch. visit Migrate temporarily from one unit group to another. Adolescent and young nulliparous adult females in estrus visit neighboring groups, then return to natal group when sexual skin deflates. Known from Gombe, 199 200 vomit

Ta and Mahale. Differs from transfer, which is permanent. 2. vocalize Call. Many calls are specific to contexts and express emotions. Includes bark, choke in tantrum, cough bark, grunt, hoo, hoot, huu, pant, pant-bark, pant-grunt, play-pant, scream, squeal in copulation, staccato call, waa bark, whimper, and wraa. Behavior accompanying vocalizations includes: clack teeth, cough, fart, hiccup, sputter, smack lip, sneeze, etc. Cf. make sound. vomit Plooij KOT. Rare, but infants do more often. See also reingest vomit and sickness. See video. See also video reingest vomit. W waa bark Goodall: Loud, sharp sound given in a variety of agonistic contexts. It is usually accompanied by an arm threat or more vigorous gestures. See also bark. See video. wad See wadge. wade Minimize contact with water, e.g. in river

crossing, and use stepping stones if available, when trees on both sides of river too far apart for overhead travel. If stepping stone unavailable, walk quadrupedal or bipedal in river, according to water depth. Cf. leap bipedal and leap quadrupedal. Bonobos wade similarly. 7. See video. wadge Chew and compress food between lower lip and teeth or between tongue and palate. Includes wadge without adding leaf and wadge by adding leaf. Neither Goodall nor Nishida et al. differentiated two forms. See also squeeze. Bonobos also wadge. Category 23. See videos. wadge by adding leaf Chew and compress meat, skin, honey, grub, egg, semen, or fruit with leaves between lower lip and teeth, or between tongue and palate. After juice extracted, remaining inedible parts such as hair, eggshell, seed,

leaf, etc. spat out. Chewed leaves from species not eaten solely, e.g. Saba comorensis, and even dry leaves are used. Fruits frequently wadged in this way at Mahale include Canthium crassum, Harungana madagascariensis, Parinari curatellifolia, Pseudospondias microcarpa, Psychotria peduncularis, W 201 202 walk in sloth position Syzigium guineense, Uapaca kirkiana and Uapaca nitida. Leaves chewed with fruit usually come from trees of eaten fruit. See video. wadge without adding leaf Extract juice by chewing and compressing some fruit, stalks of grasses or herb vines or young leaves between lower lip and teeth, or between tongue and palate. Unlike wadge by adding leaf, leaves are not added. After juice extraction, remaining inedible parts spat out. Piths of grasses such as Pennisetum purpureum, herbaceous vines such as Ipomoea rubens, or

young leaves of Brachystegia spiciformis chewed and sucked with wadge repeatedly taken out and returned to mouth. See video. wait for companion Goodall: An individual sets off, then looks back at a companion. If the latter is not following, the first stops to wait Ntologi, alpha male, often waited for adult males to follow. Bonobos wait similarly. See video. wait turn Wait for predecessor to vacate position, when target food obtainable only from specific site. Wait to groom target individual until another departs. Includes sit and stand quadrupedal. 3. See video. waiting posture See sit and stand quadrupedal, heel up for wait for companion, sit and stand quadrupedal for wait ones turn. walk backwards See retreat. walk bipedal Goodall Bipedal walk. Plooij WAB. Walk upright with much food

in arms, on muddy terrain, or to initiate charging display. Susman et al.s Bipedal walk in bonobos. See video. See also video stand bipedal. walk in sloth position Hang underneath branch using all hands and feet or combination of any three and locomote forwards. At Mahale part of solo W walk lame 203 play by immature chimpanzees. Bonobos also travel in this posture, which Kano called Sloth-walk. See video. walk lame See limp. walk quadrupedal on backs of hands Rare solo locomotor play. 9. See video. walk quadrupedal on knuckles Goodall Knuckle walking, Plooij WAQ. Quadrupedal walk with knuckles on ground or large horizontal bough. Kanos Walk. See video. walk quadrupedal on palms Walk quadrupedal, with open palm on thin horizontal branch. Rare when walking on ground. Susman

et al.s Palmigrade quadrupedalism. See videos. walk stealthily See creep. walk tripedal Walk with object in hand or walk with hand on back of another. Cf. embrace half. See video. Wamba Study area of Pan paniscus in the Equatorial Province of DRC, also called Luo Scientific Reserve. Research organized by T. Kano from 1973 to 1991 and from 1994 to 199. After civil war, research reorganized by T. Furuichi from 2002 to present. wash Seen only as wash colobus skin below. Wash includes whole process of dunk, rinse, and stamp. At least six bonobos of Lilungu wash aquatic herb or earthworms in water before consuming them. wash colobus skin Old adult male, Musa, once repeatedly dunked colobus pelt into running water, kept it immersed, rinsed it, stamped on

it on rock, repeating process several times. Musa then nibbled skin. Category 89. See Fig. 38. W 204 whimper Fig. 38 Wash colobus skin: Adult male, Musa, washed colobus skin in stream. Four chimpanzees watched rare activity watch Goodall: Gaze intently at what another is doing. If the face of the watching individual is very close to the other this is described as peering. van Hooffs Watch. Cf. stare fixedly, glance, look. Kanos Watch in bonobos. See video. watch water surface See look at water. wean Goodall Wean. Patterns of maternal rejection that encourage cessation of sucking and independence of infant from mother: Include drop infant, go ahead, ignore, reject infant, etc. Kanos Wean in bonobos. See video. wedge Infant forces itself between two others interacting socially, often

mother and grooming or mating partner. See also interfere copulation. 5. See video. W whimper Goodall: A whole series of soft, low pitched sounds, rising and falling in pitch, which may lead to crying and screaming Whimper by male infant may elicit presentation by estrous female, so serving as courtship display. Estrous female may whimper to adult male for copulatory solicitation. Cf. hoo. Bonobos call similarly. See video. whimper-scream 205 whimper-scream Goodall Cry: A combination of loud whimpers and tantrum screaming. This is commonly heard when an infant is separated from mother. At Mahale, weaning infant, especially if weaned when unusually young, whimper-screams continuously throughout mothers estrous periods. Lost call of bonobos seems distinct from that of chimpanzees in acoustic nature, and both mature and immature individuals emit

it. 3. See video. See also video throw temper tantrum. whisk fly with arm Shake arm to keep flies from converging on self or object, especially when mother carries dead infant. Cf. catch with hand. See video. whisk fly with leafy twig Sugiyama Fly whisk: Use leafy twig to fan away flies. Habitual at Ta and Budongo, present at Gombe. Recorded for Lomako bonobos. Absent at Mahale. winner support See support dominant. wipe with detached object Whiten et al. Leaf-napkin: Use leaves to clean body surface. Goodall: Use of leaves, straw, paper, etc. to wipe dirt from self or others Very rare at Mahale, with only handful of cases of rump-wiping with leaf. Customary at Kibale. Present at Assirik, Ta, and Goualougo. Penis-wiping by male after copulation customary

at Gombe and Budongo, but absent at Mahale. Wiping dirt from others not seen at Mahale. Cf. rub muzzle. Seen three times in bonobos at Wamba. See video. wipe with still-attached object See rub muzzle. wraa Goodall: ---a long drawn out, pure sounding call, neither high or low pitched Given by chimpanzees when confronted with new or disturbing objects in the environment - a human before they were habituated to humans, sometimes buffalo, python, dead bush pigs, etc See also bark. 5. See video. wrestle Goodall: Two or more youngsters grab hold of each other and often roll over as they bite, tickle, kick etc. without losing contact. Cf. play, rough and tumble. Kanos Wrestling play in bonobos. See video. W 20 wrist towards wrestle bipedal Wrestle in

bipedal stance like Japanese sumo. Lasts less than 5 s before return to usual wrestle in sitting and reclining postures. 7. See video. wrestle with fingers van Hooff Handwrestle. Goodall: Gentle fondling, holding, squeezing, tickling of one hand of intended play partner. This is how a mature individual typically seeks to initiate play with another adult. It may lead to a vigorous bout of social play. At Mahale, initiates social play in adult males and between adult males and females; overall most common pattern of social play among adults. Absent in bonobos. See video. wriggle Writhe or squirm from being heavily bitten by ants, etc. Category 17. See video. wrist towards See reach wrist toward. W Y Yalosidi Study area of Pan paniscus in Equateur Province of DRC.

Research done by T. Kano and colleagues from 1973 to 1977. yawn Goodall Yawning. Plooij YAW. Kano Yawn for the bonobos. See videos. Y 207 Discussion Our list contains 891 behavioral patterns of Mahale chimpanzees, 45 patterns that have been recorded at other sites but not at Mahale, 91 nouns concerning chimpanzees and their study sites, and 357 synonyms, totaling 1,384 terms. Thus, we have added more than 500 new terms to the previous publication. The last publication was the product of research that spanned more than 30 years. Why so many patterns have been added must be explained. First, we did not use videographic records extensively for the last publication. Although we referred to video records taken by ANC Production and East Company, they were not major

sources for the behavioral patterns. This time videotapes that add up to more than 1,500 h since 1999 were played back and watched again and again. This procedure has brought a qualitatively, as well as quantitatively, higher level of data scrutiny than before. Second, the multi-authored article that was published simultaneously with our last paper in 1999 provided fruitful discussion with many colleagues from different research sites directly or via email and alerted us to subtle differences in behavioral patterns. Third, and most important, the behavioral flexibility of chimpanzees is responsible for major revision of this paper. A comprehensive dictionary of human actions would amount to several thousand pages because human behavior is flexible and variable. If we take the number of chimpanzees we have intimately observed only

250 individuals or so for consideration, 180 pages for this encyclopedia would be remarkable. As we pointed out already, behavior as simple as social scratch is not only absent at some sites, but also the behavioral pattern of social scratch differs across sites where it exists. Chimpanzees are cultured animals whose behavior is modified during maturation after birth, but also changes even during adulthood. This was illustrated by the changes shown by migrants in the features of the grooming hand-clasp. From the viewpoint of cultural diversity, we conclude that we have currently reached only the starting point of understanding of behavioral diversity and flexibility of chimpanzees. Currently it is difficult to classify behavioral patterns into Categories 19 since detailed glossaries of behavior are available from only two populations

of chimpanzees, Gombe and Mahale, both belonging to subspecies of P. t. schweinfurthii and T. Nishida et al., Chimpanzee Behavior in the Wild: An Audio-Visual Encyclopedia, DOI 10.1007/978-4-431-53895-0 10, Springer 2010 209 210 Discussion one population of bonobos at Wamba. In this paper, we tentatively allocated categories to all the behavioral patterns on the basis of the minimal status and maximum status. For example, fish is known from all subspecies of chimpanzees, so it is very likely to be a chimpanzee universal, but it also is known from some indigenous humans, while it still unknown from bonobos. In this case, is the minimal status of fish, and is a possible maximum status. Then, we allocate fish to Categories 13. Since the minimum status is the only category that

has so far been confirmed for each behavioral pattern, we here use it as the principle means of summarizing our Table 1 Numbers of behavioral patterns in each category and its phylogenetic position Categories No. of patterns Subtotal Phylogenetic positions 1 12 2 13 23 3 15 25 35 45 5 1 2 3 17 27 37 57 7 18 28 38 78 8 19 2 4 30 127 50 31 44 12 5 58 2 2 9 5 24 5 17 3 34 1 22 13 4 2 5 44 2 39 89 9 Total 2 31 19 891 2 4 157 Likely shared by LCA Likely shared by common ancestors of Pan 125 Likely shared by Pan troglodytes 79 Likely shared by Pan troglodytes schweinfurthii 43

Likely to be Mahale customs 77 Likely to be M group customs 92 Likely to be habits of a few M group chimpanzees 54 Likely to be idiosyncracies of M group chimpanzees 891 Discussion 211 results. Thus, we have obtained 2 3 behavior patterns likely shared by the common ancestors of Homo and Pan, 157 patterns likely to be shared by commons ancestors of chimpanzees and bonobos, 12 patterns likely to be chimpanzee universals, 79 patterns likely shared by eastern subspecies, 43 patterns likely to be Mahale cultures, 77 patterns likely to be M group cultures, 93 patterns likely to be habitual among a few M group chimpanzees, and 54 patterns likely to be idiosyncracies of M group chimpanzees. We are sure that the last common ancestors of

Homo and Pan had such characteristics as drag branch, play aeroplane, female transfer, and many other patterns, but we are less sure that they engaged in patrol, ostracize, fish and many other patterns. Probably, more important, we found many candidates of cultural behavioral patterns, namely 120 patterns, in addition to those 44 patterns that have not been recorded at Mahale but have been seen elsewhere. Although this needs confirmation by future detailed, long-term studies, we speculate that like current apes and humans, our common ancestors enjoyed rich cultural diversity in addition to common basic genetic tendencies. However, most impressively, chimpanzees show a tremendous extent of diversity of individual or idiosyncratic behavioral patterns: 147!. This is the ethnographic diversity of chimpanzees that deserves much more attention as a source

of new patterns. We hope that by viewing the video clips carefully as well as reading the glossary, researchers at other sites will discover much new material for discussing the behavioral diversity of chimpanzees living in the natural environment. Rich flesh will be put on the bones of Table 1 and full interpretation will be given only after detailed information is gathered in many other sites, in particular, for Pan troglodytes troglodytes and P. t. vellerosus and Pan paniscus. Acknowledgments We thank Tanzania Commission for Science and Technology, Tanzania Wildlife Research Institute, and Tanzania National Parks for permission to do the field research, and Mahale Mountains National Park and Mahale Mountains Wildlife Research Centre for logistic support. Our gratitude goes to Hosea Y. Kayumbo, Costa Mlay, George Sabuni,

Charles Mlingwa, Erasmus Tarimo, Edeus Massawe, A.K. Seki, M. Mbaga and their staff for their friendship and encouragement; We thank Michio Nakamura, Gaku Ohashi, Tetsuya Sakamai, Crickette Sanz and Dave Morgan for allowing us to use their precious video clips; Tamotsu Asou, Richard Byrne, Satoshi Hirata, Michio Nakamura, Yukimaru Sugiyama, Rikako Tonooka, Caroline Tutin and Moyo Uehara for allowing us to use their precious photos or illustrations; Tamotsu Asou, Miho Nakamura, Mitsue Matsuya and the late Masayasu Mori for allowing us to use their videos for study; Toshimichi Nemoto and Asami Kanayama for helping us in various ways in Tanzania. We thank our colleagues who have studied chimpanzees at Mahale for sharing information on the behavioral patterns, in particular, the late Kenji Kawanaka, the late Shigeo Uehara, Hitoshige

Hayaki, John Mitani, Linda Marchant, Kazuhiko Hosaka, Akiko Matsumoto-Oda, Michio Nakamura, Noriko Itoh, Tetsuya Sakamaki and Nadia Corp. We are indebted to our field assistants, in particular, Ramadhani Nyundo, Rashidi Kitopeni, Rashidi Hawazi, Mtunda Hawazi, Mosi B. Kasagula, Hamisi B. Kasagula, Mosi Hamisi, Kabumbe Athumani, and Mosi Matumla. We are indebted to Naomi Miyamoto for assistance in editing 212 Discussion video clips. We thank Frans de Waal for constructive comments on this project and Aiko Hiraguchi and Kaoru Hashimoto for encouragement. Field research was supported by funds from the JSPS International Scientific Research Program, Global Environment Research Fund of the Ministry of Environment, the Leakey Foundation, Grant-in-Aid for JSPS Fellows. References Adang OMJ Teasing in young chimpanzees. Behaviour 88:98122 Albrecht H, Dunnett SC Chimpanzees in West Africa. Piper,

Munich Alp R Stepping-sticks and seat-sticks: new types of tools used by wild chimpanzees in Sierra Leone. Am J Primatol 41:4552 Arcadi AC, Daniel R, Boesch C Buttress drumming by wild chimpanzees: temporal patterning, phrase integration into loud calls, and preliminary evidence for individual distinctiveness. Primates 39:505518 Arcadi AC, Wrangham RW Infanticide in chimpanzees: review of cases and a new withingroup observation from the Kanyawara study group in Kibale National Park. Primates 40: 337351 Assersohn C, Whiten A, Tiwede AT, Tinka J, Karamagi J Use of leaves to inspect ectoparasites in wild chimpanzees: a third cultural variant? Primates 45:255258 Azuma S, Toyoshima A Progress report of the survey of chimpanzees in their natural habitat, Kabogo Point Area, Tanganyika. Primates 3: 170 Basabose AK, Yamagiwa J Factors affecting nesting

site choice in chimpanzees at Tshibati, Kahuzi-Biega National Park: influence of sympatric gorillas. Int J Primatol 23:2 3282 Beck BB Animal tool behavior. Garland SPTM, New York Berdecio S, Nash LT Chimpanzee visual communication. Arizona State University, Tempe, AZ, Anthropological Research Papers No. 2 Bermejo M, Illera G Tool-set for termite-fishing and honey extraction by wild chimpanzees in the Lossi Forest, Congo. Primates 40: 19 27 Bermejo M, Illera G, Sabater Pi J Animals and mushrooms consumed by bonobos: New records from Lilungu, Zaire. Int J Primatol 15:879898 Bethell E, Whiten A, Muhumuza G, Kakura J Active plant food division and sharing by wild chimpanzees. Primate Rep 5: 771 Biro D, Sousa C, Matsuzawa T Ontogeny and cultural propagation of tool use by wild chimpanzees at Bossou, Guinea:

case studies of nut cracking and leaf folding. In: Matsuzawa T, Tomonaga M, Tanaka M Cognitive development in chimpanzees. Springer, Tokyo, pp 47 508 Bodenheimer FS Insects as human food. Dr. W. Junk Publishers, Hague Boehm C Hierachy in the forest. Harvard University Press, Cambridge, MA Boesch C The effect of leopard predation on grouping patterns in forest chimpanzees. Behaviour 117:220242 Boesch C Handedness in wild chimpanzees. Int J Primatol 12:541558 Boesch C Teaching in wild chimpanzees. Anim Behav 41:530532 Boesch C Innovation in wild chimpanzees. Int J Primatol 1:11 Boesch C The real chimpanzee. Cambridge University Press, Cambridge Boesch C, Boesch H Tool use and tool making in wild chimpanzees. Folia Primatol 54:8 90 213 214 References Boesch C, Boesch-Achermann H The chimpanzees of the Ta Forest.

Oxford University Press, Oxford Boesch C, Head J, Robbins MM Complex tool sets for honey extraction among chimpanzees in Loango National Park, Gabon. J Hum Evol 5:5 05 9 Boesch C, Hohmann G, Marchant L Behavioural diversity in chimpanzees and bonobos. Cambridge University Press, Cambridge Bogart SL, Pruetz JD Ecological context of savanna chimpanzees termite fishing at Fongoli, Senegal. Am J Primatol 70: 05 12 Bogart SL, Pruetz JD, Kante D Fongoli chimpanzees eats banded mongoose. Pan Afr News 15:1517 Bonnie KE, de Waal FBM Affiliation promotes the transmission of a social custom: handclasp grooming among captive chimpanzees. Primates 47:2734 Brewer SM, McGrew WC Chimpanzee use of a tool-set to get honey. Folia Primatol 54:100104 Bygott JD Cannibalism among wild chimpanzees. Nature 238:410411 Bygott JD Agonistic behaviour and

dominance in wild chimpanzees. PhD thesis, University of Cambridge, Cambridge Byrne R, Whiten A Machiavellian intelligence. Clarendon, Oxford Call J, Tomasello M The gestural communication of apes and monkeys. Lawrence Erlbaum Associates, London Call J, Tomasello M The gestural repertoire of chimpanzees. In: Call J, Tomasello M The gestural communication of apes and monkeys. Lawrence Erlbaum Associates, London, pp 1739 Caro TM, Hauser MD Is there teaching in nonhuman animals? Q Rev Biol 7:151174 Chimpanzee Sequencing and Analysis Consortium Initial sequence of the chimpanzee genome and comparison with the human genome. Nature 437: 987 Clark AP Rank differences in the production of vocalizations by wild chimpanzees as a function of social context. Am J Primatol 31:159179 Constable JL, Ashley MV, Goodall J, Pusey AE Noninvasive paternity assignment in

Gombe chimpanzees. Mol Ecol 10:12791300 Corbalis MC From hand to mouth: The origins of language. Princeton University Press, Princeton Corp N, Hayaki H, Matsusaka T, Fujita S, Hosaka K, Kutsukake N, Nakamura M, Nakamura M, Nishie H, Shimada M, Zamma K, Wallauer W, Nishida T Prevalence of muzzle-rubbing and hand-rubbing behavior in wild chimpanzees in Mahale Mountains National Park, Tanzania. Primates 50:184189 Deblauwe I, Guislain P, Dupain J, van Elsacker L Use of a tool-set by Pan troglodytes troglodytes to obtain termites in the periphery of the Dja Biosphere Reserve, southeast Cameroon. Am J Primatol 8:1191119 Devos C, Gatti S, Levrero F New record of algae feeding and scooping by Pan t. troglodytes at Lokoue Bai in Odzala National Park, Republic of Congo. Pan Afr News 9:1921 Eibl-Eibesfeldt

I Love and hate. The natural history of behavior patterns. Holt, Rinehart and Winston, New York Enomoto T Bonobos. Maruzen, Tokyo Fawcett K, Muhumuza G Death of a wild chimpanzee community member: possible outcome of intense sexual competition? Am J Primatol 51:243247 Fay JM, Carroll RW Chimpanzee tool use for honey and termite extraction in Central Africa. Am J Primatol 34:309317 Feistner ATC, McGrew WC Food-sharing in primates: a critical review. In: Seth PK, Seth S Perspectives in primate biology, vol. 3. Today and Tomorrows, New Delhi, pp 213 Fowler A, Sommer Subsistence technology of Nigerian chimpanzees. Int J Primatol 28:9971023 Fruth BI, Hohmann G, Beuerlein MM, McGrew WC Grooming hand clasp by bonobos of Lui Kotal, Democratic Republic of Congo. Pan Afr News 13: 8 References 215

Fujimoto M, Shimada M Newly observed predation of wild birds by M group chimpanzees at Mahale, Tanzania. Pan Afr News 15:232 Furuichi T Social interactions and the life history of female Pan paniscus in Wamba, Zaire. Int J Primatol 10:173197 Furuichi T, Hashimoto C Ground beds of chimpanzees in the Kalinzu Forest, Uganda. Pan Afr News 7:2 28 Furuichi T, Thompson J The bonobos: behavior, ecology, and conservation. Springer, New York Ghiglieri MP The chimpanzees of Kibale Forest. Columbia University Press, New York Gomez J-C, Martin-Andrade B Fantasy play in apes. In: Pellegrini AD, Smith PK The nature of play. Guilford, New York, pp 139172 Goodall J The behavior of free-living chimpanzees in the Gombe Stream Reserve. Anim Behav Monogr 1:1 1311 Goodall J In the shadow of

man. Collins, London Goodall J Cultural elements in a chimpanzee community. In: Menzel EW Precultural primate behavior. S. Karger, Basel, pp 144184 Goodall J Infant killing and cannibalism in free-living chimpanzees. Folia Primatol 22:259282 Goodall J The chimpanzees of Gombe. Harvard University Press, Cambridge, MA Goodall J Glossary of chimpanzee behaviors. Jane Goodall Institute, Tucson, AZ Goodall J Unusual violence in the overthrow of an alpha male chimpanzee at Gombe. In: Nishida T, McGrew WC, Marler P, Pickford M, de Waal FBM Topics in primatology 1. Human origins. University of Tokyo Press, Tokyo, pp 131142 Goodall J, Bandora A, Bergmann E, Busse C, Matama H, Mpongo E, Pierce A, Riss D Intercommunity interactions in the chimpanzee population of the Gombe National Park. In: The Great Apes, DA Hamburg

and ER McCown, Benjamin/Cummings, Menlo Park, pp.1353 Graham CE Menstrual cycle physiology of the great apes. In: Graham CE Reproductive biology of the great apes. Academic, New York, pp 28 303 Hamai M, Nishida T, Takasaki H, Turner LA New records of within-group infanticide and cannibalism in wild chimpanzees. Primates 33:1511 2 Harris JR The nurture assumption. Brockman, New York Hasegawa T Sex differences in ranging patterns. In: Nishida T The chimpanzees of the Mahale Mountains. University of Tokyo Press, Tokyo, pp 99114 Hasegawa T, Hiraiwa-Hasegawa M, Nishida T, Takasaki H New evidence of scavenging behavior of wild chimpanzees. Curr Anthropol 24:231232 Hashimoto C, Furuichi T Possible intergroup killing in chimpanzees in the Kalinzu Forest, Uganda. Pan Afr News 12:35 Hashimoto C, Furuichi T Frequent copulations by females

and high promiscuity in chimpanzees in the Kalinzu Forest, Uganda. In: Newton-Fisher NE, Notman H, Paterson JD, Reynolds Primates of western Uganda. Springer, New York, pp 247257 Hashimoto C, Furuichi T, Tashiro Y Ant dipping and meat eating by wild chimpanzees in the Kalinzu Forest, Uganda. Primates 41:103108 Hashimoto C, Tashiro Y, Hibino E, Mulabwa M, Yangozene K, Furuichi T, Idani G, Takenaka O Longitudinal structure of a unit-group of bonobos: male philopatry and possible fusion of unit-groups. In: Furuichi T, Thompson J The bonobos: behavior, ecology, and conservation. Springer, New York, pp 107119 Hayaki H Social play of juvenile and adolescent chimpanzees in the Mahale Mountains National Park, Tanzania. Primates 2:3433 0 Hayaki H Copulation of adolescent male chimpanzees, with special reference to the influence of adult

males, in the Mahale Mountains National Park, Tanzania. Folia Primatol 44:1481 0 Hernandez-Aguilar RA, Moore J, Pickering TR Savanna chimpanzees use tools to harvest the underground storage organs of plants. Proc Natl Acad Sci USA 104:1921019213 21 References Hewes GW Primate communication and the gestural origin of language. Cur Anthropol 14:524 Hicks TC, Fouts RS, Fouts DH Chimpanzee tool use in the Ngotto Forest, Central African Republic. Am J Primatol 5:221237 Hiraiwa-Hasegawa M Sex differences in behavioral development at Mahale. In: Heltne PG, Marquardt LA Understanding chimpanzees. Harvard University Press, Cambridge, MA, pp 104115 Hiraiwa-Hasegawa M, Byrne RW, Takasaki H, Byrne JME Aggression toward large carnivores by wild chimpanzees of Mahale Mountains National Park, Tanzania. Folia Primatol 47:813 Hirata S, Myowa M, Matsuzawa T Use of leaves as

cushions to sit on wet ground by wild chimpanzees. Am J Primatol 44:215220 Hirata S, Yamakoshi G, Fujita S, Ohashi G, Matsuzawa T Capturing and toying with hyraxes by wild chimpanzees at Bossou, Guinea. Am J Primatol 53:9397 Hockings KJ, Anderson JR, Matsuzawa T Flexible feeding on cultivated underground storage organs by rainforest-dwelling chimpanzees at Bossou, West Africa. J Hum Evol 58:227233 Hockings KJ, Humle T, Anderson JR, Biro D, Sousa C, Ohashi G, Matsuzawa T Chimpanzees share forbidden fruit. PLoS One 2:e88 Hohmann G Association and social interactions between strangers and residents in bonobos. Primates 42:9199 Hohmann G, Fowler A, Sommer V, Ortmann S Frugivory and gregariousness of Salonga bonobos and Gashaka chimpanzees: the influence of abundance and nutritional quality of fruit. In: Hohmann G, Robbins MM,

Boesch C Feeding ecology in apes and other primates. Cambridge University Press, Cambridge Hohmann G, Fruth B Field observations on meat sharing among bonobos. Folia Primatol 0: 225-229 Hohmann G, Fruth B Culture in bonobos? Between-species and within-species variation in behavior. Curr Anthropol 44:5 3571 Hohmann G, Fruth B New records of prey capture and meat eating by bonobos at Lui Kotale, Salonga National Park, Democratic Republic of Congo. Folia Primatol 79:103110 van Hooff JARAM A comparative approach to the phylogeny of laughter and smiling. In Hinde RA Non-verbal communication. Cambridge University Press, Cambridge, pp 3 2439 van Hooff JARAM A structural analysis of the social behavior of a semicaptive group of chimpanzees. In: von Cranach M, Vine I Social communication and movement. Academic Press, London, pp 751

2 Hopkins WD, Morris RD Handedness in great apes: A review of findings. Int J Primatol 14:125 Hosaka K, Matsumoto-Oda A, Huffman MA, Kawanaka K Responses of chimpanzees to dead conspecifics. Primate Res 1:115 Hosaka K, Nishida T, Hamai M, Matsumoto-Oda A, Uehara S Predation of mammals by the chimpanzees of the Mahale Mountains, Tanzania. In: Galdikas BMF, Briggs N, Sheeran LK, Shapiro GL, Goodall J All apes great and small, vol. 1: African apes. Plenum/ Kluwer, New York, pp 107130 Huffman MA Current evidence for self-medication in primates: a multidisciplinary perspective. Yearb Phys Anthropol 40:171200 Huffman MA, Kalunde MS Tool-assisted predation on a squirrel by a female chimpanzee in the Mahale Mountains, Tanzania. Primates 34:9398 Humle T New record of fishing for termites by the chimpanzees of

Bossou, Guinea. Pan Afr News:34 Humle T, Matsuzawa T Ant-dipping among the chimpanzees of Bossou, Guinea, and some comparisons with other sites. Am J Primatol 58:133148 Hunt KD, Cant JG, Gebo DL, Rose MD, Walker SE, Youlatos D Standardized descriptions of primate locomotor, and postural modes. Primates 37:3 3387 References 217 Hunt KD, McGrew WC Chimpanzees in the dry habitats of Assirik, Senegal and Semliki Wildlife Reserve, Uganda. In: Boesch C, Hohmann G, Marchant LF Behavioural diversity in chimpanzees and bonobos. Cambridge University Press, Cambridge, pp 3551 Idani G Relations between unit-groups of bonobos at Wamba, Zaire: encounters and temporary fusions. Afr Study Monogr 11:15318 Idani G Social relationships between immigrant and resident bonobo females at Wamba. Folia Primatol 57:8395 Idani G Function of peering behavior among bonobos

at Wamba. Primates 3:377383 Ihobe H Interspecific interactions between wild pygmy chimpanzees and red colobus. Primates 31:109112 Ihobe H Life span of chimpanzee beds at the Mahale Mountains National Park, Tanzania. Pan Afr News 12:1012 Inagaki H, Tsukahara T A method of identifying chimpanzee hairs in lion feces. Primates 34:109112 Ingmanson EJ Tool-using behavior in wild Pan paniscus: social and ecological considerations. In: Russon AE, Bard KA, Parker ST Reaching into thought. Cambridge University Press, Cambridge, pp 190210 Inoue E, Inoue-Murayama M, Vigilant L, Takenaka O, Nishida T Relatedness in wild chimpanzees: influence of paternity, male philopatry, and demographic factors. Am J Phys Anthropol 137:25 2 2 Itani J Distribution and adaptation of chimpanzees in an arid area. In Hamburg DA, McCown ER The great apes. Benjamin/Cummings, Menlo

Park, CA, pp 5571 Itoh N, Nishida T Chimpanzee grouping patterns and food availability in Mahale Mountains National Park, Tanzania. Primates 48:879 Itoh N, Sakamaki T, Hamisi M, Kitopeni R, Bunengwa M, Matumla M, Athumani K, Mwami M, Bunengwa H A new record of invasion into the center of M group territory. Pan Afr News:810 Iverson JM, Goldin-Meadow S Why people gesture when they speak. Nature 39:228 Izawa K Unit-groups of chimpanzees and their nomadism in the savanna woodland. Primates 11:14 Jones C, Sabater Pi J Sticks used by chimpanzees in Rio Muni, west Africa. Nature 223:100101 Jones C, Sabater Pi J Comparative ecology of Gorilla gorilla and Pan troglodytes in Rio Muni, west Africa. S. Karger, Basel Kahlenberg S, Emery Thompson M, Wrangham RW Female competition over