The siphonotretide brachiopod Schizambon from the Early Ordovician of South China: ontogeny and affinity

Abstract: Abstract Schizambon is one of the earliest and most distinctive genera in the Order Siphonotretida. However, current knowledge of siphonotretide phylogeny and early evolution requires understanding of their earliest ontogeny. In this study, the new species Schizambon tongziensis from the Tungtzu Formation at Honghuayuan section in Guizhou Province, South China is described; it is also the first record of Schizambon in South China. The new well-preserved specimens shed light on the ontogeny, palaeobiogeographical distribution, phylogeny and early evolution of siphonotretide brachiopods. Characters of metamorphic shells on both valves, outlined by distinct halos, show that Schizambon had a planktotrophic ‘paterinide-type’ larva, typical of many early Palaeozoic brachiopods. Based on the newly obtained data it is inferred that the free-swimming larva of Schizambon settled directly on the ventral side of the body, and this type of ontogeny is probably true for other siphonotretides. The divergence of Schizambon from the main siphonotretide lineage probably happened prior to the Guzhangian Age, well before the occurrence of major biodiversification within the order.

linguliformea
siphonotretida
tremadocian
ontogeny
evolution
planktotrophic larva


Healed injuries, ontogeny and scleritome construction in a Late Ordovician machaeridian (Annelida, Aphroditiformia)

Abstract: Abstract Machaeridians are armoured annelids that were morphologically diverse during the Palaeozoic. The scleritome developed from fleshy protrusions at the base of each parapodium, with alternating segments giving rise to differentiated inner and outer shell plates. The elytra-like anatomy of the shell-bearing soft tissues and distinctive jaw apparatus support an affinity of machaeridians with aphroditacean scaleworms. While frequently found as disarticulated remains, whole scleritomes of machaeridians are rare. Only a few species are represented by multiple articulated individuals, rendering many aspects of our knowledge of scleritome construction and growth uncertain. Here we describe a collection of articulated scleritomes of the plumulitid machaeridian Plumulites tafennaensis Chauvel from the Upper Ordovician (Katian) Upper Tiouririne Formation of Morocco that includes individuals representing both adult and juvenile stages. These scleritomes shed light on the morphology of the anterior region in plumulitids and show an increase in the number of shell plates with increasing size, suggesting that unlike some derived lepidocoleids, at least some plumulitids added segments repeatedly during ontogeny. Shell plates with a distinctive umbo have previously been assigned exclusively to the anterior region, but our material shows that they are present along the scleritome, with corresponding sclerites on the opposite side of the skeleton showing normal morphologies. We suggest that these plates are abnormal and best explained as healed injuries, replacing plates that had been previously removed from the body, demonstrating the capacity for machaeridians to regenerate lost body parts, a feature that is widespread in living annelids.


A review of the glacial environment arthropod trace fossils Umfolozia and Warvichnium with the description of new ichnotaxa

Abstract: Abstract Trace fossils are important records of the presence and behaviour of animals in the past, especially in deposits where few body fossils are preserved. They tend to provide the main palaeobiological record for past glacial environments, and are thus very important for understanding the ecology of these palaeoenvironments. Two ichnogenera are common in glacial sediments: Umfolozia and Warvichnium. Both Umfolozia and Warvichnium are attributed to arthropods; the former occurs across the Late Palaeozoic Ice Age (LPIA) and the latter occurs during both the LPIA and the Quaternary glacial event. Here, we review the stratigraphic record, ichnotaxonomy, palaeoecology, palaeoenvironments and possible tracemakers of these two ichnogenera. In addition to assessing morphological ichnotaxobases, we use multivariate morphometric analysis and inferential statistical tests to support our ichnotaxonomic revisions. The diagnosis of Umfolozia is revised, with the description of a new ichnospecies, Umfolozia terere, and the emendation of Umfolozia riojana (formerly Umfolozia longula). We revised Warvichnium, maintaining the original Warvichnium ulbrichi and allocating two morphologies to the ichnogenus Irichnus: Irichnus saltatorius and I. paripinnatus isp. nov. These trace fossils record several behaviours, and have implications for palaeoenvironmental interpretations and the evolution of invertebrate ecosystems. Using Umfolozia and Warvichnium as case studies, we demonstrate the application of morphometric analysis in ichnotaxonomy and highlight this as a tool that may be applied to other trace-fossil groups.

arthropod trackway
ichnotaxonomy
crustacean trace fossil
statistics
morphometrics


A description of the palate and mandible of Youngina capensis (Sauropsida, Diapsida) based on synchrotron tomography, and the phylogenetic implications

Abstract: Abstract The late Permian reptile Youngina capensis (c. 254 Ma) is a non-saurian neodiapsid whose anatomy has been used to represent the reptilian condition prior to the divergence of Sauria (crown-group reptiles). However, despite being first described over 100 years ago, the anatomy of Youngina remains incompletely documented. Here we use synchrotron x-ray micro-computed tomography to document new features of the palate, braincase and mandible of Youngina. New observations include an anteriorly bifurcating vomer, dentition on the ventral surface of the parasphenoid body and cultriform process, and a strongly convex coronoid eminence. Our anatomical observations suggest that Youngina may represent a more stemward lineage among non-saurian neodiapsids and this is supported by our phylogenetic analysis, which places Youngina as an early diverging neodiapsid. Our research will benefit future studies on saurian origins by providing improved constraints on neodiapsid anatomy prior to the divergence of the reptilian crown group.

neodiapsida
sauria
phylogeny
synchrotron tomography
reptile evolution
permian


A micromorphological analysis of Bolonia lata Meunier from the Cretaceous of the Neuquén Basin (Patagonia, Argentina): new insights into the tracemaker

Abstract: Abstract Polychaetes, echinoids and gastropods have been proposed as tracemakers for Bolonia Meunier, an elongate positive epirelief trace fossil characterized by two lobes composed of biserial, subtriangular pads and a mostly heart-shaped cross-section. Here, the internal structure and micromorphology of Bolonia are described for the first time using serial thin sections from shallow-marine Lower Cretaceous intervals of the Agrio Formation (Neuquén Basin, Argentina). In vertical thin sections, a conical structure of variable width extends centrally from the base to the top of the trace. In horizontal thin sections, grains are reoriented and reworked, forming a central zigzag structure. The organic matter and clays form grain aggregates representing the internal structure of individual pads of the trace fossil. The sediment has mostly been reoriented, with grains making up subcircular pods. We interpret the central conical structure as the collapse of the medial zone of the trace fossil, which could have destroyed drain tubes, especially in the absence of early cementation. Studying horizontal thin sections, we reconstruct the diagnostic biserial pads as biserial concave structures. These are interpreted as backfilling structures produced by the aboral and lateral spines of irregular echinoids, which worked together to compact the sediment anteroposteriorly and moved alternatively, explaining the zigzag pattern in the menisci. We propose Spatangoidea or Cassiduloidea (or a closely related group) with a similar burrowing ability and spine movement as tracemakers of these Bolonia specimens.

patagonia
palaeoecology
ichnology
echinoderm
early cretaceous
thin section


First report of silicified wood from a late Pennsylvanian intramontane basin in the Pyrenees: systematic affinities and palaeoecological implications

Abstract: Abstract The first anatomically preserved wood specimens of an upland Carboniferous flora from the Iberian Peninsula are reported from the Erillcastell Basin (Eastern Pyrenees, Catalonia, Spain). Two taxa are described, a calamitacean Equisetales (Arthropitys sp.) and a Cordaitales (Dadoxylon sp.). The Arthropitys specimen has fusiform multiseriate rays composed of square parenchyma cells with conspicuous uniseriate or multiseriate simple pits. These pits are located near the transverse walls and occasionally in the tangential walls. The tracheids vary in lumen size, with scalariform-bordered pits on their radial walls and multiseriate pits in their cross-field areas. The Dadoxylon specimen commonly has uniseriate fusiform rays. The tracheids are long, with a square shape in transverse section. Their radial walls bear araucarian pitting with a uniseriate to triseriate arrangement. The pits are circular with a spindle-shaped aperture. Comparison of the Erillcastell specimens with coeval species from Europe indicates that they could belong to new species. The good preservation of the new fossil wood yields significant palaeoenvironmental information. The lack of marked growth rings in both specimens and the presence of tyloses in Dadoxylon suggest that the climate in the intramontane basins of the Pyrenees was slightly seasonal towards the end of the Carboniferous. This contrasts with the marked seasonality of coeval lowland basins. Such upland habitats may have enhanced the survival of plants adapted to humid conditions in a global context of increasing aridity.

cordaites
calamites
silicified wood
upland flora
pyrenees
iberian peninsula


Trilobites of Thailand's Cambrian–Ordovician Tarutao Group and their geological setting

Abstract: Abstract Tuff-bearing upper Cambrian to lowermost Ordovician strata on Ko Tarutao island, Satun province, southernmost peninsular Thailand, contain a rich trilobite fauna relevant to global biostratigraphy, peri-Gondwanan palaeogeography and shifting evolutionary mode. This area of Sibumasu, a lower Palaeozoic marginal Gondwanan terrane, is shown to have been closely associated with Australia, North China (Sino-Korea) and other continental fragments from the supercontinent's northern equatorial sector, including South China at that time. Shared faunas also suggest a Kazakhstani and Laurentian association. Collections from eight sections yielded 10 newly discovered species and one new genus from ancient shoreface and inner shelf siliciclastic deposits. With the new taxa and revision of taxa known previously, we refine the age of the upper two formations of the Tarutao Group to the middle of Cambrian Stage 10, and lower–middle Tremadocian. Two biozones are erected for Sibumasu: the Eosaukia buravasi Zone, encompassing all Cambrian sections from Ko Tarutao, and the Asaphellus charoenmiti Zone, encompassing the Tremadocian fauna discussed herein. The new genus is Tarutaoia and new species are Tsinania sirindhornae, Pseudokoldinioidia maneekuti, Pagodia? uhleini, Asaphellus charoenmiti, Tarutaoia techawani, Jiia talowaois, Caznaia imsamuti, Anderssonella undulata, Lophosaukia nuchanongi and Corbinia perforata. Other taxa reported for the first time from Tarutao are Mansuyia? sp., Parakoldinioidia callosa Qian, Pseudagnostus sp., Homagnostus sp., Haniwa mucronata Shergold, Haniwa sosanensis? Kobayashi, Lichengia simplex Shergold, Pacootasaukia sp., Wuhuia? sp., Plethopeltella sp., Apatokephalus sp., Akoldinioidia sp. 1 and Koldinioidia sp.

thailand trilobita cambrian ordovician sibumasu gondwana


The last Baru (Crocodylia, Mekosuchinae): a new species of ‘cleaver-headed crocodile’ from central Australia and the turnover of crocodylians during the Late Miocene in Australia

Abstract: Abstract Baru is a genus that includes several large mekosuchine crocodylians from the Oligo-Miocene of Australia. Here we describe Baru iylwenpeny sp. nov. from a large sample of cranial bones from the Upper Miocene Alcoota Local Fauna of the Northern Territory. Baru iylwenpeny can be diagnosed by several autapomorphies that include, but are not limited to: extreme reduction of the pneumatic foramina associated with the diverticula that invade the bones of the suspensorium; and enlargement of the postcaniniform maxillary teeth resulting in crowding of the postcaniniform alveoli and a reduction in the number of maxillary alveoli to 12. The new species is the geologically youngest known member of the genus. While species of Baru are a ubiquitous component of Oligo-Miocene crocodylian assemblages from northern Australia, they are absent from all known Plio-Pleistocene sites, suggesting the extinction of the genus by the latest Miocene. We suggest that the marked taxonomic turnover of crocodylian assemblages between the Upper Miocene Alcoota Local Fauna and the Lower Pliocene is the result of a short period of severe aridity during the latest Miocene, causing widespread crocodylian extinctions across Australia, followed by the establishment of taxonomically novel crocodylian faunas in the Pliocene. This event mirrors similar contemporary losses of crocodylian diversity in Africa and South America, although these cases are partly driven by local tectonic events. It is likely that late Cenozoic global cooling also played a role in these extinctions.

baru crocodile mekosuchinae miocene australia extinction


Evolutionary process of extremely twisted heteromorph ammonites from the Upper Cretaceous of Japan

Abstract: Abstract The heteromorph ammonite Pravitoceras sigmoidale with extremely twisted retroversal hook and its probable ancestors, Didymoceras awajiense and D. morozumii, occur mainly in the Upper Cretaceous Izumi and Sotoizumi groups in southwest Japan. Their morphological characteristics were investigated in this study. Morphotypes I–II were recognized in D. morozumii. Morphotypes III–V were recognized in D. awajiense. Morphotypes VI–VIII were recognized in P. sigmoidale. Moreover, all morphotypes were recognized in both the dextral and sinistral forms of these three species. Based on the morphological and stratigraphic relationships, it is considered that the evolutionary process of these species is divided into three steps: (1) shift from morphotypes I–II to III; (2) increase in variation from morphotypes III to III–V; and (3) shift from morphotypes III–V to VI–VIII. By comparing these morphotypes based on the ‘growing tube model’ belonging to the moving frame analysis, it is seen that there are no significant gaps of coiling patterns in the evolutionary processes of this lineage. Meanwhile, morphotypes III–V of D. awajiense have a larger morphological variation than morphotypes I–II of D. morozumii and morphotypes VI–VIII of P. sigmoidale. These results, and the radiometric ages from previous studies, suggest that D. awajiense, with considerable variation between ancestor and descendant species, existed in a short interval. It seems that P. sigmoidale occurred after the continuous but rapid and drastic morphological evolution from D. morozumii via D. awajiense.

ammonite cretaceous didymoceras heteromorph nostoceratidae pravitoceras


Newly discovered morphology of the Silurian sea spider Haliestes and its implications

Abstract: Abstract The three-dimensionally preserved Haliestes dasos from the Silurian (Wenlock) Lagerstätte is the most complete fossil sea spider and the oldest unambiguous pycnogonid known from the fossil record. The discovery of two new specimens to add to the holotype reveals new features including proximal annulations of the appendages and segmentation of the trunk end, critical details for comparison with pycnogonids from the Devonian (Emsian) Hunsrück Slate and for the interpretation of the evolutionary significance of Palaeozoic genera. There is some evidence of sexual dimorphism. Haliestes dasos was nektobenthic and its morphology indicates an unusual mode of feeding compared with living pycnogonids. The new morphological features of H. dasos are closely similar to those in Palaeoisopus problematicus from the Hunsrück Slate and it clearly belongs, together with that species, in stem Pycnogonida and not the crown group.

herefordshire lagerstätte
silurian
arthropoda
pycnogonida
pantopoda
haliestes


The Late Miocene mammals from the Humahuaca Basin (northwestern Argentina) provide new evidence on the initial stages of the Great American Biotic Interchange

Abstract: Abstract We describe new Late Miocene mammalian specimens from the Maimará Formation (Late Miocene to Early Pliocene) exposed at Humahuaca Basin (23°–24°S), northwestern Argentina (NWA), and analyse their taxonomy and relevance for our understanding of the initial stages of the Great American Biotic Interchange (GABI). The stratigraphical and geochronological control of the studied specimens indicates a time window of c. 6.6–5.8 Ma. These data are crucial for establishing the oldest records of the Holarctic immigrants of the GABI. The first record of cricetid rodents from the Maimará Formation is reported. Moreover, with an age of c. 6 Ma it is the first appearance datum (FAD) of these rodents in South America. The age of the procyonid Cyonasua recorded in this unit is estimated between c. 6.6 and c. 6.4 Ma. The record of procyonids and cricetids in the same continuous sedimentary sequence suggests that the time interval between the dispersion of both groups into the continent during GABI was c. 1 myr. Of the autochthonous mammals from the Maimará Formation, the first records of litopterns, chlamyphorid and mylodontine xenarthrans, and caviomorph rodents (including new species Pithanotomys? solisae and Palaeocavia humahuaquense) are described. FADs of immigrant and autochthonous mammals in NWA suggest a major faunal turnover during GABI at the Messinian, probably linked to global climatic changes and the tectonic activity that affected the Humahuaca Basin at that time. Hypsodonty, a dominant feature among Maimaran mammals, may be related to a landscape strongly influenced by the Andean uplift.

late miocene
humahuaca basin
great american biotic interchange
cricetid
procyonid
faunal turnover


Evaluating growth in Macrospondylus bollensis (Crocodylomorpha, Teleosauroidea) in the Toarcian Posidonia Shale, Germany

Abstract: Abstract The study of how organisms grow is a fundamental aspect of palaeontology. Growth in teleosauroids is poorly understood and little studied, especially in an ontogenetic sense. We investigate growth rates of the most common and abundant teleosauroid, Macrospondylus bollensis, in which a large sample of multiple body sizes is available from the Posidonienschiefer Formation (Posidonia Shale) of southwestern Germany. We perform linear regression analyses on 62 specimens of Macrospondylus (16 juveniles, 7 subadults and 39 adults) using 21 cranial and postcranial measurements. Our results show that juvenile, subadult and adult individuals have near-isometric or isometric growth throughout much of the body. Notably, we find that in Macrospondylus: (1) the femur grows at a faster rate than the skull and hindlimb zeugopodium; (2) the forelimb and hindlimb grow at the same rate; and (3) there is distinct ontogenetic signal in the growth of the orbit and supratemporal fenestra. We also find that limb scaling in Macrospondylus is somewhat comparable to that seen in the extant gavialids Gavialis gangeticus and Tomistoma schlegelii. Last, we examine evolutionary allometry in skull length relative to body size in Crocodylomorpha using femoral length as a proxy, which shows a near-isometric trend. Non-thalattosuchian crocodylomorphs (with the exception of one pholidosaurid and one dryosaurid taxon) are differentiated from thalattosuchians due to their shorter skulls, as previously suggested, but the scaling relationship with femur length remains unchanged.

teleosauroidea
crocodylomorpha
evolutionary allometry
ontogenetic allometry
macrospondylus bollensis
posidonia shale formation


Elucidating the morphology and ecology of Eoandromeda octobrachiata from the Ediacaran of South Australia

Abstract: Abstract Eoandromeda octobrachiata is a poorly understood Ediacaran organism, with spiral octoradial arms, found in South Australia and South China. The informal Nilpena member of the Rawnsley Quartzite, Flinders Ranges in South Australia preserves more than 200 specimens of Eoandromeda. Here we use the novel application of rotational geometric morphometrics together with palaeoenvironmental information to provide a better insight into their palaeobiology and ecology, and to address conflicting hypotheses regarding mode of life and taxonomic affinity. We find that Eoandromeda probably had a radially symmetrical shape in life, was cone shaped and had a high relief off the microbial mat. Analysis of the symmetric and asymmetric shape components showed that they deform strongly in the direction of palaeocurrent, therefore they are thought to be made of a flexible material. Almost all specimens are compressed flat. Specimens that appear to have not fully collapsed support the idea that Eoandromeda was probably cone shaped and suggest that they possibly collapsed spirally. Our shape analysis, along with observed morphological features, support a benthic rather than pelagic mode of life. Morphological and ecological inconsistencies do not fully support the hypothesis of a Ctenophora taxonomic affinity.

ediacara
eoandromeda
south australia
morphometrics
benthic


Katian (Late Ordovician) trilobites of the North Qilian Mountains and their palaeogeographical implications for the Proto-Tethys Archipelagic Ocean

Abstract: Abstract Trilobites from the middle Koumenzi Formation (Katian, Upper Ordovician) of the North Qilian Mountains, Menyuan, northeastern Qinghai Province are systematically documented for the first time. The fauna consists of five families, seven genera and seven species, one of which is new (Remopleurides zhangi), showing a close relationship to those of the Kazakh terranes (such as the Chu-Ili terrane, Chingiz-Tarbagatai area and the Karatau-Naryn and North-Tien Shan microcontinents), and the North China and Laurentia palaeoplates during the Katian (Late Ordovician). The cluster and non-metric multidimensional scaling analyses of the Middle–Late Ordovician (late Darriwilian–Katian) trilobite faunas with 299 genera or subgenera from 46 horizons of 37 areas, provide valuable information for the palaeogeographical reconstruction of the Proto-Tethys Archipelagic Ocean (PTAO) of this interval. The Qilian terrane and adjacent areas are essential components of the PTAO, some of which include the Qilian terrane, the North Qilian Mountains area, the Altun faulted terrane, the Hexi Corridor area and the East Qinling terrane. Their relative positions in the PTAO are inferred by the palaeobiogeography of trilobite faunas. Furthermore, based on discussions of the spatiotemporal distribution of those faunas, the Pliomerina and/or Sinocybele Province of the Middle–Late Ordovician (late Darriwilian–Katian) age is defined as a trilobite faunal province of the PTAO. Moreover, a distinct faunal subprovince, essentially comprising the South China Palaeoplate and its neighbours (e.g. Tarim, Annamia, Sibuma, East Qinling, Turkestan-Alai and probably Talesh), might be surrounded by the equatorial cold-water tongue.

late ordovician
trilobite
north qilian mountains
proto-tethys archipelagic ocean (ptao)
palaeogeographical reconstruction
faunal province


Three-dimensional reconstruction, taphonomic and petrological data suggest that the oldest record of bioturbation is a body fossil coquina

Abstract: Abstract Fossil material assigned to Nenoxites from the late Ediacaran Khatyspyt Formation of Arctic Siberia (550–544 Ma) has been presented as evidence for bioturbation prior to the basal Cambrian boundary. However, that ichnological interpretation has been challenged, and descriptions of similar material from other global localities support a body fossil origin. Here we combine x-ray computed tomography, scanning electron microscopy and petrographic methods to evaluate the body or trace fossil nature of Nenoxites from the Khatyspyt Formation. The fossilized structures consist of densely packed chains of three-dimensionally preserved silicic, bowl-shaped elements surrounded by distinct sedimentary halos, in a dolomitized matrix. Individual bowl-shaped elements can exhibit diffuse mineralogical boundaries and bridging connections between elements, both considered here to result from silicification and dolomitization during diagenesis. This new morphological and petrological evidence, in conjunction with recent studies of the late Ediacaran tubular taxa Ordinilunulatus and Shaanxilithes from China, suggest that the Khatyspyt specimens most probably reflect a coquina deposit of Shaanxilithes-like body fossils. Our data support the possibility of Shaanxilithes-like organisms representing total group eumetazoans.

ediacaran sedimentology taphonomy nenoxites bioturbation petrology


Two new species of small-bodied pachycephalosaurine (Dinosauria, Marginocephalia) from the uppermost Cretaceous of North America suggest hidden diversity in well-sampled formations

Abstract: Abstract Here we report two new small-bodied pachycephalosaurines: one from the Dinosaur Park Formation of Alberta and the other from the Hell Creek Formation of Montana, each represented by an isolated squamosal. These two new specimens are approximately the same size as squamosals of Sphaerotholus buchholtzae, and possess several overlapping morphologies with the genus (such as a strongly posteroventrally projecting and laterally tapering parietosquamosal bar). Therefore, these two new specimens are identified as belonging to the genus Sphaerotholus. However, a suite of characters and combinations differentiate these two new specimens from the other three Sphaerotholus species. Most notably, they possess multiple posterior and lateral node rows on the squamosal, and lack a parietosquamosal node (a single row and a parietosquamosal node define these other species in the genus). Specifically, the Alberta specimen possesses two posterior and two lateral node rows, whereas the Montana specimen has three of each. This analysis provides phylogenetic support for the evolutionary hypothesis that a single-rowed and parietosquamosal node-less taxon underwent a cladogenetic event that gave rise to a single-rowed and parietosquamosal node-bearing lineage and a multi-rowed, parietosquamosal node-less lineage (the two taxa described herein). In turn, these two Sphaerotholus lineages possibly underwent anagenetic change throughout the latest Maastrichtian. Additionally, these new taxa suggest that while large-bodied taxa from well sampled formations are to be found with decreasing frequency, the inverse holds true for small-bodied taxa and that pachycephalosaurines were more diverse than previously recognized, and continued to diversify up until the very end of the Cretaceous.

pachycephalosauria
sphaerotholus
anagenesis
cladogenesis
dinosaur park formation
hell creek formation


The early Paleocene (Danian) climate of Svalbard based on palaeobotanical data

Abstract: Abstract The early Paleocene (Danian) Barentsburg flora from the Firkanten Formation, Svalbard, provides a valuable insight into the environment and climate of the Arctic early in the warm Palaeogene prior to the Paleocene–Eocene Thermal Maximum (PETM) event. The flora includes c. 50 species of ferns, ginkgos, conifers and angiosperms. Angiosperms predominate and are represented by the families Platanaceae, Cercidiphyllaceae, Trochodendraceae, Hamamelidaceae, Fagaceae, Betulaceae, Nyssaceae and several taxa of uncertain taxonomic position. Palaeoclimate parameters, calculated using a new calibration of the Climate Leaf Analysis Multivariate Program (CLAMP), indicate that Svalbard in the early Paleocene was characterized by a temperate, maritime, humid, seasonal climate with cool mild winters and warm summers. Humidity was highest during winter months. Short-term winter snow cover and light frosts were likely to have been frequent. The CLAMP analysis yields a mean annual temperature of 10.1 ± 1.98°С, a warmest month mean temperature of 19.2 ± 2.49°C and a coldest month mean temperature of 1.7 ± 3.24°С. The growing season lasted c. 6 months. Precipitation during this growth period was 572.4 ± 296.50 mm, while vapour pressure deficit and potential evapotranspiration indicate very high humidity year-round, with summer being only slightly drier than winter. Taken together with data from other early Paleocene circum-Arctic sites, these results point to highly uniform, temperate and extremely humid conditions around the margins of the early Cenozoic Arctic Ocean. This condition represents the warm background condition that dominated in the Arctic prior to the anomalous extreme warmth of the early Eocene.

climate leaf analysis multivariate program (clamp)


A taxonomic revision and cranial description of Terrestrisuchus gracilis (Archosauria, Crocodylomorpha) from the Upper Triassic of Pant-y-Ffynnon Quarry (southern Wales)

Abstract: Abstract Non-crocodyliform crocodylomorphs, formerly referred to the informal group ‘Sphenosuchia’, are the earliest known crocodylomorph precursors of extant crocodylians. They are therefore crucial for our understanding of early crocodylian evolution and the origin of typical crocodylian characteristics, such as the formation of a secondary palate, complex cranial pneumaticity, and a reinforced braincase. Terrestrisuchus gracilis, known from the Upper Triassic fissure fill deposits of Pant-y-Ffynnon in southern Wales, is represented by almost 200 specimens, comprising articulated, partially associated, and isolated remains. In this contribution we provide a taxonomic revision of Terrestrisuchus gracilis and redescribe its cranial anatomy in detail, based in part on novel micro-computed tomography data. The posterior skull region is extensively pneumatized as indicated, among other features, by large pre- and postcarotid recesses on the parabasisphenoid, and a large pneumatic cavity in the articular of the mandible. In contrast, the quadrate forms only a small, unfused contact with the prootic, suggesting that complex pneumatization of the postorbital region pre-dated the co-ossification of the quadrate and braincase in Crocodylomorpha. Terrestrisuchus gracilis preserves an ossified basihyal, which represents the first occurrence of this bone in non-avemetatarsalian archosaurs. Finally, we show that Terrestrisuchus gracilis was probably cathemeral (i.e. active in a range of light levels), based on a phylogenetic flexible discriminant analysis of the relative dimensions of the sclerotic ring and orbit.

terrestrisuchus
crocodylomorpha
triassic
ct scanning
cranial osteology
braincase pneumaticity


An enigmatic structure in the tail of vetulicolians from the Cambrian Chengjiang biota, South China

Abstract: Abstract Cambrian vetulicolians have mosaic characteristics of both deuterostomes and protostomes, which has important implications for the origin and early evolution of the Deuterostomia. They are intriguing in their bizarre body plan with a series of pharyngeal gill slits. The anterior section is characterized by five pairs of gill pouches, while the paddle-like posterior part is composed of seven or more segments. Although the overall external morphology and some internal anatomies of vetulicolians, such as pharyngeal cavity, gill system and alimentary canal, have largely been clarified, mysteries remain regarding some other internal structures, and their functional interpretations are highly controversial. In this study we identify an important but hitherto unrecognized feature: a sub-rounded, wrinkled structure located ventrally at the posterior section (or ‘tail’) of weakly sclerotized members of vetulicolians, including Xidazoon stephanus and Didazoon haoae, from the early Cambrian Chengjiang biota. The enigmatic structure consistently appears at or between the third and fourth segments of the posterior section. We show that the new structure is an internal organ in the body cavity of vetulicolians and infer that it might have functioned for reproduction, excretion or digestion. The finding of this enigmatic structure from X. stephanus and D. haoae enriches our understanding of vetulicolians and might facilitate further exploration of the anatomy and physiology of early deuterostomes.

vetulicolian
enigmatic structure
deuterostome
early cambrian
chengjiang biota


A unique Late Cretaceous fossil wood assemblage from Chilean Patagonia provides clues to a high-latitude continental environment

Abstract: Abstract Fossil plants, including large trunks, stems, some branches, and twigs, were collected from the Maastrichtian (68.9 Ma), upper Dorotea Formation in the Magallanes–Austral Basin, 16 km north of the Cerro Guido–Las Chinas complex in the southern Chilean Magallanes region. These fossil trunks range from 0.2 to 2.2 m in length. Petrographic slides were made in three sections (transverse, radial and tangential) and analysed under a light microscope to study the permineralized fossils. The woods and stems belong to Austroginkgoxylon gen. et sp. nov., Agathoxylon antarcticum, Podocarpoxylon paradoxi sp. nov., Podocarpoxylon mazzonii, Palmoxylon subantarcticae and Notomalvaceoxylon magallanense gen. et sp. nov. The growth rings of gymnosperms and anatomical characters of angiosperms were analysed to obtain palaeoecological data. Interactions between gymnosperm roots growing into the secondary xylem of an angiosperm (nurse logs) are recorded. The data obtained from the fossil woods suggest warm and humid conditions in this southern South American locality during the Late Cretaceous, providing a unique opportunity to study continental environments at high southern latitudes, which are poorly represented on a global scale.

gymnosperm
angiosperm
growth ring
wood anatomy
cretaceous
patagonia


Evidence for the evolutionary history and diversity of fossil sweetgums: leaves and associated capitate reproductive structures of Liquidambar from the Eocene of Hainan Island, South China

Abstract: Abstract Extant species of Liquidambar L. exhibit a disjunct distribution between western and eastern Asia and eastern North America to Central America, with the highest species diversity being in China. In this study, two new species of Liquidambar are described from the Eocene of Hainan Island, South China: L. hainanensis on basis of the leaf fossils, and L. ovoidea based on compressions of infructescences. The co-occurrences of L. hainanensis leaves, associated infructescences of L. ovoidea and dispersed pantoporate pollen grains have the combination of morphological features peculiar to the genera Liquidambar and former Semiliquidambar and Altingia. This is considered further palaeobotanical justification for the placement of these three genera into a single genus, Liquidambar, proposed earlier based on molecular markers. Our data provide evidence for the occurrence of an ancestral polymorphic group of Liquidambar species (L. maomingensis with associated capitate infructescences, L. bella, L. hainanensis and L. ovoidea) during the Eocene in South China, and corroborate the view that South China could have been a centre of Liquidambar speciation during the Eocene. The morphological similarity of L. hainanensis leaves to those of the extant species L. orientalis from Western Asia and the North American L. styraciflua may support the importance of both the North Atlantic Land Bridge and the Bering Land Bridge for the dispersal of Liquidambar. Analysis of fossil species of Liquidambar from eastern Asia also contributes to further understanding of patterns in the taxonomic diversity and evolutionary history of this genus.

liquidambar
eocene
leaf fossil
infructescence
pantoporate pollen grain
hainan island


First Eocene–Miocene anuran fossils from Peruvian Amazonia: insights into neotropical frog evolution and diversity

Abstract: Abstract Anurans are one of the most diverse vertebrate groups, particularly in Amazonia, where species richness exceeds that of anywhere else. Amazonian frogs belong to three main lineages (Hyloidea, Microhylidae and Pipidae), each of which diversified during the Cenozoic. However, due to the virtual absence of an anuran fossil record in that area, the evolutionary history of modern lineages has so far remained accessible only via molecular data. In recent decades, a series of field campaigns in Peruvian Amazonia led to the discovery of an unparalleled set of anuran bone fragments, scattered across different sites spanning the Eocene–Miocene time interval. Here, we describe these first Palaeogene and early Neogene anurans from Peru with a focus on humeral and ilial morphology, identifying five humeral and five ilial morphotypes. Humeral morphotypes suggest the presence of different lineages of Brachycephaloidea in Peruvian fossil assemblages, whereas ilial morphotypes suggest the presence of Leptodactylidae, although leptodactylid-like ilia also occur in some extant brachycephaloids. Pipids were also identified based on both humeral and ilial fragments. This study fills a major temporal and geographical gap in the evolutionary history of South American anurans, while further uncovering a lack of knowledge in the skeletal morphology of extant anuran families, as well as their inter- and intra-species variability.

anura amazonia brachycephaloidea eocene–miocene fossil pipidae


Redefining the Huayquerian Stage (Upper Miocene to Lower Pliocene) of the South American chronostratigraphic scale based on biostratigraphical analyses and geochronological dating

Abstract: Abstract The Huayquerian Stage of the South American chronostratigraphic scheme (named for the Huayquerías del Este, Argentina) was originally based on a poorly known mammal association of six taxa from the Huayquerías Formation. We studied the geology, age and fauna of the Neogene sequence in this area, including the Huayquerías, Tunuyán and Bajada Grande formations. The sequence comprises a monotonous succession of synorogenic epiclastic sediments deposited under arid to semi-arid conditions. Zircon U–Pb dates from 10 tuffaceous levels (7.2–1.6 Ma) place deposition of the Huayquerías Formation during the late Tortonian or Messinian to early Zanclean, the Tunuyán Formation during the Zanclean–Piacenzian, and the Bajada Grande Formation during the Piacenzian–Calabrian. We present 43 and 31 new mammal taxon records for the Huayquerías and Tunuyán formations, respectively. Progressive faunal change was observed along the sequence. The first records of the Chaco tortoise Chelonoidis chilensis and the notoungulate Xotodon major, and the latest records of Interatheriidae and Typotheriopsis (notoungulates), Metacaremys calfucalel, Phtoramys hidalguense and Lagostomus pretrichodactyla (rodents), Chasicotatus ameghinoi and Macroeuphractus morenoi (xenarthrans) are reported. The faunal associations of the Huayquerías and lower Tunuyán formations are highly similar to each other, and to other coeval localities in Argentina. The Macroeuphractus morenoi Assemblage Biozone is proposed as the basis for redefining the Huayquerian Stage, due to the co-occurrence of three taxa with wide geographical distribution in southern South America: Macroeuphractus morenoi, Pseudotypotherium subinsigne and Lagostomus pretrichodactyla. The age of this biozone is constrained at c. 8–5 Ma in its type area.

south american chronostratigraphic scale


The first European woolly rhinoceros mitogenomes, retrieved from cave hyena coprolites, suggest long-term phylogeographic differentiation

Abstract: The woolly rhinoceros (Coelodonta antiquitatis) is an iconic species of the Eurasian Pleistocene megafauna, which was abundant in Eurasia in the Pleistocene until its demise beginning approximately 10 000 years ago. Despite the early recovery of several specimens from well-known European archaeological sites, including its type specimen (Blumenbach 1799), no genomes of European populations were available so far, and all available genomic data originated exclusively from Siberian populations. Using coprolites of cave hyenas (Crocuta crocuta spelea) recovered from Middle Palaeolithic layers of two caves in Germany (Bockstein-Loch and Hohlenstein-Stadel), we isolated and enriched predator and prey DNA to assemble the first European woolly rhinoceros mitogenomes, in addition to cave hyena mitogenomes. Both coprolite samples produced copious sequences assigned to C. crocuta (27% and 59% mitogenome coverage, respectively) and woolly rhinoceros (Coelodonta antiquitatis; 27% and 81% coverage, respectively). The sequences suggested considerable DNA degradation, which may limit the conclusions to be drawn; however, the mitogenomes of European woolly rhinoceros are genetically distinct from the Siberian woolly rhinoceros, and analyses of the more complete mitogenome suggest a split of the populations potentially coinciding with the earliest fossil records of woolly rhinoceros in Europe.

ancient dna
predator
prey
hyaenidae
rhinocerotidae
mammal


<i>Gilsonicaris</i> from the Lower Devonian Hunsrück slate is a eunicidan annelid and not the oldest crown anostracan crustacean

Abstract: The Lower Devonian (Lower Emsian, −400 Myr) roof slates of the Hunsrück in southeastern Germany have delivered a highly diverse and exceptionally preserved marine fauna that provides a unique snapshot into the anatomy and ecology of a wide range of Palaeozoic animals. Several of the described taxa, however, remain enigmatic in their affinity, at least until new pyritized features hidden under the surface of the slate are revealed using X-ray radiography or micro-computed tomography (µCT). Here, we redescribe such an enigmatic fossil, the putative anostracan crustacean Gilsonicaris rhenanus Van Straelen, 1943. Using µCT scanning, we unveil unprecedented details of its anatomy, including a ventral oral opening and four pairs of recalcitrant jaw elements. These jaws are morphologically consistent with the scolecodonts of eunicidan polychaetes, which along with the gross anatomy of the body and head unambiguously identifies G. rhenanus as a polychaete rather than an arthropod. While this discovery firmly discards the Early Devonian record of crown anostracans in the fossil record, it adds a new record of eunicidan soft tissues, which are surprisingly rare considering the abundant microfossil record of scolecodonts.

branchiopoda
polychaeta
scolecodonts
soft-tissue
x-ray microtomography
early devonian


Palaeoecological deductions from osteohistology

Abstract: Palaeoecological deductions are vital for understanding the evolution and diversification of species within prehistoric environments. This review highlights the multitude of ways in which the microanatomy and microscopic structure of bones enables palaeoecological deductions. The occurrence of growth marks in bones is discussed, and their usefulness in deducing the ontogenetic status and age of individuals is considered, as well as how such marks in bones permit the assessment of the growth dynamics of individuals and species. Here osteohistology is shown to provide insight into the structure of past populations, as well as ecological relationships between individuals. In addition, the response of bones to trauma, disease and moulting is considered. Finally, I explore how osteohistology can give insight into ecomorphological adaptations, such as filter feeding, probe feeding and saltatorial locomotion. Methodological advances in three-dimensional microtomography and synchrotron scanning bodes well for future studies in osteohistology and despite some compromises in terms of tissue identity, circumvents the crucial issue of destructive analyses.

palaeohistology
bone histology
microanatomy
palaeoecology


A genetic glimpse of the Chinese straight-tusked elephants

Abstract: Straight-tusked elephants (genus: Palaeoloxodon) including their island dwarf forms are extinct enigmatic members of the Pleistocene megafauna and the most common Pleistocene elephants after the mammoths. Their taxonomic placement has been revised several times. Using palaeogenomic evidence, previous studies suggested that the European P. antiquus has a hybrid origin, but no molecular data have been retrieved from their Asian counterparts, leaving a gap in our knowledge of the global phylogeography and population dynamics of Palaeoloxodon. Here, we captured a high-quality complete mitogenome from a Pleistocene Elephantidae molar (CADG841) from Northern China, which was previously morphologically assigned to the genus Elephas (Asian elephant), and partial mitochondrial sequences (838 bp) of another Palaeoloxodon sp. specimen (CADG1074) from Northeastern China. We found that both Chinese specimens cluster with a 244 000-year-old P. antiquus (specimen name: WE) from Western Europe, suggesting that this clade may represent a population with a large spatial span across Eurasia. Based on the fossil record and the molecular dating of both the divergences of different Palaeoloxodon mitochondrial clades and previously determined hybridization events, we propose that this Eurasian-wide WE clade provides evidence for an earlier migration and/or another hybridization event that happened in the evolutionary history of straight-tusked elephants.

ancient dna
mitogenome
palaeoloxodon
asia
phylogenetic tree


Bridging two oceans: small toothed cetaceans (Odontoceti) from the Late Miocene Chagres Formation, eastern Caribbean (Colon, Panama)

Abstract: Fossil cetaceans are often found in Miocene marine outcrops across the globe. However, because this record is not homogeneous, the dissimilar increase in occurrences, along with the sampling bias has created regions with extensive records and others with great scarcity. Among these, the Caribbean has remained enigmatic due to the lack of well-preserved cetacean fossils. Here, we report new Caribbean fossil cetaceans from the Upper Miocene Chagres Formation exposed along Piña beach, Eastern Panama, including a scaphokogiine kogiid, an Acrophyseter-like physeteroid and the phocoenid Piscolithax. Along with previous records of the iniid Isthminia panamensis and the kogiine Nanokogia isthmia, the Chagres cetacean fauna shows some similarities with other Late Miocene cetacean communities such as the Californias in the North Pacific, although their closest affinities lie with the eastern South Pacific Pisco Formation, Peru. Such findings indicate that though deep and intermediate Caribbean–Pacific water interchange was reduced by the Middle Miocene due to the shallowing of the Central American Seaway, shallow waters marine connection that persisted until the Pliocene might have facilitated the dispersal of coastal species across both sides of the Isthmus.

panama
miocene
odontocetes
pisco formation
chagres formation


Cambrian origin but no early burst in functional disparity for Class Bivalvia

Abstract: Both the Cambrian explosion, more than half a billion years ago, and its Ordovician aftermath some 35 Myr later, are often framed as episodes of widespread ecological opportunity, but not all clades originating during this interval showed prolific rises in morphological or functional disparity. In a direct analysis of functional disparity, instead of the more commonly used proxy of morphological disparity, we find that ecological functions of Class Bivalvia arose concordantly with and even lagged behind taxonomic diversification, rather than the early-burst pattern expected for clades originating in supposedly open ecological landscapes. Unlike several other clades originating in the Cambrian explosion, the bivalves' belated acquisition of key anatomical novelties imposed a macroevolutionary lag, and even when those novelties evolved in the Early Ordovician, functional disparity never surpassed taxonomic diversity. Beyond this early period of animal evolution, the founding and subsequent diversification of new major clades and their functions might be expected to follow the pattern of the early bivalves—one where interactions between highly dynamic environmental and biotic landscapes and evolutionary contingencies need not promote prolific functional innovation.

bivalve
early burst
cambrian explosion
functional disparity
diversity-disparity
adaptive radiation


Anthropogenic predictors of varying Holocene occurrence for Europe's large mammal fauna

Abstract: Understanding how species respond to different anthropogenic pressures is essential for conservation planning. The archaeological record has great potential to inform extinction risk assessment by providing evidence on past human-caused biodiversity loss, but identifying specific drivers of past declines from environmental archives has proved challenging. We used 17 684 Holocene zooarchaeological records for 15 European large mammal species together with data on past environmental conditions and anthropogenic activities across Europe, to assess the ability of environmental archives to determine the relative importance of different human pressures in shaping faunal distributions through time. Site occupancy probability showed differing significant relationships with environmental covariates for all species, and nine species also showed significant relationships with anthropogenic covariates (human population density, % cropland, % grazing land). Across-species differences in negative relationships with covariates provide ecological insights for understanding extinction dynamics: some mammals (red deer, aurochs, wolf, wildcat, lynx, pine marten and beech marten) were more vulnerable to past human–environmental interactions, and differing single and synergistic anthropogenic factors influenced likelihood of past occurrence across species. Our results provide new evidence for pre-industrial population fragmentation and depletion in European mammals, and demonstrate the usefulness of historical baselines for understanding species' varying long-term sensitivity to multiple threats.

environmental archives
extinction filter
extinction risk
historical baselines
quaternary extinction
zooarchaeology


The earliest pollen-loaded insects from the Lower Permian of Russia

Abstract: Recent fossil discoveries suggest that the coevolution of insect pollinators and gymnosperms started long before the appearance of flowering plants. One of the keys to understanding the origins of pollination relationships is fossil insects with gymnosperm pollen attached to the body surface. Such fossils are exceedingly rare to find, especially from the Palaeozoic, a time when ambers with insect inclusions were absent. Here, we report compression fossils of Early Permian tillyardembiid insects (Polyneoptera) preserved with pollen on their heads, thoraces, legs and abdomens. This is the earliest finding of pollen-bearing insects, predating the previous oldest record from the Middle Jurassic by ca 120 Ma. Judging by the pollen composition, tillyardembiids visited a narrow range of host plants, including Rufloriaceae (Cordaitales). While it is impossible to say for certain whether tillyardembiids as pollen consumers contributed to pollination, a trophic specialization of this kind could be considered an evolutionary precursor of pollination mutualism.

plant–insect interactions
pollination
palaeozoic


Beetle elytra: evolution, modifications and biological functions

Abstract: Conversion of forewings into hardened covers, elytra, was a ground-breaking morphological adaptation that has contributed to the extraordinary evolutionary success of beetles. Nevertheless, the knowledge of the functional aspects of these structures is still fragmentary and scattered across a large number of studies. Here, we have synthesized the presently available information on the evolution, development, modifications and biological functions of this crucial evolutionary novelty. The formation of elytra took place in the earliest evolution of Coleoptera, very likely already in the Carboniferous, and was achieved through the gradual process of progressive forewing sclerotization and the formation of inward directed epipleura and a secluded sub-elytral space. In many lineages of modern beetles, the elytra have been distinctly modified. This includes multiple surface modifications, a rigid connection or fusion of the elytra, or partial or complete reduction. Beetle elytra can be involved in a very broad spectrum of functions: mechanical protection of hind wings and body, anti-predator strategies, thermoregulation and water saving, water harvesting, flight, hind wing folding, diving and swimming, self-cleaning and burrow cleaning, phoresy of symbiotic organisms, mating and courtship, and acoustic communication. We postulate that the potential of the elytra to take over multiple tasks has enormously contributed to the unparalleled diversification of beetles.

forewing evolution development coleoptera morphology biomaterial


Elbow-joint morphology in the North American ‘cheetah-like’ cat <i>Miracinonyx trumani</i>

Abstract: The North American cheetah-like cat Miracinonyx trumani is an extinct species that roamed the Pleistocene prairies 13 000 years ago. Although M. trumani is more closely related to the cougar (Puma concolor) than to the living cheetah (Acinonyx jubatus), it is believed that both A. jubatus and M. trumani possess a highly specialized skeleton for fast-running, including limbs adapted for speed at the expense of restricting the ability of prey grappling. However, forelimb dexterity of M. trumani has not been yet investigated. Here, we quantify the 3D-shape of the humerus distal epiphysis as a proxy for elbow-joint morphology in a sample of living cats to determine whether the extinct M. trumani was specialized to kill open-country prey using predatory behaviour based on fast running across the prairies and steppe terrains of the North American Pleistocene. We show that M. trumani had an elbow morphology intermediate to that of P. concolor and A. jubatus, suggesting that M. trumani had a less specialized pursuit predatory behaviour than A. jubatus. We propose that M. trumani probably deployed a unique predatory behaviour without modern analogues. Our results bring into question the degree of ecomorphological convergence between M. trumani and its Old World vicar A. jubatus.

miracinonyx trumani
predatory behaviour
elbow joint
convergence
pronghorn


The unbearable uncertainty of panarthropod relationships

Abstract: Panarthropoda, the clade comprising the phyla Onychophora, Tardigrada and Euarthropoda, encompasses the largest majority of animal biodiversity. The relationships among the phyla are contested and resolution is key to understanding the evolutionary assembly of panarthropod bodyplans. Molecular phylogenetic analyses generally support monophyly of Onychophora and Euarthropoda to the exclusion of Tardigrada (Lobopodia hypothesis), which is also supported by some analyses of morphological data. However, analyses of morphological data have also been interpreted to support monophyly of Tardigrada and Euarthropoda to the exclusion of Onychophora (Tactopoda hypothesis). Support has also been found for a clade of Onychophora and Tardigrada that excludes Euarthropoda (Protarthropoda hypothesis). Here we show, using a diversity of phylogenetic inference methods, that morphological datasets cannot discriminate statistically between the Lobopodia, Tactopoda and Protarthropoda hypotheses. Since the relationships among the living clades of panarthropod phyla cannot be discriminated based on morphological data, we call into question the accuracy of morphology-based phylogenies of Panarthropoda that include fossil species and the evolutionary hypotheses based upon them.

panarthropoda
tactopoda
lobopodia
protarthropoda
phylogeny
topology testing


Palaeopathological evidence for intraspecific combat in ankylosaurid dinosaurs

Abstract: Ankylosaurid dinosaurs were heavily armoured herbivores with tails modified into club-like weapons. These tail clubs have widely been considered defensive adaptations wielded against predatory theropod dinosaurs. Here we argue instead that ankylosaurid tail clubs were sexually selected structures used primarily for intraspecific combat. We found pathological osteoderms (armour plates) in the holotype specimen of Zuul crurivastator, which are localized to the flanks in the hip region rather than distributed randomly across the body, consistent with injuries inflicted by lateral tail-swinging and ritualized combat. We failed to find convincing evidence for predation as a key selective pressure in the evolution of the tail club. High variation in tail club size through time, and delayed ontogenetic growth of the tail club further support the sexual selection hypothesis. There is little doubt that the tail club could have been used in defence when needed, but our results suggest that sexual selection drove the evolution of this impressive weapon. This changes the prevailing view of ankylosaurs, suggesting they were behaviorally complex animals that likely engaged in ritualized combat for social dominance as in other ornithischian dinosaurs and mammals.

dinosauria
ankylosauria
sexual selection
animal weaponry


The origin of modern patterns of continental diversity in Mauritiinae palms: the Neotropical museum and the Afrotropical graveyard

Abstract: While the latitudinal diversity gradient has received much attention, biodiversity and species richness also vary between continents across similar latitudes. Fossil information can be used to understand the evolutionary mechanisms that generated such variation between continents of similar latitudes. We integrated fossil data into a phylogenetic analysis of the Mauritiinae palms, whose extant diversity is restricted to the Neotropics, but extended across Africa and India during most of the Cenozoic. Mauritiinae diverged from its sister lineage Raphiinae ca 106 Ma. Using ancestral state estimation and a lineage through time analysis, we found that diversity arose globally during the late Cretaceous and Palaeocene across South America, Africa and India. The Palaeocene–Eocene transition (ca 56 Ma) marked the end of global Mauritiinae expansion, and the beginning of their decline in both Africa and India. Mauritiinae disappeared from the Indian subcontinent and Africa at the end of the Eocene and the Miocene, respectively. By contrast, Neotropical diversity steadily increased over the last 80 Myr. Taken together, our results suggest that the Neotropics functioned as a continental-scale refuge for Mauritiinae palms, where lineages survived and diversified while global climatic changes that drastically reduced rainforests led to their demise on other continents.

diversification
evolution
longitudinal diversity gradient
palms
palynology


An early nimravid from California and the rise of hypercarnivorous mammals after the middle Eocene climatic optimum

Abstract: Carnivoraforms (crown carnivorans and their closest relatives) first occupied hypercarnivorous niches near the dawn of the late Eocene, 40–37 Ma. This followed the decline or extinction of earlier carnivorous groups, Mesonychia and Oxyaenodonta, leaving carnivoraforms and hyaenodontan meat-eaters as high trophic level consumers. The pattern of this change and the relative contributions of the taxonomic groups has hitherto been unclear. We report a new genus and species of the sabretoothed mammalian carnivore family Nimravidae, Pangurban egiae, from the Eocene Pomerado Conglomerate of southern California, with strongly derived hypercarnivorous features. While geochronologically the oldest named nimravid in North America, Pangurban egiae is recovered as phylogenetically derived, with affinities to Hoplophoneus. This provides unequivocal evidence for rapid radiation and spread of nimravid carnivores across Asia and North America and constrains the timing of early divergences within the family. Pangurban egiae narrows the gap between convergent iterations of sabretoothed mammalian carnivores and demonstrates swift diversification of the hypercarnivorous nimravids during a period of global climatic instability. Furthermore, it highlights the top-to-bottom restructuring North American ecosystems underwent during the Eocene–Oligocene transition, resulting in carnivoraforms taking positions as trophic specialists for the first time, a niche they still occupy today.

sabretooth
carnassial
hoplophoneus
duchesnean
evolution
eocene–oligocenebiotic transition


A possible home for a bizarre Carboniferous animal: is <i>Typhloesus</i> a pelagic gastropod?

Abstract: By contrast to many previously enigmatic Palaeozoic fossils, the Carboniferous metazoan Typhloesus has defied phylogenetic placement. Here, we document new features, including possible phosphatized muscle tissues and a hitherto unrecognized feeding apparatus with two sets of ca 20 spinose teeth whose closest similarities appear to lie with the molluscan radula. The ribbon-like structure, located well behind the mouth area and deep into the anterior part of the body, is interpreted as being in an inverted proboscis configuration. Gut contents, mostly conodonts, in the midgut area demonstrate that Typhloesus was an active predator. This animal was capable of propelling itself in the water column using its flexible body and a prominent posterior fin. The affinity of Typhloesus as a pelagic mollusc remains problematic but may lie more closely with the gastropods. Heteropod gastropods share with Typhloesus an active predatory lifestyle and have a comparable general body organization, albeit they possess characteristic aragonitic shells and their origins in the Jurassic post-date Typhloesus. Typhloesus may represent an independent radiation of Mid-Palaeozoic pelagic gastropods.


The emblematic South African therocephalian <i>Euchambersia</i> in China: a new link in the dispersal of late Permian vertebrates across Pangea

Abstract: Therapsids were widely distributed in Pangea in the late Permian. South Africa in Gondwana and Russia in Laurasia are the principal areas recording tetrapods (including therapsids) of this age. More recent field explorations have increased the importance of Chinese late Permian fossil assemblages. This is clearly reflected in the discovery of several new therocephalians from the Naobaogou Formation in Nei Mongol. Here, we report a therocephalian from that unit identified as a new species of the emblematic South African taxon Euchambersia. The new species, Euchambersia liuyudongi, is represented by a well-preserved skull and mandible showing a well-developed maxillary fossa and the absence of postcanine teeth. This is the third akidnognathid therocephalian recovered from the Naobaougou Formation, but oddly, the two basal Chinese akidnognathids previously known were recovered from a younger unit of the formation than the derived E. liuyudongi. This is the first time that the same therocephalian genus has been recorded in northern and southern continents, making the record of the Naobaougou Formation key to understanding the evolution of late Permian continental fauna in general, and of akidnognathid therocephalians in particular.

therocephalia
lopingian
naobaougou formation
china
south africa


Giant gar from directly above the Cretaceous–Palaeogene boundary suggests healthy freshwater ecosystems existed within thousands of years of the asteroid impact

Abstract: The Cretaceous–Palaeogene (K–Pg) mass extinction was responsible for the destruction of global ecosystems and loss of approximately three-quarters of species diversity 66 million years ago. Large-bodied land vertebrates suffered high extinction rates, whereas small-bodied vertebrates living in freshwater ecosystems were buffered from the worst effects. Here, we report a new species of large-bodied (1.4–1.5 m) gar based on a complete skeleton from the Williston Basin of North America. The new species was recovered 18 cm above the K–Pg boundary, making it one of the oldest articulated vertebrate fossils from the Cenozoic. The presence of this freshwater macropredator approximately 1.5–2.5 thousand years after the asteroid impact suggests the rapid recovery and reassembly of North American freshwater food webs and ecosystems after the mass extinction.

gar
k–pg
lilliput effect
lepisosteidae
fish


Genetic evidence for post-glacial expansion from a southern refugium in the eastern moa (<i>Emeus crassus</i>)

Abstract: Cycles of glacial expansion and contraction throughout the Pleistocene drove increases and decreases, respectively, in the geographical range and population size of many animal species. Genetic data have revealed that during glacial maxima the distribution of many Eurasian animals was restricted to small refugial areas, from which species expanded to reoccupy parts of their former range as the climate warmed. It has been suggested that the extinct eastern moa (Emeus crassus)—a large, flightless bird from New Zealand—behaved analogously during glacial maxima, possibly surviving only in a restricted area of lowland habitat in the southern South Island of New Zealand during the Last Glacial Maximum (LGM). However, previous studies have lacked the power and geographical sampling to explicitly test this hypothesis using genetic data. Here we analyse 46 ancient mitochondrial genomes from Late Pleistocene and Holocene bones of the eastern moa from across their post-LGM distribution. Our results are consistent with a post-LGM increase in the population size and genetic diversity of eastern moa. We also demonstrate that genetic diversity was higher in eastern moa from the southern extent of their range, supporting the hypothesis that they expanded from a single glacial refugium following the LGM.

ancient dna
megafauna
new zealand
phylogeography
quaternary


Evolution of amniote dentine apposition rates

Abstract: In amniotes, daily rates of dentine formation in non-ever-growing teeth range from less than 1 to over 25 μm per day. The latter value has been suggested to represent the upper limit of odontoblast activity in non-ever-growing teeth, a hypothesis supported by the lack of scaling between dentine apposition rates and body mass in Dinosauria. To determine the correlates and potential controls of dentine apposition rate, we assembled a dataset of apposition rates, metabolic rates and body masses for ca 80 amniote taxa of diverse ecologies and diets. We used phylogenetic regression to test for scaling relationships and reconstruct ancestral states of daily dentine apposition across Amniota. We find no relationship between body mass and daily dentine apposition rate (DDAR) for non-ever-growing teeth in Amniota as a whole or within major clades. Metabolic rate, the number of tooth generations, diet and habitat also do not predict or correspond with DDARs. Similar DDARs are found in large terrestrial mammals, dinosaurs and marine reptiles, whereas primates, cetaceans and some smaller marine reptiles independently evolved exceptionally slow rates. Life-history factors may explain the evolution of dentine apposition rates, which evolved rapidly at the origin of major clades.


Dietary isotopes of Madagascar's extinct megafauna reveal Holocene browsing and grazing guilds

Abstract: Megafauna play a disproportionate role in developing and maintaining their biomes, by regulating plant dispersal, community structure and nutrient cycling. Understanding the ecological roles of extinct megafaunal communities, for example through dietary reconstruction using isotope analysis, is necessary to determine pre-human states and set evidence-based restoration goals. We use δ13C and δ15N isotopic analyses to reconstruct Holocene feeding guilds in Madagascar's extinct megaherbivores, which included elephant birds, hippopotami and giant tortoises that occurred across multiple habitats and elevations. We compare isotopic data from seven taxa and two elephant bird eggshell morphotypes against contemporary regional floral baselines to infer dietary subsistence strategies. Most taxa show high consumption of C3 and/or CAM plants, providing evidence of widespread browsing ecology. However, Aepyornis hildebrandti, an elephant bird restricted to the central highlands region, has isotope values with much higher δ13C values than other taxa. This species is interpreted as having obtained up to 48% of its diet from C4 grasses. These findings provide new evidence for distinct browsing and grazing guilds in Madagascar's Holocene megaherbivore fauna, with implications for past regional distribution of ecosystems dominated by endemic C4 grasses.

elephant bird
hippopotamus
isotope ecology
megafauna
quaternary extinction


Body-axis organization in tetrapods: a model-system to disentangle the developmental origins of convergent evolution in deep time

Abstract: Convergent evolution is a central concept in evolutionary theory but the underlying mechanism has been largely debated since On the Origin of Species. Previous hypotheses predict that developmental constraints make some morphologies more likely to arise than others and natural selection discards those of the lowest fitness. However, the quantification of the role and strength of natural selection and developmental constraint in shaping convergent phenotypes on macroevolutionary timescales is challenging because the information regarding performance and development is not directly available. Accordingly, current knowledge of how embryonic development and natural selection drive phenotypic evolution in vertebrates has been extended from studies performed at short temporal scales. We propose here the organization of the tetrapod body-axis as a model system to investigate the developmental origins of convergent evolution over hundreds of millions of years. The quantification of the primary developmental mechanisms driving body-axis organization (i.e. somitogenesis, homeotic effects and differential growth) can be inferred from vertebral counts, and recent techniques of three-dimensional computational biomechanics have the necessary potential to reveal organismal performance even in fossil forms. The combination of both approaches offers a novel and robust methodological framework to test competing hypotheses on the functional and developmental drivers of phenotypic evolution and evolutionary convergence.

macroevolution
development
phenotypic evolution
tetrapod axis


The impact of molecular data on the phylogenetic position of the putative oldest crown crocodilian and the age of the clade

Abstract: The use of molecular data for living groups is vital for interpreting fossils, especially when morphology-only analyses retrieve problematic phylogenies for living forms. These topological discrepancies impact on the inferred phylogenetic position of many fossil taxa. In Crocodylia, morphology-based phylogenetic inferences differ fundamentally in placing Gavialis basal to all other living forms, whereas molecular data consistently unite it with crocodylids. The Cenomanian Portugalosuchus azenhae was recently described as the oldest crown crocodilian, with affinities to Gavialis, based on morphology-only analyses, thus representing a potentially important new molecular clock calibration. Here, we performed analyses incorporating DNA data into these morphological datasets, using scaffold and supermatrix (total evidence) approaches, in order to evaluate the position of basal crocodylians, including Portugalosuchus. Our analyses incorporating DNA data robustly recovered Portugalosuchus outside Crocodylia (as well as thoracosaurs, planocraniids and Borealosuchus spp.), questioning the status of Portugalosuchus as crown crocodilian and any future use as a node calibration in molecular clock studies. Finally, we discuss the impact of ambiguous fossil calibration and how, with the increasing size of phylogenomic datasets, the molecular scaffold might be an efficient (though imperfect) approximation of more rigorous but demanding supermatrix analyses.

crocodylia
molecular data
divergence-age
calibration
portugalosuchus


Ostracods had colonized estuaries by the late Silurian

Abstract: The fossil record of terrestrialization documents notable shifts in the environmental and physiological tolerances of many animal and plant groups. However, for certain significant components of modern freshwater and terrestrial environments, the transition out of marine settings remains largely unconstrained. Ostracod crustaceans occupy an exceptional range of modern aquatic environments and are invaluable palaeoenvironmental indicators in the fossil record. However, pre-Carboniferous records of supposed non-marine and marginal marine ostracods are sparse, and the timing of their marine to non-marine transition has proven elusive. Here, we reassess the early environmental history of ostracods in light of new assemblages from the late Silurian of Vietnam. Two, low diversity but distinct ostracod assemblages are associated with estuarine deposits. This occurrence is consistent with previous incidental reports of ostracods occupying marginal and brackish settings through the late Silurian and Devonian. Therefore, ostracods were pioneering the occupation of marginal marine and estuarine settings 60 Myr before the Carboniferous and they were a component of the early phase of transition from marine to non-marine environments.

silurian ostracods pioneer colonizers estuary


Growth variability, dimensional scaling, and the interpretation of osteohistological growth data

Abstract: Osteohistological data are commonly used to study the life history of extant and extinct tetrapods. While recent advances have permitted detailed reconstructions of growth patterns, physiology and other features using these data, they are most commonly used in assessments of ontogenetic stage and relative growth in extinct animals. These methods have seen widespread adoption in recent years, rapidly becoming a common component of the taxonomic description of new fossil taxa, but are often applied without close consideration of the sources of variation present or the dimensional scaling relationships that exist among different osteohistological measurements. Here, we use a combination of theoretical models and empirical data from a range of extant and extinct tetrapods to review sources of variability in common osteohistological measurements, their dimensional scaling relationships and the resulting interpretations that can be made from those data. In particular, we provide recommendations on the usage and interpretation of growth mark spacing/zonal thickness data, when these are likely to be unreliable, and under what conditions they can provide useful inferences for studies of growth and life history.

osteohistology ontogeny growth dinosaur mammal bird


Causal analysis of the temperature impact on deep-sea biodiversity

Abstract: The deep sea comprises more than 90% of the ocean; therefore, understanding the controlling factors of biodiversity in the deep sea is of great importance for predicting future changes in the functioning of the ocean system. Consensus has recently been increasing on two plausible factors that have often been discussed as the drivers of deep‐sea species richness in the contexts of the species‐energy and physiological tolerance hypotheses: (i) seafloor particulate organic carbon (POC) derived from primary production in the euphotic zone and (ii) temperature. Nonetheless, factors that drive deep-sea biodiversity are still actively debated potentially owing to a mirage of correlations (sign and magnitude are generally time dependent), which are often found in nonlinear, complex ecological systems, making the characterization of causalities difficult. Here, we tested the causal influences of POC flux and temperature on species richness using long-term palaeoecological datasets derived from sediment core samples and convergent cross mapping, a numerical method for characterizing causal relationships in complex systems. The results showed that temperature, but not POC flux, influenced species richness over 103–104-year time scales. The temperature–richness relationship in the deep sea suggests that human-induced future climate change may, under some conditions, affect deep-sea ecosystems through deep-water circulation changes rather than surface productivity changes.


Giant baleen whales emerged from a cold southern cradle

Abstract: Baleen whales (mysticetes) include the largest animals on the Earth. How they achieved such gigantic sizes remains debated, with previous research focusing primarily on when mysticetes became large, rather than where. Here, we describe an edentulous baleen whale fossil (21.12–16.39 mega annum (Ma)) from South Australia. With an estimated body length of 9 m, it is the largest mysticete from the Early Miocene. Analysing body size through time shows that ancient baleen whales from the Southern Hemisphere were larger than their northern counterparts. This pattern seemingly persists for much of the Cenozoic, even though southern specimens contribute only 19% to the global mysticete fossil record. Our findings contrast with previous ideas of a single abrupt shift towards larger size during the Plio-Pleistocene, which we here interpret as a glacially driven Northern Hemisphere phenomenon. Our results highlight the importance of incorporating Southern Hemisphere fossils into macroevolutionary patterns, especially in light of the high productivity of Southern Ocean environments.

mysticeti
chaeomysticeti
southernhemisphere
body size
gigantism


Convergent evolution of ventral adaptations for enrolment in trilobites and extant euarthropods

Abstract: The ability to enrol for protection is an effective defensive strategy that has convergently evolved multiple times in disparate animal groups ranging from euarthropods to mammals. Enrolment is a staple habit of trilobites, and their biomineralized dorsal exoskeleton offered a versatile substrate for the evolution of interlocking devices. However, it is unknown whether trilobites also featured ventral adaptations for enrolment. Here, we report ventral exoskeletal adaptations that facilitate enrolment in exceptionally preserved trilobites from the Middle Ordovician Walcott–Rust Quarry in New York State, USA. Walcott–Rust trilobites reveal the intricate three-dimensional organization of the non-biomineralized ventral anatomy preserved as calcite casts, including the spatial relationship between the articulated sternites (i.e. ventral exoskeletal plates) and the wedge-shaped protopodites. Enrolment in trilobites is achieved by ventrally dipping the anterior margin of the sternites during trunk flexure, facilitated by the presence of flexible membranes, and with the close coupling of the wedge-shaped protopodites. Comparisons with the ventral morphology of extant glomerid millipedes and terrestrial isopods reveal similar mechanisms used for enrolment. The wedge-shaped protopodites of trilobites closely resemble the gnathobasic coxa/protopodite of extant horseshoe crabs. We propose that the trilobites' wedge-shaped protopodite simultaneously facilitated tight enrolment and gnathobasic feeding with the trunk appendages.

walcott–rust
exceptional preservation
glomerida
trilobita
limulus
isopoda


Millennial processes of population decline, range contraction and near extinction of the European bison

Abstract: European bison (Bison bonasus) were widespread throughout Europe during the late Pleistocene. However, the contributions of environmental change and humans to their near extinction have never been resolved. Using process-explicit models, fossils and ancient DNA, we disentangle the combinations of threatening processes that drove population declines and regional extinctions of European bison through space and across time. We show that the population size of European bison declined abruptly at the termination of the Pleistocene in response to rapid environmental change, hunting by humans and their interaction. Human activities prevented populations of European bison from rebounding in the Holocene, despite improved environmental conditions. Hunting caused range loss in the north and east of its distribution, while land use change was responsible for losses in the west and south. Advances in hunting technologies from 1500 CE were needed to simulate low abundances observed in 1870 CE. While our findings show that humans were an important driver of the extinction of the European bison in the wild, vast areas of its range vanished during the Pleistocene–Holocene transition because of post-glacial environmental change. These areas of its former range have been climatically unsuitable for millennia and should not be considered in reintroduction efforts.

extinction dynamics
rewilding
process-basedmodel
megafauna
range dynamics
conservation biogeography


Uncovering the Holocene roots of contemporary disease-scapes: bringing archaeology into One Health

Abstract: The accelerating pace of emerging zoonotic diseases in the twenty-first century has motivated cross-disciplinary collaboration on One Health approaches, combining microbiology, veterinary and environmental sciences, and epidemiology for outbreak prevention and mitigation. Such outbreaks are often caused by spillovers attributed to human activities that encroach on wildlife habitats and ecosystems, such as land use change, industrialized food production, urbanization and animal trade. While the origin of anthropogenic effects on animal ecology and biogeography can be traced to the Late Pleistocene, the archaeological record—a long-term archive of human–animal–environmental interactions—has largely been untapped in these One Health approaches, thus limiting our understanding of these dynamics over time. In this review, we examine how humans, as niche constructors, have facilitated new host species and ‘disease-scapes’ from the Late Pleistocene to the Anthropocene, by viewing zooarchaeological, bioarchaeological and palaeoecological data with a One Health perspective. We also highlight how new biomolecular tools and advances in the ‘-omics’ can be holistically coupled with archaeological and palaeoecological reconstructions in the service of studying zoonotic disease emergence and re-emergence.

niche construction theory
zoonotic diseases
disease ecology


Contrasting terrestrial and marine ecospace dynamics after the end-Triassic mass extinction event

Abstract: Mass extinctions have fundamentally altered the structure of the biosphere throughout Earth's history. The ecological severity of mass extinctions is well studied in marine ecosystems by categorizing marine taxa into functional groups based on ‘ecospace’ approaches, but the ecological response of terrestrial ecosystems to mass extinctions is less well understood due to the lack of a comparable methodology. Here, we present a new terrestrial ecospace framework that categorizes fauna into functional groups as defined by tiering, motility and feeding traits. We applied the new terrestrial and traditional marine ecospace analyses to data from the Paleobiology Database across the end-Triassic mass extinction—a time of catastrophic global warming—to compare changes between the marine and terrestrial biospheres. We found that terrestrial functional groups experienced higher extinction severity, that taxonomic and functional richness are more tightly coupled in the terrestrial, and that the terrestrial realm continued to experience high ecological dissimilarity in the wake of the extinction. Although signals of extinction severity and ecological turnover are sensitive to the quality of the terrestrial fossil record, our findings suggest greater ecological pressure from the end-Triassic mass extinction on terrestrial ecosystems than marine ecosystems, contributing to more prolonged terrestrial ecological flux.

palaeoecology
functional ecology
ecospace
triassic–jurassic
mass extinction


Evolutionary ecomorphology for the twenty-first century: examples from mammalian carnivores

Abstract: Carnivores (cats, dogs and kin) are a diverse group of mammals that inhabit a remarkable range of ecological niches. While the relationship between ecology and morphology has long been of interest in carnivorans, the application of quantitative techniques has resulted in a recent explosion of work in the field. Therefore, they provide a case study of how quantitative techniques, such as geometric morphometrics (GMM), have impacted our ability to tease apart complex ecological signals from skeletal anatomy, and the implications for our understanding of the relationships between form, function and ecological specialization. This review provides a synthesis of current research on carnivoran ecomorphology, with the goal of illustrating the complex interaction between ecology and morphology in the skeleton. We explore the ecomorphological diversity across major carnivoran lineages and anatomical systems. We examine cranial elements (skull, sensory systems) and postcranial elements (limbs, vertebral column) to reveal mosaic patterns of adaptation related to feeding and hunting strategies, locomotion and habitat preference. We highlight the crucial role that new approaches have played in advancing our understanding of carnivoran ecomorphology, while addressing challenges that remain in the field, such as ecological classifications, form–function relationships and multi-element analysis, offering new avenues for future research.

carnivora
ecomorphology
geometric morphometrics


From teeth to pad: tooth loss and development of keratinous structures in sirenians

Abstract: Sirenians are a well-known example of morphological adaptation to a shallow-water grazing diet characterized by a modified feeding apparatus and orofacial morphology. Such adaptations were accompanied by an anterior tooth reduction associated with the development of keratinized pads, the evolution of which remains elusive. Among sirenians, the recently extinct Steller's sea cow represents a special case for being completely toothless. Here, we used μ-CT scans of sirenian crania to understand how motor-sensor systems associated with tooth innervation responded to innovations such as keratinized pads and continuous dental replacement. In addition, we surveyed nine genes associated with dental reduction for signatures of loss of function. Our results reveal how patterns of innervation changed with modifications of the dental formula, especially continuous replacement in manatees. Both our morphological and genomic data show that dental development was not completely lost in the edentulous Steller's sea cows. By tracing the phylogenetic history of tooth innervation, we illustrate the role of development in promoting the innervation of keratinized pads, similar to the secondary use of dental canals for innervating neomorphic keratinized structures in other tetrapod groups.

sirenians
tooth loss
keratinous pad
steller’ssea cow
dental pseudogenes


A Morrison stem gekkotan reveals gecko evolution and Jurassic biogeography

Abstract: Geckos are a speciose and globally distributed clade of Squamata (lizards, including snakes and amphisbaenians) that are characterized by a host of modifications for nocturnal, scansorial and insectivorous ecologies. They are among the oldest divergences in the lizard crown, so understanding the origin of geckoes (Gekkota) is essential to understanding the origin of Squamata, the most species-rich extant tetrapod clade. However, the poor fossil record of gekkotans has obscured the sequence and timing of the assembly of their distinctive morphology. Here, we describe the first North American stem gekkotan based on a three-dimensionally preserved skull from the Morrison Formation of western North America. Despite its Late Jurassic age, the new species already possesses several key characteristics of the gekkotan skull along with retained ancestral features. We show that this new stem gekkotan, and several previously named species of uncertain phylogenetic relationships, comprise a widespread clade of early crown lizards, substantiating faunal homogeneity in Laurasia during the Late Jurassic that extended across disparate ecological, body-size and physiological classes.

squamata gekkota phylogenetics macroevolution biogeography


Quantifying the complexity of plant reproductive structures reveals a history of morphological and functional integration

Abstract: Vascular plant reproductive structures have undoubtedly become more complex through time, evolving highly differentiated parts that interact in specialized ways. But quantifying these patterns at broad scales is challenging because lineages produce disparate reproductive structures that are often difficult to compare and homologize. We develop a novel approach for analysing interactions within reproductive structures using networks, treating component parts as nodes and a suite of physical and functional interactions among parts as edges. We apply this approach to the plant fossil record, showing that interactions have generally increased through time and that the concentration of these interactions has shifted towards differentiated surrounding organs, resulting in more compact, functionally integrated structures. These processes are widespread across plant lineages, but their extent and timing vary with reproductive biology; in particular, seed-producing structures show them more strongly than spore or pollen-producing structures. Our results demonstrate that major reproductive innovations like the origin of seeds and angiospermy were associated with increased integration through greater interactions among parts. But they also reveal that for certain groups, particularly Mesozoic gymnosperms, millions of years elapsed between the origin of reproductive innovations and increased interactions among parts within their reproductive structures.

macroevolution networks gymnosperms angiosperms


Large-scale application of palaeoproteomics (Zooarchaeology by Mass Spectrometry; ZooMS) in two Palaeolithic faunal assemblages from China

Abstract: The application of Zooarchaeology by Mass Spectrometry (ZooMS) on Pleistocene sites in Europe and northern Asia has resulted in the discovery of important new hominin fossils and has expanded the range of identified fauna. However, no systematic, large-scale application of ZooMS on Palaeolithic sites in East Asia has been attempted thus far. Here, we analyse 866 morphologically non-diagnostic bones from Jinsitai Cave in northeast China and Yumidong Cave in South China, from archaeological horizons dating to 150–10 ka BP. Bones from both sites revealed a high degree of collagen preservation and potentially time-related deamidation patterns, despite being located in very distinct environmental settings. At Jinsitai, we identified 31 camel bones, five of which were radiocarbon dated to 37–20 ka BP. All dated specimens correspond to colder periods of Marine Isotope Stages 3 and 2. We regard the presence of camels at Jinsitai as evidence of wild camels being a megafauna taxon targeted, most likely by early modern humans, during their expansion across northeast Asia. This large-scale application of ZooMS in China highlights the potential of the method for furthering our knowledge of the palaeoanthropological and zooarchaeological records of East Asia.

zooms
palaeoproteomics
deamidation
radiocarbon dating
palaeolithic
camels


Colonial green algae in the Cambrian plankton

Abstract: The fossil record indicates a major turnover in marine phytoplankton across the Ediacaran–Cambrian transition, coincident with the rise of animal-rich ecosystems. However, the diversity, affinities and ecologies of Cambrian phytoplankton are poorly understood, leaving unclear the role of animal interactions and the drivers of diversification. New exceptionally preserved acritarchs (problematic organic-walled microfossils) from the late early Cambrian (around 510 Ma) reveal colonial organization characterized by rings and plates of interconnected, geometrically arranged cells. The assemblage exhibits a wide but gradational variation in cell size, ornamentation and intercell connection, interpreted as representing one or more species with determinate (coenobial) colony formation via cell division, aggregation and growth by cell expansion. An equivalent strategy is known only among green algae, specifically chlorophycean chlorophytes. The fossils differ in detail from modern freshwater examples and apparently represent an earlier convergent radiation in marine settings. Known trade-offs between sinking risk and predator avoidance in colonial phytoplankton point to adaptations triggered by intensifying grazing pressure during a Cambrian metazoan invasion of the water column. The new fossils reveal that not all small acritarchs are unicellular resting cysts, and support an early Palaeozoic prominence of green algal phytoplankton as predicted by molecular biomarkers.

colonial
algae
cambrian
plankton
phytoplankton
cambrian explosion


Musculature of an Early Cambrian cycloneuralian animal

Abstract: Cycloneuralians are ecdysozoans with a fossil record extending to the Early Cambrian Fortunian Age and represented mostly by cuticular integuments. However, internal anatomies of Fortunian cycloneuralians are virtually unknown, hampering our understanding of their functional morphology and phylogenetic relationships. Here we report the exceptional preservation of cycloneuralian introvert musculature in Fortunian rocks of South China. The musculature consists of an introvert body-wall muscular grid of four circular and 36 radially arranged longitudinal muscle bundles, as well as an introvert circular muscle associated with 19 roughly radially arranged, short retractors. Collectively, these features support at least a scalidophoran affinity, and the absence of muscles associated with a mouth cone and scalids further indicates a priapulan affinity. As in modern scalidophorans, the fossil musculature, and particularly the introvert circular muscle retractors, may have controlled introvert inversion and facilitated locomotion and feeding. This work supports the evolution of scalidophoran-like or priapulan-like introvert musculature in cycloneuralians at the beginning of the Cambrian Period.

introvert musculature
priapulida
scalidophora
cycloneuralia
cambrian fortunian
south china


The origins of colour patterns in fossil insects revealed by maturation experiments

Abstract: Many fossil insects show monochromatic colour patterns that may provide valuable insights into ancient insect behaviour and ecology. Whether these patterns reflect original pigmentary coloration is, however, unknown, and their formation mechanism has not been investigated. Here, we performed thermal maturation experiments on extant beetles with melanin-based colour patterns. Scanning electron microscopy reveals that melanin-rich cuticle is more resistant to degradation than melanin-poor cuticle: with progressive maturation, melanin-poor cuticle regions experience preferential thinning and loss, yet melanin-rich cuticle remains. Comparative analysis of fossil insects with monotonal colour patterns confirms that the variations in tone correspond to variations in preserved cuticle thickness. These preserved colour patterns can thus be plausibly explained as melanin-based patterning. Recognition of melanin-based colour patterns in fossil insects opens new avenues for interpreting the evolution of insect coloration and behaviour through deep time.

fossil insect
colour pattern
melanin
taphonomy
insect cuticle
maturation experiments


Infectious diseases may have arrested the southward advance of microblades in Upper Palaeolithic East Asia

Abstract: An unsolved archaeological puzzle of the East Asian Upper Palaeolithic is why the southward expansion of an innovative lithic technology represented by microblades stalled at the Qinling–Huaihe Line. It has been suggested that the southward migration of foragers with microblades stopped there, which is consistent with ancient DNA studies showing that populations to the north and south of this line had differentiated genetically by 19 000 years ago. Many infectious pathogens are believed to have been associated with hominins since the Palaeolithic, and zoonotic pathogens in particular are prevalent at lower latitudes, which may have produced a disease barrier. We propose a mathematical model to argue that mortality due to infectious diseases may have arrested the wave-of-advance of the technologically advantaged foragers from the north.

microblades
ancient dna
diffusion equation
infectious disease
qinling–huaihe line


Fossilization can mislead analyses of phenotypic disparity

Abstract: Analyses of morphological disparity can incorporate living and fossil taxa to facilitate the exploration of how phenotypic variation changes through time. However, taphonomic processes introduce non-random patterns of data loss in fossil data and their impact on perceptions of disparity is unclear. To address this, we characterize how measures of disparity change when simulated and empirical data are degraded through random and structured data loss. We demonstrate that both types of data loss can distort the disparity of clades, and that the magnitude and direction of these changes varies between the most commonly employed distance metrics and disparity indices. The inclusion of extant taxa and exceptionally preserved fossils mitigates these distortions and clarifies the full extent of the data lost, most of which would otherwise go uncharacterized. This facilitates the use of ancestral state estimation and evolutionary simulations to further control for the effects of data loss. Where the addition of such reference taxa is not possible, we urge caution in the extrapolation of general patterns in disparity from datasets that characterize subsets of phenotype, which may represent no more than the traits that they sample.

disparity morphology simulation fossilization decay biostratinomy