Search

BackgroundMegaloolithid eggs have long been associated with sauropod dinosaurs. Despite their extensive and worldwide fossil record, interpretations of egg size and shape, clutch morphology, and incubation strategy vary. The Pinyes locality in the Upper Cretaceous Tremp Formation in the southern Pyrenees, Catalonia provides new information for addressing these issues. Nine horizons containing Megaloolithus siruguei clutches are exposed near the village of Coll de Nargó. Tectonic deformation in the study area strongly influenced egg size and shape, which could potentially lead to misinterpretation of reproductive biology if 2D and 3D maps are not corrected for bed dip that results from tectonism.Methodology/findingsDetailed taphonomic study and three-dimensional modelling of fossil eggs show that intact M. siruguei clutches contained 20-28 eggs, which is substantially larger than commonly reported from Europe and India. Linear and grouped eggs occur in three superimposed levels and form an asymmetric, elongate, bowl-shaped profile in lateral view. Computed tomography data support previous interpretations that the eggs hatched within the substrate. Megaloolithid clutch sizes reported from other European and Indian localities are typically less than 15 eggs; however, these clutches often include linear or grouped eggs that resemble those of the larger Pinyes clutches and may reflect preservation of incomplete clutches.Conclusions/significanceWe propose that 25 eggs represent a typical megaloolithid clutch size and smaller egg clusters that display linear or grouped egg arrangements reported at Pinyes and other localities may represent eroded remnants of larger clutches. The similarity of megaloolithid clutch morphology from localities worldwide strongly suggests common reproductive behaviour. The distinct clutch geometry at Pinyes and other localities likely resulted from the asymmetrical, inclined, and laterally compressed titanosaur pes unguals of the female, using the hind foot for scratch-digging during nest excavation.

We consider sex differences in human facial morphology in the context of developmental change. We show that at puberty, the height of the upper face, between the lip and the brow, develops differently in males and females, and that these differences are not explicable in terms of sex differences in body size. We find the same dimorphism in the faces of human ancestors. We propose that the relative shortening in men and lengthening in women of the anterior upper face at puberty is the mechanistic consequence of extreme maxillary rotation during ontogeny. A link between this developmental model and sexual dimorphism is made for the first time, and provides a new set of morphological criteria to sex human crania. This finding has important implications for the role of sexual selection in the evolution of anthropoid faces and for theories of human facial attractiveness.

Integration into the nuclear genome of germ line cells can lead to vertical inheritance of retroviral genes as host alleles. For other viruses, germ line integration has only rarely been documented. Nonetheless, we identified endogenous viral elements (EVEs) derived from ten non-retroviral families by systematic in silico screening of animal genomes, including the first endogenous representatives of double-stranded RNA, reverse-transcribing DNA, and segmented RNA viruses, and the first endogenous DNA viruses in mammalian genomes. Phylogenetic and genomic analysis of EVEs across multiple host species revealed novel information about the origin and evolution of diverse virus groups. Furthermore, several of the elements identified here encode intact open reading frames or are expressed as mRNA. For one element in the primate lineage, we provide statistically robust evidence for exaptation. Our findings establish that genetic material derived from all known viral genome types and replication strategies can enter the animal germ line, greatly broadening the scope of paleovirological studies and indicating a more significant evolutionary role for gene flow from virus to animal genomes than has previously been recognized., Author Summary The presence of retrovirus sequences in animal genomes has been recognized since the 1970s, but is readily explained by the fact that these viruses integrate into chromosomal DNA as part of their normal replication cycle. Unexpectedly, however, we identified a large and diverse population of sequences in animal genomes that are derived from non-retroviral viruses. Analysis of these sequences—which represent all known virus genome types and replication strategies—reveals new information about the evolutionary history of viruses, in many cases providing the first and only direct evidence for their ancient origins. Additionally, we provide evidence that the functionality of one of these sequences has been maintained in the host genome over many millions of years, raising the possibility that captured viral sequences may have played a larger than expected role in host evolution.

Perhaps more than any other biological discipline, the study of animal viruses is confined to the present. Virions are simply not the stuff of which robust fossils are made. Phylogenetic analysis can help by revealing deep relationships between extant viral lineages, yet such reconstructions lack detail (telling us nothing about transitional or extinct viral forms, the movement of viruses between species, or the timing of major events in viral evolution), and molecular clock estimates are notoriously imprecise when applied to viruses [1]. Until recently, ancient endogenous retroviruses (ERVs) were the closest thing to a fossil record available to scientists with a proclivity for combining virology and natural history. Happily, a trio of recent studies appearing in PLoS Genetics [2], PLoS Biology [3], and PLoS Pathogens [4] reveal an unexpected wealth of non-retroviral virus sequences embedded in the genome sequence databases, a virtual equivalent of the Burgess Shale, ripe for excavation by eager paleovirologists. Retroviral infection occasionally results in the deposition of a provirus in a host's germline DNA. While germline integration of a provirus may be an exceedingly rare event, across the great expanse of evolutionary time millions of ERV loci have accumulated in animal genomes. Because retroviruses replicate through an integrated DNA intermediate, it is not difficult to imagine how ERVs are generated. For other animal viruses, which do not normally integrate their genomes into host DNA, the formation of germline insertions should be far less likely. Nonetheless, reports of non-retroviral specimens being unearthed from the genomes of animal species are on the rise. Notable examples include functional expression of nudivirus-related structural genes in the genomes of parasitic wasps [5]; Ebolavirus-like sequences, related to modern filoviruses, present in multiple mammalian genomes [6]; and sequences resembling the Bornavirus nucleoprotein gene (N) in the genomes of various mammals including primates, rodents, and elephants [7]. Even some herpesviruses have a propensity for occasional germline insertion and thus, the potential for vertical inheritance [8]. Now, Belyi et al. [4] and Katzourakis and Gifford [2], have unearthed diverse collections of non-retroviral sequences buried in whole genome sequence data from an impressive array of host organisms, including mammals, marsupials, birds, rodents, and insects, using modern viral sequences as bioinformatic probes. A third study from Gilbert and Feschotte specifically reevaluates the macroevolution of hepadnaviruses based on the sequence and distribution of hepadnavirus-like fossils in the genomes of passerine birds [3]. To cope with this newfound abundance, the authors of one of the studies suggest the acronym EVE (for endogenous viral element) as a general term to encompass all virus-derived genomic loci [2]. Two of the studies also took a closer look at a previously described class of EVEs, called EBLNs (for endogenous Bornavirus-like N genes) [2], [4], [7]. While most EVEs were either defective at the time of insertion or rendered functionless by the accumulation of random mutations over the course of millions of years, EBLNs are striking in retaining largely intact protein-coding sequences. In fact, in silico simulations of EBLN evolution estimate that these elements should have accumulated ∼10–20 stop codons since the time of genome insertion. That the EBLN coding sequences appear relatively unscathed suggests that these particular elements provide (or at times provided) a selectively advantageous function, subjecting them to purifying selection. The possibility is not without precedent: for example, at least one human ERV has evolved to provide a cellular function [9], and there are several examples of ERVs that have been subverted by host evolution to serve as inhibitors of retroviral infection [10]–[14]. As a group, viruses are polyphyletic, as evidenced by the variety of unique genome types and distinctive replication strategies they collectively employ. There are double-stranded DNA viruses and single-stranded DNA viruses, double-stranded and single-stranded RNA viruses, and viruses with segmented genomes; among those with single-stranded RNA, there are those with positive polarity (the genome resembles an mRNA) and those with negative sense genomes. Each genome type represents a different starting point for takeover of the host cell, and each requires a different strategy for achieving this fundamental task. For example, replication of some viruses is confined entirely to the cytoplasm, whereas others involve synthesis of DNA or RNA in the nucleus. While the fossil record is still dominated by retroviral sequences, the inventory of known EVE loci now appears to include representatives of all the basic replication strategies exemplified by modern viruses. Non-retroviral EVEs are typically subgenomic, derived from just one or a few viral genes instead of entire viral genomes. Insertion site duplications bracketing some EVEs suggest that retrotransposition in trans, by retrotransposons or possibly retroviruses, may be a predominant mechanism of EVE formation. In fact, for RNA viruses that replicate in the cytoplasm (e.g., filoviruses and rhabdoviruses), retrotransposition is the most plausible mechanism for EVE formation. In such cases, it will be interesting to determine whether the more abundant EVE sequences share some common feature(s) conferring a propensity for retrotransposition. In contrast, hepadnavirus “fossils” lack the hallmarks of retrotransposition (such as flanking insertion-site duplications and poly-A tails), and may instead have resulted from non-homologous end joining and insertion of viral DNA directly into the host genome [3]. When incorporated into phylogenetic trees, many EVEs group as sister taxa to their modern counterparts. Thus, they are not evolutionary intermediates on the path to extant viruses, but rather extinct lineages sharing a common ancestor with modern viruses. From this, one can infer that most of the distinctive replication strategies employed by modern viruses probably originated hundreds of millions of years ago. While virologists intuitively understand this (given the widespread distribution of viruses among living organisms), EVEs constitute direct, physical evidence that modern viral lineages have very ancient roots (Table 1). That modern viruses and ancient EVE sequences are still recognizably related is astonishing, given that they are separated by millions of years of exogenous viral evolution. Table 1 Estimated Minimum Age of Select EVEs. The catalog of EVEs is impressive for what it contains, but even more so for what it does not. Why? Because the known EVEs probably represent a minor and highly skewed sampling of viral prehistory. Minor, because the odds that infection of an individual organism will result in fixation of an EVE are exceedingly small (Figure 1). Skewed, because some viruses may be more prone to germline insertion than others (that retroviral insertions greatly outnumber other EVEs is a particularly striking example of a virus-dependent bias). Thus, as impressive in scope and variety as the EVEs are, they may represent but a drop in the ocean of all the viruses that have buffeted host organisms across the ages. Figure 1 Formation of a hypothetical EVE and relationship to modern viruses. The current EVE record may have other limitations. Just how far back does the EVE fossil record extend? Erosion due to the steady accumulation of mutations must impose an upper limit on how far back the viral fossil record can be deciphered, and theoretical predictions of that limit would be useful. Even in the absence of sequence degradation, some EVEs may be easier to detect than others. For example, the studies described here relied on known viral sequences as queries: if our genomes also harbor ancient viral sequences for which there is no modern counterpart, how would we recognize them for what they once were?

BackgroundIn the Plio-Pleistocene, the hominin foot evolved from a grasping appendage to a stiff, propulsive lever. Central to this transition was the development of the longitudinal arch, a structure that helps store elastic energy and stiffen the foot during bipedal locomotion. Direct evidence for arch evolution, however, has been somewhat elusive given the failure of soft-tissue to fossilize. Paleoanthropologists have relied on footprints and bony correlates of arch development, though little consensus has emerged as to when the arch evolved.Methodology/principal findingsHere, we present evidence from radiographs of modern humans (n = 261) that the set of the distal tibia in the sagittal plane, henceforth referred to as the tibial arch angle, is related to rearfoot arching. Non-human primates have a posteriorly directed tibial arch angle, while most humans have an anteriorly directed tibial arch angle. Those humans with a posteriorly directed tibial arch angle (8%) have significantly lower talocalcaneal and talar declination angles, both measures of an asymptomatic flatfoot. Application of these results to the hominin fossil record reveals that a well developed rearfoot arch had evolved in Australopithecus afarensis. However, as in humans today, Australopithecus populations exhibited individual variation in foot morphology and arch development, and "Lucy" (A.L. 288-1), a 3.18 Myr-old female Australopithecus, likely possessed asymptomatic flat feet. Additional distal tibiae from the Plio-Pleistocene show variation in tibial arch angles, including two early Homo tibiae that also have slightly posteriorly directed tibial arch angles.Conclusions/significanceThis study finds that the rearfoot arch was present in the genus Australopithecus. However, the female Australopithecus afarensis "Lucy" has an ankle morphology consistent with non-pathological flat-footedness. This study suggests that, as in humans today, there was variation in arch development in Plio-Pleistocene hominins.

The size and flight mechanics of giant pterosaurs have received considerable research interest for the last century but are confused by conflicting interpretations of pterosaur biology and flight capabilities. Avian biomechanical parameters have often been applied to pterosaurs in such research but, due to considerable differences in avian and pterosaur anatomy, have lead to systematic errors interpreting pterosaur flight mechanics. Such assumptions have lead to assertions that giant pterosaurs were extremely lightweight to facilitate flight or, if more realistic masses are assumed, were flightless. Reappraisal of the proportions, scaling and morphology of giant pterosaur fossils suggests that bird and pterosaur wing structure, gross anatomy and launch kinematics are too different to be considered mechanically interchangeable. Conclusions assuming such interchangeability--including those indicating that giant pterosaurs were flightless--are found to be based on inaccurate and poorly supported assumptions of structural scaling and launch kinematics. Pterosaur bone strength and flap-gliding performance demonstrate that giant pterosaur anatomy was capable of generating sufficient lift and thrust for powered flight as well as resisting flight loading stresses. The retention of flight characteristics across giant pterosaur skeletons and their considerable robustness compared to similarly-massed terrestrial animals suggest that giant pterosaurs were not flightless. Moreover, the term 'giant pterosaur' includes at least two radically different forms with very distinct palaeoecological signatures and, accordingly, all but the most basic sweeping conclusions about giant pterosaur flight should be treated with caution. Reappraisal of giant pterosaur material also reveals that the size of the largest pterosaurs, previously suggested to have wingspans up to 13 m and masses up to 544 kg, have been overestimated. Scaling of fragmentary giant pterosaur remains have been misled by distorted fossils or used inappropriate scaling techniques, indicating that 10-11 m wingspans and masses of 200-250 kg are the most reliable upper estimates of known pterosaur size.

The polar bear is the only living ursid with a fully carnivorous diet. Despite a number of well-documented craniodental adaptations for a diet of seal flesh and blubber, molecular and paleontological data indicate that this morphologically distinct species evolved less than a million years ago from the omnivorous brown bear. To better understand the evolution of this dietary specialization, we used phylogenetic tests to estimate the rate of morphological specialization in polar bears. We then used finite element analysis (FEA) to compare the limits of feeding performance in the polar bear skull to that of the phylogenetically and geographically close brown bear. Results indicate that extremely rapid evolution of semi-aquatic adaptations and dietary specialization in the polar bear lineage produced a cranial morphology that is weaker than that of brown bears and less suited to processing tough omnivorous or herbivorous diets. Our results suggest that continuation of current climate trends could affect polar bears by not only eliminating their primary food source, but also through competition with northward advancing, generalized brown populations for resources that they are ill-equipped to utilize.

BackgroundThe ceratopsians represent one of the last dinosaurian radiations. Traditionally the only universally accepted speciose clade within the group was the Ceratopsidae. However, recent discoveries and phylogenetic analyses have led to the recognition of a new speciose clade, the Leptoceratopsidae, which is predominantly known from the Upper Cretaceous of North America.Methodology/principal findingsHere we report a new leptoceratopsid taxon, Zhuchengceratops inexpectus gen. et sp. nov., based on a partial, articulated skeleton recovered from the Upper Cretaceous Wangshi Group of Zhucheng, Shandong Province, China. Although Zhuchengceratops is significantly different from other known leptoceratopsids, it is recovered as a derived member of the group by our phylogenetic analysis. Furthermore, Zhuchengceratops exhibits several features previously unknown in leptoceratopsids but seen in ceratopsids and their close relatives, suggesting that the distribution of morphological features within ceratopsians is more complex than previously realized.Conclusion/significanceThe discovery of Zhuchengceratops increases both the taxonomic diversity and the morphological disparity of the Leptoceratopsidae, providing further support for the hypothesis that this clade represents a successful radiation of horned dinosaurs in parallel with the Ceratopsidae in the Late Cretaceous. This documents a surprising case of the coexistence and radiation of two closely-related lineages with contrasting suites of jaw and dental features that probably reflect adaptation to different food resources.

BackgroundBasal iguanodontian dinosaurs were extremely successful animals, found in great abundance and diversity almost worldwide during the Early Cretaceous. In contrast to Europe and Asia, the North American record of Early Cretaceous basal iguanodonts has until recently been limited largely to skulls and skeletons of Tenontosaurus tilletti.Methodology/principal findingsHerein we describe two new basal iguanodonts from the Yellow Cat Member of the Cedar Mountain Formation of eastern Utah, each known from a partial skull and skeleton. Iguanacolossus fortis gen. et sp. nov. and Hippodraco scutodens gen. et sp. nov. are each diagnosed by a single autapomorphy and a unique combination of characters.Conclusions/significanceIguanacolossus and Hippodraco add greatly to our knowledge of North American basal iguanodonts and prompt a new comprehensive phylogenetic analysis of basal iguanodont relationships. This analysis indicates that North American Early Cretaceous basal iguanodonts are more basal than their contemporaries in Europe and Asia.

The extant beaver, Castor, has played an important role shaping landscapes and ecosystems in Eurasia and North America, yet the origins and early evolution of this lineage remain poorly understood. Here we use a geometric morphometric approach to help re-evaluate the phylogenetic affinities of a fossil skull from the Late Miocene of China. This specimen was originally considered Sinocastor, and later transferred to Castor. The aim of this study was to determine whether this form is an early member of Castor, or if it represents a lineage outside of Castor. The specimen was compared to 38 specimens of modern Castor (both C. canadensis and C. fiber) as well as fossil specimens of C. fiber (Pleistocene), C. californicus (Pliocene) and the early castorids Steneofiber eseri (early Miocene). The results show that the specimen falls outside the Castor morphospace and that compared to Castor, Sinocastor possesses a: 1) narrower post-orbital constriction, 2) anteroposteriorly shortened basioccipital depression, 3) shortened incisive foramen, 4) more posteriorly located palatine foramen, 5) longer rostrum, and 6) longer braincase. Also the specimen shows a much shallower basiocciptal depression than what is seen in living Castor, as well as prominently rooted molars. We conclude that Sinocastor is a valid genus. Given the prevalence of apparently primitive traits, Sinocastor might be a near relative of the lineage that gave rise to Castor, implying a possible Asiatic origin for Castor.

BackgroundMineralized and permineralized bone is the most common form of fossilization in the vertebrate record. Preservation of gross soft tissues is extremely rare, but recent studies have suggested that primary soft tissues and biomolecules are more commonly preserved within preserved bones than had been presumed. Some of these claims have been challenged, with presentation of evidence suggesting that some of the structures are microbial artifacts, not primary soft tissues. The identification of biomolecules in fossil vertebrate extracts from a specimen of Brachylophosaurus canadensis has shown the interpretation of preserved organic remains as microbial biofilm to be highly unlikely. These discussions also propose a variety of potential mechanisms that would permit the preservation of soft-tissues in vertebrate fossils over geologic time.Methodology/principal findingsThis study experimentally examines the role of microbial biofilms in soft-tissue preservation in vertebrate fossils by quantitatively establishing the growth and morphology of biofilms on extant archosaur bone. These results are microscopically and morphologically compared with soft-tissue extracts from vertebrate fossils from the Hell Creek Formation of southeastern Montana (Latest Maastrichtian) in order to investigate the potential role of microbial biofilms on the preservation of fossil bone and bound organic matter in a variety of taphonomic settings. Based on these analyses, we highlight a mechanism whereby this bound organic matter may be preserved.Conclusions/significanceResults of the study indicate that the crystallization of microbial biofilms on decomposing organic matter within vertebrate bone in early taphonomic stages may contribute to the preservation of primary soft tissues deeper in the bone structure.

Background The evidence of several forms of arthritis has been well documented in the fossil record. However, for pre-Cenozoic vertebrates, especially regarding reptiles, this record is rather scarce. In this work we present a case report of spondarthritis found in a vertebral series that belonged to a carnivorous archosaurian reptile from the Lower Triassic (~245 million years old) of the South African Karoo. Methodology/Principal Findings Neutron tomography confirmed macroscopic data, revealing the ossification of the entire intervertebral disc space (both annulus fibrosus and nucleus pulposus), which supports the diagnosis of spondarthritis. Conclusions/Significance The presence of spondarthritis in the new specimen represents by far the earliest evidence of any form of arthritis in the fossil record. The present find is nearly 100 million years older than the previous oldest report of this pathology, based on a Late Jurassic dinosaur. Spondarthritis may have indirectly contributed to the death of the animal under study.

BackgroundDebate regarding the monophyly and relationships of the avian order Pelecaniformes represents a classic example of discord between morphological and molecular estimates of phylogeny. This lack of consensus hampers interpretation of the group's fossil record, which has major implications for understanding patterns of character evolution (e.g., the evolution of wing-propelled diving) and temporal diversification (e.g., the origins of modern families). Relationships of the Pelecaniformes were inferred through parsimony analyses of an osteological dataset encompassing 59 taxa and 464 characters. The relationships of the Plotopteridae, an extinct family of wing-propelled divers, and several other fossil pelecaniforms (Limnofregata, Prophaethon, Lithoptila, ?Borvocarbo stoeffelensis) were also assessed. The antiquity of these taxa and their purported status as stem members of extant families makes them valuable for studies of higher-level avian diversification.Methodology/principal findingsPelecaniform monophyly is not recovered, with Phaethontidae recovered as distantly related to all other pelecaniforms, which are supported as a monophyletic Steganopodes. Some anatomical partitions of the dataset possess different phylogenetic signals, and partitioned analyses reveal that these discrepancies are localized outside of Steganopodes, and primarily due to a few labile taxa. The Plotopteridae are recovered as the sister taxon to Phalacrocoracoidea, and the relationships of other fossil pelecaniforms representing key calibration points are well supported, including Limnofregata (sister taxon to Fregatidae), Prophaethon and Lithoptila (successive sister taxa to Phaethontidae), and ?Borvocarbo stoeffelensis (sister taxon to Phalacrocoracidae). These relationships are invariant when 'backbone' constraints based on recent avian phylogenies are imposed.Conclusions/significanceRelationships of extant pelecaniforms inferred from morphology are more congruent with molecular phylogenies than previously assumed, though notable conflicts remain. The phylogenetic position of the Plotopteridae implies that wing-propelled diving evolved independently in plotopterids and penguins, representing a remarkable case of convergent evolution. Despite robust support for the placement of fossil taxa representing key calibration points, the successive outgroup relationships of several "stem fossil + crown family" clades are variable and poorly supported across recent studies of avian phylogeny. Thus, the impact these fossils have on inferred patterns of temporal diversification depends heavily on the resolution of deep nodes in avian phylogeny.

Human remains, some as recent as 15 thousand years, from Liang Bua (LB) on the Indonesian island of Flores have been attributed to a new species, Homo floresiensis. The definition includes a mosaic of features, some like modern humans (hence derived: genus Homo), some like modern apes and australopithecines (hence primitive: not species sapiens), and some unique (hence new species: floresiensis). Conversely, because only modern humans (H. sapiens) are known in this region in the last 40 thousand years, these individuals have also been suggested to be genetic human dwarfs. Such dwarfs resemble small humans and do not show the mosaic combination of the most complete individuals, LB1 and LB6, so this idea has been largely dismissed. We have previously shown that some features of the cranium of hypothyroid cretins are like those of LB1. Here we examine cretin postcrania to see if they show anatomical mosaics like H. floresiensis. We find that hypothyroid cretins share at least 10 postcranial features with Homo floresiensis and unaffected humans not found in apes (or australopithecines when materials permit). They share with H. floresiensis, modern apes and australopithecines at least 11 postcranial features not found in unaffected humans. They share with H. floresiensis, at least 8 features not found in apes, australopithecines or unaffected humans. Sixteen features can be rendered metrically and multivariate analyses demonstrate that H. floresiensis co-locates with cretins, both being markedly separate from humans and chimpanzees (P0.999). We therefore conclude that LB1 and LB6, at least, are, most likely, endemic cretins from a population of unaffected Homo sapiens. This is consistent with recent hypothyroid endemic cretinism throughout Indonesia, including the nearby island of Bali.

Background Stramenopiles constitute a large and diverse eukaryotic clade that is currently poorly characterized from both phylogenetic and temporal perspectives at deeper taxonomic levels. To better understand this group, and in particular the photosynthetic stramenopiles (Ochrophyta), we analyzed sequence data from 135 taxa representing most major lineages. Our analytical approach utilized several recently developed methods that more realistically model the temporal evolutionary process. Methodology/Principal Findings Phylogenetic reconstruction employed a Bayesian joint rate- and pattern-heterogeneity model to reconstruct the evolutionary history of these taxa. Inferred phylogenetic resolution was generally high at all taxonomic levels, sister-class relationships in particular receiving good statistical support. A signal for heterotachy was detected in clustered portions of the tree, although this does not seem to have had a major influence on topological inference. Divergence time estimates, assuming a lognormally-distributed relaxed molecular clock while accommodating topological uncertainty, were broadly congruent over alternative temporal prior distributions. These data suggest that Ochrophyta originated near the Proterozoic-Phanerozoic boundary, diverging from their sister-taxon Oomycota. The evolution of the major ochrophyte lineages appears to have proceeded gradually thereafter, with most lineages coming into existence by ∼200 million years ago. Conclusions/Significance The evolutionary timescale of the autotrophic stramenopiles reconstructed here is generally older than previously inferred from molecular clocks. However, this more ancient timescale nevertheless casts serious doubt on the taxonomic validity of putative xanthophyte/phaeophyte fossils from the Proterozoic, which predate by as much as a half billion years or more the age suggested by our molecular genetic data. If these fossils truly represent crown stramenopile lineages, then this would imply that molecular rate evolution in this group proceeds in a fashion that is fundamentally incompatible with the relaxed molecular clock model employed here. A more likely scenario is that there is considerable convergent morphological evolution within Heterokonta, and that these fossils have been taxonomically misdiagnosed.

Background There have been numerous studies on dinosaur biogeographic distribution patterns. However, these distribution data have not yet been applied to ecological questions. Ecological studies of dinosaurs have tended to focus on reconstructing individual taxa, usually through comparisons to modern analogs. Fewer studies have sought to determine if the ecological structure of fossil assemblages is preserved and, if so, how dinosaur communities varied. Climate is a major component driving differences between communities. If the ecological structure of a fossil locality is preserved, we expect that dinosaur assemblages from similar environments will share a similar ecological structure. Methodology/Principal Findings This study applies Ecological Structure Analysis (ESA) to a dataset of 100+ dinosaur taxa arranged into twelve composite fossil assemblages from around the world. Each assemblage was assigned a climate zone (biome) based on its location. Dinosaur taxa were placed into ecomorphological categories. The proportion of each category creates an ecological profile for the assemblage, which were compared using cluster and principal components analyses. Assemblages grouped according to biome, with most coming from arid or semi-arid/seasonal climates. Differences between assemblages are tied to the proportion of large high-browsing vs. small ground-foraging herbivores, which separates arid from semi-arid and moister environments, respectively. However, the effects of historical, taphonomic, and other environmental factors are still evident. Conclusions/Significance This study is the first to show that the general ecological structure of Late Jurassic dinosaur assemblages is preserved at large scales and can be assessed quantitatively. Despite a broad similarity of climatic conditions, a degree of ecological variation is observed between assemblages, from arid to moist. Taxonomic differences between Asia and the other regions demonstrate at least one case of ecosystem convergence. The proportion of different ecomorphs, which reflects the prevailing climatic and environmental conditions present during fossil deposition, may therefore be used to differentiate Late Jurassic dinosaur fossil assemblages. This method is broadly applicable to different taxa and times, allowing one to address questions of evolutionary, biogeographic, and climatic importance.

BACKGROUND: The cusp homology of Lagomorpha has long been problematic largely because their teeth are highly derived relative to their more typically tribosphenic ancestors. Within this context, the lagomorph central cusp has been particularly difficult to homologize with other tribosphenic cusps; authors have previously considered it the paracone, protocone, metacone, amphicone, or an entirely new cusp. METHODOLOGY/PRINCIPAL FINDINGS: Here we present newly described fossil duplicidentates (Lagomorpha and Mimotonidae) in the context of a well-constrained phylogeny to establish a nomenclatural system for cusps based on the tribosphenic pattern. We show that the central cusp of lagomorphs is homologous with the metaconule of other mammals. We also show that the buccal acquisition of a second cusp on the premolars (molarization) within duplicidentates is atypical with respect to other mammalian lineages; within the earliest lagomorphs, a second buccal cusp is added mesially to an isolated buccal cusp. CONCLUSIONS/SIGNIFICANCE: The distal shift of the 'ancestral' paracone within early duplicidentates amounts to the changing of a paracone into a metacone in these lineages. For this reason, we support a strictly topological approach to cusp names, and suggest a discontinuity in nomenclature to capture the complexity of the interplay between evolutionary history and the developmental process that have produced cusp patterns in duplicidentates.

BackgroundDuring much of the Late Cretaceous, a shallow, epeiric sea divided North America into eastern and western landmasses. The western landmass, known as Laramidia, although diminutive in size, witnessed a major evolutionary radiation of dinosaurs. Other than hadrosaurs (duck-billed dinosaurs), the most common dinosaurs were ceratopsids (large-bodied horned dinosaurs), currently known only from Laramidia and Asia. Remarkably, previous studies have postulated the occurrence of latitudinally arrayed dinosaur "provinces," or "biomes," on Laramidia. Yet this hypothesis has been challenged on multiple fronts and has remained poorly tested.Methodology/principal findingsHere we describe two new, co-occurring ceratopsids from the Upper Cretaceous Kaiparowits Formation of Utah that provide the strongest support to date for the dinosaur provincialism hypothesis. Both pertain to the clade of ceratopsids known as Chasmosaurinae, dramatically increasing representation of this group from the southern portion of the Western Interior Basin of North America. Utahceratops gettyi gen. et sp. nov.-characterized by short, rounded, laterally projecting supraorbital horncores and an elongate frill with a deep median embayment-is recovered as the sister taxon to Pentaceratops sternbergii from the late Campanian of New Mexico. Kosmoceratops richardsoni gen. et sp. nov.-characterized by elongate, laterally projecting supraorbital horncores and a short, broad frill adorned with ten well developed hooks-has the most ornate skull of any known dinosaur and is closely allied to Chasmosaurus irvinensis from the late Campanian of Alberta.Conclusions/significanceConsidered in unison, the phylogenetic, stratigraphic, and biogeographic evidence documents distinct, co-occurring chasmosaurine taxa north and south on the diminutive landmass of Laramidia. The famous Triceratops and all other, more nested chasmosaurines are postulated as descendants of forms previously restricted to the southern portion of Laramidia. Results further suggest the presence of latitudinally arrayed evolutionary centers of endemism within chasmosaurine ceratopsids during the late Campanian, the first documented occurrence of intracontinental endemism within dinosaurs.

The South American phorusrhacid bird radiation comprised at least 18 species of small to gigantic terrestrial predators for which there are no close modern analogs. Here we perform functional analyses of the skull of the medium-sized (,40 kg) patagornithine phorusrhacid Andalgalornis steulleti (upper Miocene-lower Pliocene, Andalgalá Formation, Catamarca, Argentina) to assess its mechanical performance in a comparative context. Based on computed tomographic (CT) scanning and morphological analysis, the skull of Andalgalornis steulleti is interpreted as showing features reflecting loss of intracranial immobility. Discrete anatomical attributes permitting such cranial kinesis are widespread phorusrhacids outgroups, but this is the first clear evidence of loss of cranial kinesis in a gruiform bird and may be among the best documented cases among all birds. This apomorphic loss is interpreted as an adaptation for enhanced craniofacial rigidity, particularly with regard to sagittal loading. We apply a Finite Element approach to a three-dimensional (3D) model of the skull. Based on regression analysis we estimate the bite force of Andalgalornis at the bill tip to be 133 N. Relative to results obtained from Finite Element Analysis of one of its closest living relatives (seriema) and a large predatory bird (eagle), the phorusrhacid's skull shows relatively high stress under lateral loadings, but low stress where force is applied dorsoventrally (sagittally) and in ''pullback'' simulations. Given the relative weakness of the skull mediolaterally, it seems unlikely that Andalgalornis engaged in potentially risky behaviors that involved subduing large, struggling prey with its beak. We suggest that it either consumed smaller prey that could be killed and consumed more safely (e.g., swallowed whole) or that it used multiple well-targeted sagittal strikes with the beak in a repetitive attack-and-retreat strategy., Facultad de Ciencias Naturales y Museo

BACKGROUND: The Green River Formation (early Eocene, about 42-53 Ma) at and near Fossil Butte National Monument in Wyoming, USA, is world famous for its exquisitely preserved freshwater teleost fish in the former Fossil Lake. Nonetheless, trace fossils attributed to fish interacting with the lake bottom are apparently rare, and have not been associated directly with any fish species. Here we interpret the first known feeding and swimming trace fossil of the teleost Notogoneus osculus Cope (Teleostei: Gonorynchidae), which is also represented as a body fossil in the same stratum. METHODOLOGY/PRINCIPAL FINDINGS: A standard description of the trace fossil, identified as Undichna cf. U. simplicatas, was augmented by high-resolution digital images and spatial and mathematical analyses, which allowed for detailed interpretations of the anatomy, swimming mode, feeding behavior, and body size of the tracemaker. Our analysis indicates that the tracemaker was about 45 cm long; used its caudal, anal, and pelvic fins (the posterior half of its body) to make the swimming traces; and used a ventrally oriented mouth to make overlapping feeding marks. We hypothesize that the tracemaker was an adult Notogoneus osculus. CONCLUSIONS/SIGNIFICANCE: Our results are the first to link a specific teleost tracemaker with a trace fossil from the Green River Formation, while also interpreting the size and relative age of the tracemaker. The normal feeding and swimming behaviors indicated by the trace fossil indicate temporarily oxygenated benthic conditions in the deepest part of Fossil Lake, counter to most paleoecological interpretations of this deposit. Lastly, our spatial and mathematical analyses significantly update and advance previous approaches to the study of teleost trace fossils.

BackgroundAs we know from modern species, nursery areas are essential shark habitats for vulnerable young. Nurseries are typically highly productive, shallow-water habitats that are characterized by the presence of juveniles and neonates. It has been suggested that in these areas, sharks can find ample food resources and protection from predators. Based on the fossil record, we know that the extinct Carcharocles megalodon was the biggest shark that ever lived. Previous proposed paleo-nursery areas for this species were based on the anecdotal presence of juvenile fossil teeth accompanied by fossil marine mammals. We now present the first definitive evidence of ancient nurseries for C. megalodon from the late Miocene of Panama, about 10 million years ago.Methodology/principal findingsWe collected and measured fossil shark teeth of C. megalodon, within the highly productive, shallow marine Gatun Formation from the Miocene of Panama. Surprisingly, and in contrast to other fossil accumulations, the majority of the teeth from Gatun are very small. Here we compare the tooth sizes from the Gatun with specimens from different, but analogous localities. In addition we calculate the total length of the individuals found in Gatun. These comparisons and estimates suggest that the small size of Gatun's C. megalodon is neither related to a small population of this species nor the tooth position within the jaw. Thus, the individuals from Gatun were mostly juveniles and neonates, with estimated body lengths between 2 and 10.5 meters.Conclusions/significanceWe propose that the Miocene Gatun Formation represents the first documented paleo-nursery area for C. megalodon from the Neotropics, and one of the few recorded in the fossil record for an extinct selachian. We therefore show that sharks have used nursery areas at least for 10 millions of years as an adaptive strategy during their life histories.

Modern grylloblattids are one of the least diverse of the modern insect orders. The thorax changes in morphology might be associated with the changes of the function of the forelegs, wing loss, changes in behavior and adaptation to habitat. As temperature is the main barrier for migration of modern grylloblattids, the range of each species is extremely limited. The potential distribution areas of grylloblattids remain unclear. A second new species of ice crawlers (Insecta: Grylloblattodea), Grylloblattella cheni Bai, Wang et Yang sp. nov., is described from China. The distribution map and key to species of Grylloblattella are given. A comparison of the thorax of extant and extinct Grylloblattodea is presented, with an emphasis on the pronotum using geometric morphometric analysis, which may reflect thorax adaptation and the evolution of Grylloblattodea. Potential global distribution of grylloblattids is inferred. Highly diversified pronota of extinct Grylloblattodea may reflect diverse habitats and niches. The relatively homogeneous pronota of modern grylloblattids might be explained by two hypotheses: synapomorphy or convergent evolution. Most fossils of Grylloblattodea contain an obviously longer meso- and metathorax than prothorax. The length of the meso- and metathorax of modern grylloblattids is normally shorter than the prothorax. This may be associated with the wing loss, which is accompanied by muscle reduction and changes to the thoracic skeleton system. Threats to grylloblattids and several conservation comments are also provided.

BackgroundThe two forms of white rhinoceros; northern and southern, have had contrasting conservation histories. The Northern form, once fairly numerous is now critically endangered, while the southern form has recovered from a few individuals to a population of a few thousand. Since their last taxonomic assessment over three decades ago, new material and analytical techniques have become available, necessitating a review of available information and re-assessment of the taxonomy.ResultsDental morphology and cranial anatomy clearly diagnosed the southern and northern forms. The differentiation was well supported by dental metrics, cranial growth and craniometry, and corresponded with differences in post-cranial skeleton, external measurements and external features. No distinctive differences were found in the limited descriptions of their behavior and ecology. Fossil history indicated the antiquity of the genus, dating back at least to early Pliocene and evolution into a number of diagnosable forms. The fossil skulls examined fell outside the two extant forms in the craniometric analysis. Genetic divergence between the two forms was consistent across both nuclear and mitochondrial genomes, and indicated a separation of over a million years.ConclusionsOn re-assessing the taxonomy of the two forms we find them to be morphologically and genetically distinct, warranting the recognition of the taxa formerly designated as subspecies; Ceratotherium simum simum the southern form and Ceratotherium simum cottoni the northern form, as two distinct species Ceratotherium simum and Ceratotherium cottoni respectively. The recognition of the northern form as a distinct species has profound implications for its conservation.

A new snake from Upper Cretaceous rocks in India is found with hatchling sauropod dinosaurs, demonstrating that large, gape-limited snakes were probably capable of taking in moderate-sized vertebrate prey., Derived large-mouthed snakes (macrostomatans) possess numerous specializations in their skull and lower jaws that allow them to consume large vertebrate prey. In contrast, basal snakes lack these adaptations and feed primarily on small prey items. The sequence of osteological and behavioral modifications involved in the evolution of the macrostomatan condition has remained an open question because of disagreement about the origin and interrelationships of snakes, the paucity of well-preserved early snake fossils on many continental landmasses, and the lack of information about the feeding ecology of early snakes. We report on a partial skeleton of a new 3.5-m-long snake, Sanajeh indicus gen. et sp. nov., recovered from Upper Cretaceous rocks of western India. S. indicus was fossilized in association with a sauropod dinosaur egg clutch, coiled around an egg and adjacent to the remains of a ca. 0.5-m-long hatchling. Multiple snake-egg associations at the site strongly suggest that S. indicus frequented nesting grounds and preyed on hatchling sauropods. We interpret this pattern as “ethofossil” preservation of feeding behavior. S. indicus lacks specializations of modern egg-eaters and of macrostomatans, and skull and vertebral synapomorphies place it in an intermediate position in snake phylogeny. Sanajeh and its large-bodied madtsoiid sister taxa Yurlunggur camfieldensis and Wonambi naracoortensis from the Neogene of Australia show specializations for intraoral prey transport but lack the adaptations for wide gape that characterize living macrostomatan snakes. The Dholi Dungri fossils are the second definitive association between sauropod eggs and embryonic or hatchling remains. New fossils from western India provide direct evidence of feeding ecology in a Mesozoic snake and demonstrate predation risks for hatchling sauropod dinosaurs. Our results suggest that large body size and jaw mobility afforded some non-macrostomatan snakes a greater diversity of prey items than previously suspected on the basis of extant basal snakes., Author Summary Snakes first appear in the fossil record towards the end of the dinosaur era, approximately 98 million years ago. Snake fossils from that time are fragmentary, usually consisting of parts of the backbone. Relatively complete snake fossils preserving skulls and occasionally hindlimbs are quite rare and have only been found in marine sediments in Afro-Arabia and Europe or in terrestrial sediments in South America. Early snake phylogeny remains controversial, in part because of the paucity of early fossils. We describe a new 3.5-m-long snake from the Late Cretaceous of western India that is preserved in an extraordinary setting—within a sauropod dinosaur nest, coiled around an egg and adjacent the remains of a ca. 0.5-m-long hatchling. Other snake-egg associations at the same site suggest that the new snake frequented nesting grounds and preyed on hatchling sauropods. We named this new snake Sanajeh indicus because of its provenance and its somewhat limited oral gape. Sanajeh broadens the geographical distribution of early snakes and helps resolve their phylogenetic affinities. We conclude that large body size and jaw mobility afforded some early snakes a greater diversity of prey items than previously suspected.

Background: Although variation provides the raw material for natural selection and evolution, few empirical data exist about the factors controlling morphological variation. Because developmental constraints on variation are expected to act by influencing trait correlations, studies of modularity offer promising approaches that quantify and summarize patterns of trait relationships. Modules, highly-correlated and semi-autonomous sets of traits, are observed at many levels of biological organization, from genes to colonies. The evolutionary significance of modularity is considerable, with potential effects including constraining the variation of individual traits, circumventing pleiotropy and canalization, and facilitating the transformation of functional structures. Despite these important consequences, there has been little empirical study of how modularity influences morphological evolution on a macroevolutionary scale. Here, we conduct the first morphometric analysis of modularity and disparity in two clades of placental mammals, Primates and Carnivora, and test if trait integration within modules constrains or facilitates morphological evolution.Principal Findings: We used both randomization methods and direct comparisons of landmark variance to compare disparity in the six cranial modules identified in previous studies. The cranial base, a highly-integrated module, showed significantly low disparity in Primates and low landmark variance in both Primates and Carnivora. The vault, zygomatic-pterygoid and orbit modules, characterized by low trait integration, displayed significantly high disparity within Carnivora. 14 of 24 results from analyses of disparity show no significant relationship between module integration and morphological disparity. Of the ten significant or marginally significant results, eight support the hypothesis that integration within modules constrains morphological evolution in the placental skull. Only the molar module, a highly-integrated and functionally important module, showed significantly high disparity in Carnivora, in support of the facilitation hypothesis.Conclusions: This analysis of within-module disparity suggested that strong integration of traits had little influence on morphological evolution over large time scales. However, where significant results were found, the primary effect of strong integration of traits was to constrain morphological variation. Thus, within Primates and Carnivora, there was some support for the hypothesis that integration of traits within cranial modules limits morphological evolution, presumably by limiting the variation of individual traits.

International audience; Background: Dental microwear analyses are commonly used to deduce the diet of extinct mammals. Conventional methods rely on the user identifying features within a 2D image. However, recent interdisciplinary research has lead to the development of an advanced methodology that is free of observer error, based on the automated quantification of 3D surfaces by combining confocal microscopy with scale-sensitive fractal analysis. This method has already proved to be very efficient in detecting dietary differences between species. Focusing on a finer, intra-specific scale of analysis, the aim of this study is to test this method's ability to track such differences between individuals from a single population. Methodology/Principal Findings: For the purposes of this study, the 3D molar microwear of 78 individuals from a well-known population of extant roe deer (Capreolus caprelous) is quantified. Multivariate statistical analyses indicate significant seasonal and sexual differences in individual dental microwear design. These are probably the consequence of seasonal variations in fruit, seed and leaf availability, as well as differences in feeding preference between males and females due to distinct energy requirements during periods of rutting, gestation or giving birth. Nevertheless, further investigations using two-block Partial Least-Squares analysis show no strong relationship between individual stomach contents and microwear texture. This is an expected result, assuming that stomach contents are composed of food items ingested during the last few hours whereas dental microwear texture records the physical properties of items eaten over periods of days or weeks. Conclusions/Significance: Microwear 3D scale-sensitive fractal analysis does detect differences in diet ranging from the inter-feeding styles scale to the intra-population between-season and between-sex scales. It is therefore a possible tool, to be used with caution, in the further exploration of the feeding biology and ecology of extinct mammals.

BackgroundDebates over the evolution of hominin bipedalism, a defining human characteristic, revolve around whether early bipeds walked more like humans, with energetically efficient extended hind limbs, or more like apes with flexed hind limbs. The 3.6 million year old hominin footprints at Laetoli, Tanzania represent the earliest direct evidence of hominin bipedalism. Determining the kinematics of Laetoli hominins will allow us to understand whether selection acted to decrease energy costs of bipedalism by 3.6 Ma.Methodology/principal findingsUsing an experimental design, we show that the Laetoli hominins walked with weight transfer most similar to the economical extended limb bipedalism of humans. Humans walked through a sand trackway using both extended limb bipedalism, and more flexed limb bipedalism. Footprint morphology from extended limb trials matches weight distribution patterns found in the Laetoli footprints.ConclusionsThese results provide us with the earliest direct evidence of kinematically human-like bipedalism currently known, and show that extended limb bipedalism evolved long before the appearance of the genus Homo. Since extended-limb bipedalism is more energetically economical than ape-like bipedalism, energy expenditure was likely an important selection pressure on hominin bipeds by 3.6 Ma.

Molecular and morphological evidence unite the hemichordates and echinoderms as the Ambulacraria, but their earliest history remains almost entirely conjectural. This is on account of the morphological disparity of the ambulacrarians and a paucity of obvious stem-groups. We describe here a new taxon Herpetogaster collinsi gen. et sp. nov. from the Burgess Shale (Middle Cambrian) Lagerstätte. This soft-bodied vermiform animal has a pair of elongate dendritic oral tentacles, a flexible stolon with an attachment disc, and a re-curved trunk with at least 13 segments that is directed dextrally. A differentiated but un-looped gut is enclosed in a sac suspended by mesenteries. It consists of a short pharynx, a conspicuous lenticular stomach, followed by a narrow intestine sub-equal in length. This new taxon, together with the Lower Cambrian Phlogites and more intriguingly the hitherto enigmatic discoidal eldoniids (Cambrian-Devonian), form a distinctive clade (herein the cambroernids). Although one hypothesis of their relationships would look to the lophotrochozoans (specifically the entoprocts), we suggest that the evidence is more consistent with their being primitive deuterostomes, with specific comparisons being made to the pterobranch hemichordates and pre-radial echinoderms. On this basis some of the earliest ambulacrarians are interpreted as soft-bodied animals with a muscular stalk, and possessing prominent tentacles.

Inferring the behavior and function of ancient organisms is hard. Some paleontologists would say that it cannot be done because such hypotheses can never be testable, whereas others would say that this is surely a prime task for paleontology—to seek to bring ancient organisms back to life. These issues have long troubled paleontologists. The founder of comparative anatomy, Georges Cuvier (1769–1832), insisted on the common pattern of the skeleton of living and fossil vertebrates and that anatomy could be reconstructed with confidence from incomplete fossil remains. Further, he argued that the skeleton of a living or extinct animal held unequivocal clues about function and behavior. Cuvier saw his mission to establish rules for comparative anatomy that would allow paleontologists to make certain statement with clarity and confidence [1], a key principle today, what one might call “evidence-based reconstruction” (for example, sharp teeth indicate a diet of meat rather than plants, or mammalian characters in the teeth indicate that the unknown animal was endothermic and nourished its young from mammary glands) as opposed to speculation (“this dinosaur was purple because I guess it was”).

BackgroundThe fossil record reveals surprising crocodile diversity in the Neogene of Africa, but relationships with their living relatives and the biogeographic origins of the modern African crocodylian fauna are poorly understood. A Plio-Pleistocene crocodile from Olduvai Gorge, Tanzania, represents a new extinct species and shows that high crocodylian diversity in Africa persisted after the Miocene. It had prominent triangular "horns" over the ears and a relatively deep snout, these resemble those of the recently extinct Malagasy crocodile Voay robustus, but the new species lacks features found among osteolaemines and shares derived similarities with living species of Crocodylus.Methodology/principal findingsThe holotype consists of a partial skull and skeleton and was collected on the surface between two tuffs dated to approximately 1.84 million years (Ma), in the same interval near the type localities for the hominids Homo habilis and Australopithecus boisei. It was compared with previously-collected material from Olduvai Gorge referable to the same species. Phylogenetic analysis places the new form within or adjacent to crown Crocodylus.Conclusions/significanceThe new crocodile species was the largest predator encountered by our ancestors at Olduvai Gorge, as indicated by hominid specimens preserving crocodile bite marks from these sites. The new species also reinforces the emerging view of high crocodylian diversity throughout the Neogene, and it represents one of the few extinct species referable to crown genus Crocodylus.

Biodiversity is in a crisis caused by multiple human impacts on the environment [1]–[3]. The immediate and critical tasks in addressing this crisis are to examine global biodiversity patterns and document changes through time and space in order to understand the factors contributing to loss of biodiversity [4]. Meeting this challenge has emerged as a global priority [5]–[7] that requires approaches to mobilize data across broad geographic and taxonomic ranges. The community of vertebrate natural history collections has begun to meet this challenge by establishing social and technological infrastructures that provide open access to species occurrence data through broad participation and funding from the US National Science Foundation (NSF) and the Global Biodiversity Information Facility (GBIF). One result is VertNet, a publicly accessible database of vertebrate biodiversity data from natural history collections around the world. VertNet currently consists of four existing global vertebrate networks: Mammal Networked Information System (MaNIS) (http://manisnet.org – mammals [8]); Ornithological Information System (ORNIS) (http://ornisnet.org – birds); HerpNET (http://herpnet.org – amphibians and reptiles); and FishNet 2 (http://www.fishnet2.net – fishes). These networks collectively mobilize over 52 million records from over 70 institutions, which represent about 70% of all the vertebrate species occurrence data that are accessible through GBIF. VertNet was created to develop the tools and infrastructure necessary to make the data in these distributed networks available in a standard format to maximize their potential for understanding and protecting biodiversity. GBIF and VertNet work synergistically to enhance biodiversity data mobilization efforts. GBIF has identified the important role that VertNet will play in its new emphasis on decentralization of services and applications [5]. In particular, VertNet provides important data maintenance services, including data cleaning and indexing, thus removing development and deployment burdens for many fundamental tasks from GBIF. Data from VertNet are currently accessed through the four networks by a broad audience at a high frequency of about 2.5 million records per week. The networks continue to grow, even those whose extramural funding have expired; MaNIS, for example, has grown from 17 contributing institutions under the original grant to 38 institutions, with a waiting list of 31 more that will be added to the network as time and resources permit. The same enthusiasm is mirrored across the other three vertebrate networks. This rapid growth demonstrates an important sociological shift from skepticism to enthusiasm for data sharing. Ironically, the success of these networks has become their biggest challenge, straining original architectures and demanding a scalable and more sustainable solution. Below, we provide perspectives on the sociological and technical developments that brought vertebrate biodiversity networks to this point and discuss solutions to the immediate and anticipated challenges.

BackgroundThe holotype of the theropod non-avian dinosaur Microraptor gui from the Early Cretaceous of China shows extensive preservation of feathers in a halo around the body and with flight feathers associated with both the fore and hindlimbs. It has been questioned as to whether or not the feathers did extend into the halo to reach the body, or had disassociated and moved before preservation. This taxon has important implications for the origin of flight in birds and the possibility of a four-winged gliding phase.Methodology/principal findingsExamination of the specimen under ultraviolet light reveals that these feathers actually reach the body of the animal and were not disassociated from the bones. Instead they may have been chemically altered by the body tissues of the animal meaning that they did not carbonise close into the animal or more likely were covered by other decaying tissue, though evidence of their presence remains.Conclusions/significanceThese UV images show that the feathers preserved on the slab are genuinely associated with the skeleton and that their arrangement and orientation is likely correct. The methods used here to reveal hidden features of the specimen may be applicable to other specimens from the fossil beds of Liaoning that produced Microraptor.

Background Abundant fossils from the Ediacaran and Cambrian showing cnidarian grade grossly suggest that cnidarian diversification occurred earlier than that of other eumetazoans. However, fossils of possible soft-bodied polyps are scanty and modern corals are dated back only to the Middle Triassic, although molecular phylogenetic results support the idea that anthozoans represent the first major branch of the Cnidaria. Because of difficulties in taxonomic assignments owing to imperfect preservation of fossil cnidarian candidates, little is known about forms ancestral to those of living groups. Methods and Findings We have analyzed the soft-bodied polypoid microfossils Eolympia pediculata gen. et sp. nov. from the lowest Cambrian Kuanchuanpu Formation in southern China by scanning electron microscopy and computer-aided microtomography after isolating fossils from sedimentary rocks by acetic acid maceration. The fossils, about a half mm in body size, are preserved with 18 mesenteries including directives bilaterally arranged, 18 tentacles and a stalk-like pedicle. The pedicle suggests a sexual life cycle, while asexual reproduction by transverse fission also is inferred by circumferential grooves on the body column. Conclusions The features found in the present fossils fall within the morphological spectrum of modern Hexacorallia excluding Ceriantharia, and thus Eolympia pediculata could be a stem member for this group. The fossils also demonstrate that basic features characterizing modern hexacorallians such as bilateral symmetry and the reproductive system have deep roots in the Early Cambrian., Funding was provided by the National Science Foundation of China (http://www.nsfc.gov.cn/) grants 40830208, 40602003, 50702005 to J. Han and D. G. Shu, and by MOST Special Fund from the State Key Laboratory of Continental Dynamics, Northwest University, China (http://sklcd.nwu.edu.cn/) to J. Han and D. G. Shu. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

BackgroundNeandertals and the Middle Paleolithic persisted in the Iberian Peninsula south of the Ebro drainage system for several millennia beyond their assimilation/replacement elsewhere in Europe. As only modern humans are associated with the later stages of the Aurignacian, the duration of this persistence pattern can be assessed via the dating of diagnostic occurrences of such stages.Methodology/principal findingsUsing AMS radiocarbon and advanced pretreatment techniques, we dated a set of stratigraphically associated faunal samples from an Aurignacian III-IV context excavated at the Portuguese cave site of Pego do Diabo. Our results establish a secure terminus ante quem of ca. 34,500 calendar years ago for the assimilation/replacement process in westernmost Eurasia. Combined with the chronology of the regional Late Mousterian and with less precise dating evidence for the Aurignacian II, they place the denouement of that process in the 37th millennium before present.Conclusions/significanceThese findings have implications for the understanding of the emergence of anatomical modernity in the Old World as a whole, support explanations of the archaic features of the Lagar Velho child's anatomy that invoke evolutionarily significant Neandertal/modern admixture at the time of contact, and counter suggestions that Neandertals could have survived in southwest Iberia until as late as the Last Glacial Maximum.

11 páginas, 5 figuras, 1 tabla., [Background]Evolutionary studies of insular biotas are based mainly on extant taxa, although such biotas represent artificial subsets of original faunas because of human-caused extinctions of indigenous species augmented by introduced exotic taxa. This makes it difficult to obtain a full understanding of the history of ecological interactions between extant sympatric species. Morphological bill variation of Fringilla coelebs and F. teydea (common and blue chaffinches) has been previously studied in the North Atlantic Macaronesian archipelagos. Character displacement between both species has been argued to explain bill sizes in sympatry. However, this explanation is incomplete, as similar patterns of bill size have been recorded in F. coelebs populations from islands with and without F. teydea., [Methodology/Principal Findings]The discovery of a new extinct species in Tenerife (Canary Islands), here named Carduelis aurelioi n. sp. (slender-billed greenfinch), provides the opportunity to study ancient ecological interactions among Macaronesian finches. To help understand the evolutionary histories of forest granivores in space and time, we have performed a multidisciplinary study combining: (1) morphological analyses and radiocarbon dating (11,460±60 yr BP) of the new taxon and, (2) molecular divergence among the extant finch species and populations in order to infer colonization times (1.99 and 1.09 My for F. teydea and F. coelebs respectively)., [Conclusion/Significance]C. aurelioi, F. coelebs and F. teydea co-habited in Tenerife for at least one million years. The unique anatomical trends of the new species, namely chaffinch-like beak and modified hind and forelimbs, reveal that there was a process of divergence of resource competition traits among the three sympatric finches. The results of our study, combined with the presence of more extinct greenfinches in other Macaronesian islands with significant variation in their beak sizes, suggests that the character displacement has influenced patterns of divergence in bill size and shape on other Macaronesian islands as well.

BACKGROUND: Tyrannosaurus rex was one of the largest terrestrial carnivores of all time, and consequently its ecology and diet have been the focus of much discussion. However, there is little direct evidence of diet or feeding habits in this species. METHODOLOGY/PRINCIPAL FINDINGS: Examination of museum collections has revealed four specimens of Tyrannosaurus rex that bear tooth marks made by large, carnivorous dinosaurs. Because Tyrannosaurus is the only large carnivore known from the Late Maastrichtian of western North America, we infer that Tyrannosaurus made these tooth marks. CONCLUSIONS/SIGNIFICANCE: The marks are interpreted as feeding traces and these fossils therefore record instances of cannibalism. Given that this behavior has a low preservation potential, cannibalism seems to have been a surprisingly common behavior in Tyrannosaurus, and this behavior may have been relatively common in carnivorous dinosaurs.

Altres ajuts: Generalitat de Catalunya 2006 FI 00065 i beca de viatge 2008 BE1 00370 Background: The morphology of human pollical distal phalanges (PDP) closely reflects the adaptation of human hands for refined precision grip with pad-to-pad contact. The presence of these precision grip-related traits in the PDP of fossil hominins has been related to human-like hand proportions (i.e. short hands with a long thumb) enabling the thumb and finger pads to contact. Although this has been traditionally linked to the appearance of stone tool-making, the alternative hypothesis of an earlier origin-related to the freeing of the hands thanks to the advent of terrestrial bipedalism-is also possible given the human-like intrinsic hand proportion found in australopiths. - Methodology/Principal Findings: We perform morphofunctional and morphometric (bivariate and multivariate) analyses of most available hominin pollical distal phalanges, including Orrorin, Australopithecus, Paranthropous and fossil Homo, in order to investigate their morphological affinities. Our results indicate that the thumb morphology of the early biped Orrorin is more human-like than that of australopiths, in spite of its ancient chronology (ca. 6 Ma). Moreover, Orrorin already displays typical human-like features related to precision grasping. - Conclusions: These results reinforce previous hypotheses relating the origin of refined manipulation of natural objects-not stone tool-making-with the relaxation of locomotor selection pressures on the forelimbs. This suggests that human hand length proportions are largely plesiomorphic, in the sense that they more closely resemble the relatively short-handed Miocene apes than the elongated hand pattern of extant hominoids. With the advent of terrestrial bipedalism, these hand proportions may have been co-opted by early hominins for enhanced manipulative capabilities that, in turn, would have been later co-opted for stone tool-making in the genus Homo, more encephalized than the previous australopiths. This hypothesis remains may be further tested by the finding of more complete hands of unequivocally biped early hominins.

This study compares three extant elephants species - forest, savanna, and Asian - to extinct mammoth and mastodon. Surprisingly, forest and savanna elephants in Africa today are as distinct from each other as mammoth and Asian elephants., To elucidate the history of living and extinct elephantids, we generated 39,763 bp of aligned nuclear DNA sequence across 375 loci for African savanna elephant, African forest elephant, Asian elephant, the extinct American mastodon, and the woolly mammoth. Our data establish that the Asian elephant is the closest living relative of the extinct mammoth in the nuclear genome, extending previous findings from mitochondrial DNA analyses. We also find that savanna and forest elephants, which some have argued are the same species, are as or more divergent in the nuclear genome as mammoths and Asian elephants, which are considered to be distinct genera, thus resolving a long-standing debate about the appropriate taxonomic classification of the African elephants. Finally, we document a much larger effective population size in forest elephants compared with the other elephantid taxa, likely reflecting species differences in ancient geographic structure and range and differences in life history traits such as variance in male reproductive success., Author Summary The living elephants are the last survivors of a once highly successful mammalian order, the Proboscidea, which includes extinct species such as the iconic woolly mammoth (Mammuthus primigenius) and the American mastodon (Mammut americanum). Despite numerous studies, the phylogenetic relationships of the modern elephants to the woolly mammoth, as well as the taxonomic status of the African elephants of the genus Loxodonta, remain controversial. This is in large part due to the fact that both the woolly mammoth and the American mastodon (the closest outgroup to elephants and mammoths available for genetic studies) are extinct, posing considerable technical hurdles for comparative genetic analysis. We have used a combination of modern DNA sequencing and targeted PCR amplification to obtain a large data set for comparing American mastodon, woolly mammoth, Asian elephant, African savanna elephant, and African forest elephant. We unequivocally establish that the Asian elephant is the sister species to the woolly mammoth. A surprising finding from our study is that the divergence of African savanna and forest elephants—which some have argued to be two populations of the same species—is about as ancient as the divergence of Asian elephants and mammoths. Given their ancient divergence, we conclude that African savanna and forest elephants should be classified as two distinct species.

BackgroundGlobal temperature increased by approximately half a degree (Celsius) within the last 150 years. Even this moderate warming had major impacts on Earth's ecological and biological systems, especially in the Arctic where the magnitude of abiotic changes even exceeds those in temperate and tropical biomes. Therefore, understanding the biological consequences of climate change on high latitudes is of critical importance for future conservation of the species living in this habitat. The past 25,000 years can be used as a model for such changes, as they were marked by prominent climatic changes that influenced geographical distribution, demographic history and pattern of genetic variation of many extant species. We sequenced ancient and modern DNA of the collared lemming (Dicrostonyx torquatus), which is a key species of the arctic biota, from a single site (Pymva Shor, Northern Pre Urals, Russia) to see if climate warming events after the Last Glacial Maximum had detectable effects on the genetic variation of this arctic rodent species, which is strongly associated with a cold and dry climate.ResultsUsing three dimensional network reconstructions we found a dramatic decline in genetic diversity following the LGM. Model-based approaches such as Approximate Bayesian Computation and Markov Chain Monte Carlo based Bayesian inference show that there is evidence for a population decline in the collared lemming following the LGM, with the population size dropping to a minimum during the Greenland Interstadial 1 (Bølling/Allerød) warming phase at 14.5 kyrs BP.ConclusionOur results show that previous climate warming events had a strong influence on genetic diversity and population size of collared lemmings. Due to its already severely compromised genetic diversity a similar population reduction as a result of the predicted future climate change could completely abolish the remaining genetic diversity in this population. Local population extinctions of collared lemmings would have severe effects on the arctic ecosystem, as collared lemmings are a key species in the trophic interactions and ecosystem processes in the Arctic.

Earth has experienced five major extinction events in the past 450 million years. Many scientists suggest we are now witnessing a sixth, driven by human impacts. However, it has been difficult to quantify the real extent of the current extinction episode, either for a given taxonomic group at the continental scale or for the worldwide biota, largely because comparisons of pre-anthropogenic and anthropogenic biodiversity baselines have been unavailable. Here, we compute those baselines for mammals of temperate North America, using a sampling-standardized rich fossil record to reconstruct species-area relationships for a series of time slices ranging from 30 million to 500 years ago. We show that shortly after humans first arrived in North America, mammalian diversity dropped to become at least 15%-42% too low compared to the "normal" diversity baseline that had existed for millions of years. While the Holocene reduction in North American mammal diversity has long been recognized qualitatively, our results provide a quantitative measure that clarifies how significant the diversity reduction actually was. If mass extinctions are defined as loss of at least 75% of species on a global scale, our data suggest that North American mammals had already progressed one-fifth to more than halfway (depending on biogeographic province) towards that benchmark, even before industrialized society began to affect them. Data currently are not available to make similar quantitative estimates for other continents, but qualitative declines in Holocene mammal diversity are also widely recognized in South America, Eurasia, and Australia. Extending our methodology to mammals in these areas, as well as to other taxa where possible, would provide a reasonable way to assess the magnitude of global extinction, the biodiversity impact of extinctions of currently threatened species, and the efficacy of conservation efforts into the future.

BackgroundOne of the great unresolved controversies in paleobiology is whether extinct dinosaurs were endothermic, ectothermic, or some combination thereof, and when endothermy first evolved in the lineage leading to birds. Although it is well established that high, sustained growth rates and, presumably, high activity levels are ancestral for dinosaurs and pterosaurs (clade Ornithodira), other independent lines of evidence for high metabolic rates, locomotor costs, or endothermy are needed. For example, some studies have suggested that, because large dinosaurs may have been homeothermic due to their size alone and could have had heat loss problems, ectothermy would be a more plausible metabolic strategy for such animals.Methodology/principal findingsHere we describe two new biomechanical approaches for reconstructing the metabolic rate of 14 extinct bipedal dinosauriforms during walking and running. These methods, well validated for extant animals, indicate that during walking and slow running the metabolic rate of at least the larger extinct dinosaurs exceeded the maximum aerobic capabilities of modern ectotherms, falling instead within the range of modern birds and mammals. Estimated metabolic rates for smaller dinosaurs are more ambiguous, but generally approach or exceed the ectotherm boundary.Conclusions/significanceOur results support the hypothesis that endothermy was widespread in at least larger non-avian dinosaurs. It was plausibly ancestral for all dinosauriforms (perhaps Ornithodira), but this is perhaps more strongly indicated by high growth rates than by locomotor costs. The polarity of the evolution of endothermy indicates that rapid growth, insulation, erect postures, and perhaps aerobic power predated advanced "avian" lung structure and high locomotor costs.

International audience; BACKGROUND: The fish order Cypriniformes is one of the most diverse ray-finned fish groups in the world with more than 3000 recognized species. Cypriniformes are characterized by a striking distribution of their dentition: namely the absence of oral teeth and presence of pharyngeal teeth on the last gill arch (fifth ceratobranchial). Despite this limited localisation, the diversity of tooth patterns in Cypriniformes is astonishing. Here we provide a further description of this diversity using X-ray microtomography and we map the resulting dental characters on a phylogenetic tree to explore evolutionary trends. RESULTS: We performed a pilot survey of dental formulae and individual tooth shapes in 34 adult species of Cypriniformes by X-ray microtomography (using either conventional X-ray machine, or synchrotron microtomography when necessary) or by dissecting. By mapping morphological results in a phylogenetic tree, it emerges that the two super-families Cobitoidea and Cyprinoidea have followed two distinct evolutionary pathways. Furthermore, our analysis supports the hypothesis of a three-row dentition as ancestral for Cyprinoidea and a general trend in tooth row reduction in most derived lineages. Yet, this general scheme must be considered with caution as several events of tooth row gain and loss have occurred during evolutionary history of Cyprinoidea. SIGNIFICANCE: Dentition diversity in Cypriniformes constitutes an excellent model to study the evolution of complex morphological structures. This morphological survey clearly advocates for extending the use of X-ray microtomography to study tooth morphology in Cypriniformes. Yet, our survey also underlines that improved knowledge of Cypriniformes life traits, such as feeding habits, is required as current knowledge is not sufficient to conclude on the link between diet and dental morphology.

Background Archaeopteryx is the oldest and most primitive known bird (Avialae). It is believed that the growth and energetic physiology of basalmost birds such as Archaeopteryx were inherited in their entirety from non-avialan dinosaurs. This hypothesis predicts that the long bones in these birds formed using rapidly growing, well-vascularized woven tissue typical of non-avialan dinosaurs. Methodology/Principal Findings We report that Archaeopteryx long bones are composed of nearly avascular parallel-fibered bone. This is among the slowest growing osseous tissues and is common in ectothermic reptiles. These findings dispute the hypothesis that non-avialan dinosaur growth and physiology were inherited in totality by the first birds. Examining these findings in a phylogenetic context required intensive sampling of outgroup dinosaurs and basalmost birds. Our results demonstrate the presence of a scale-dependent maniraptoran histological continuum that Archaeopteryx and other basalmost birds follow. Growth analysis for Archaeopteryx suggests that these animals showed exponential growth rates like non-avialan dinosaurs, three times slower than living precocial birds, but still within the lowermost range for all endothermic vertebrates. Conclusions/Significance The unexpected histology of Archaeopteryx and other basalmost birds is actually consistent with retention of the phylogenetically earlier paravian dinosaur condition when size is considered. The first birds were simply feathered dinosaurs with respect to growth and energetic physiology. The evolution of the novel pattern in modern forms occurred later in the group's history.

BackgroundDuring the Ordovician the global diversity increased dramatically at family, genus and species levels. Partially the diversification is explained by an increased nutrient, and phytoplankton availability in the open water. Cephalopods are among the top predators of today's open oceans. Their Ordovician occurrences, diversity evolution and abundance pattern potentially provides information on the evolution of the pelagic food chain.Methodology/principal findingsWe reconstructed the cephalopod departure from originally exclusively neritic habitats into the pelagic zone by the compilation of occurrence data in offshore paleoenvironments from the Paleobiology Database, and from own data, by evidence of the functional morphology, and the taphonomy of selected cephalopod faunas. The occurrence data show, that cephalopod associations in offshore depositional settings and black shales are characterized by a specific composition, often dominated by orthocerids and lituitids. The siphuncle and conch form of these cephalopods indicate a dominant lifestyle as pelagic, vertical migrants. The frequency distribution of conch sizes and the pattern of epibionts indicate an autochthonous origin of the majority of orthocerid and lituitid shells. The consistent concentration of these cephalopods in deep subtidal sediments, starting from the middle Tremadocian indicates the occupation of the pelagic zone early in the Early Ordovician and a subsequent diversification which peaked during the Darriwilian.Conclusions/significanceThe exploitation of the pelagic realm started synchronously in several independent invertebrate clades during the latest Cambrian to Middle Ordovician. The initial rise and diversification of pelagic cephalopods during the Early and Middle Ordovician indicates the establishment of a pelagic food chain sustainable enough for the development of a diverse fauna of large predators. The earliest pelagic cephalopods were slowly swimming vertical migrants. The appearance and early diversification of pelagic cephalopods is interpreted as a consequence of the increased food availability in the open water since the latest Cambrian.

BackgroundThe largest living lizard species, Varanus komodoensis Ouwens 1912, is vulnerable to extinction, being restricted to a few isolated islands in eastern Indonesia, between Java and Australia, where it is the dominant terrestrial carnivore. Understanding how large-bodied varanids responded to past environmental change underpins long-term management of V. komodoensis populations.Methodology/principal findingsWe reconstruct the palaeobiogeography of Neogene giant varanids and identify a new (unnamed) species from the island of Timor. Our data reject the long-held perception that V. komodoensis became a giant because of insular evolution or as a specialist hunter of pygmy Stegodon. Phyletic giantism, coupled with a westward dispersal from mainland Australia, provides the most parsimonious explanation for the palaeodistribution of V. komodoensis and the newly identified species of giant varanid from Timor. Pliocene giant varanid fossils from Australia are morphologically referable to V. komodoensis suggesting an ultimate origin for V. komodoensis on mainland Australia (>3.8 million years ago). Varanus komodoensis body size has remained stable over the last 900,000 years (ka) on Flores, a time marked by major faunal turnovers, extinction of the island's megafauna, the arrival of early hominids by 880 ka, co-existence with Homo floresiensis, and the arrival of modern humans by 10 ka. Within the last 2000 years their populations have contracted severely.Conclusions/significanceGiant varanids were once a ubiquitous part of Subcontinental Eurasian and Australasian faunas during the Neogene. Extinction played a pivotal role in the reduction of their ranges and diversity throughout the late Quaternary, leaving only V. komodoensis as an isolated long-term survivor. The events over the last two millennia now threaten its future survival.

BackgroundTyrannosaurus rex and other tyrannosaurid fossils often display multiple, smooth-edged full-thickness erosive lesions on the mandible, either unilaterally or bilaterally. The cause of these lesions in the Tyrannosaurus rex specimen FMNH PR2081 (known informally by the name 'Sue') has previously been attributed to actinomycosis, a bacterial bone infection, or bite wounds from other tyrannosaurids.Methodology/principal findingsWe conducted an extensive survey of tyrannosaurid specimens and identified ten individuals with full-thickness erosive lesions. These lesions were described, measured and photographed for comparison with one another. We also conducted an extensive survey of related archosaurs for similar lesions. We show here that these lesions are consistent with those caused by an avian parasitic infection called trichomonosis, which causes similar abnormalities on the mandible of modern birds, in particular raptors.Conclusions/significanceThis finding represents the first evidence for the ancient evolutionary origin of an avian transmissible disease in non-avian theropod dinosaurs. It also provides a valuable insight into the palaeobiology of these now extinct animals. Based on the frequency with which these lesions occur, we hypothesize that tyrannosaurids were commonly infected by a Trichomonas gallinae-like protozoan. For tyrannosaurid populations, the only non-avian dinosaur group that show trichomonosis-type lesions, it is likely that the disease became endemic and spread as a result of antagonistic intraspecific behavior, consumption of prey infected by a Trichomonas gallinae-like protozoan and possibly even cannibalism. The severity of trichomonosis-related lesions in specimens such as Tyrannosaurus rex FMNH PR2081 and Tyrannosaurus rex MOR 980, strongly suggests that these animals died as a direct result of this disease, mostly likely through starvation.

Background: The spread of agriculture into Europe and the ancestry of the first European farmers have been subjects of debate and controversy among geneticists, archaeologists, linguists and anthropologists. Debates have centred on the extent to which the transition was associated with the active migration of people as opposed to the diffusion of cultural practices. Recent studies have shown that patterns of human cranial shape variation can be employed as a reliable proxy for the neutral genetic relationships of human populations. Methodology/Principal Findings: Here, we employ measurements of Mesolithic (hunter-gatherers) and Neolithic (farmers) crania from Southwest Asia and Europe to test several alternative population dispersal and hunter-farmer gene-flow models. We base our alternative hypothetical models on a null evolutionary model of isolation-by-geographic and temporal distance. Partial Mantel tests were used to assess the congruence between craniometric distance and each of the geographic model matrices, while controlling for temporal distance. Our results demonstrate that the craniometric data fit a model of continuous dispersal of people (and their genes) from Southwest Asia to Europe significantly better than a null model of cultural diffusion. Conclusions/Significance: Therefore, this study does not support the assertion that farming in Europe solely involved the adoption of technologies and ideas from Southwest Asia by indigenous Mesolithic hunter-gatherers. Moreover, the results highlight the utility of craniometric data for assessing patterns of past population dispersal and gene flow.

BackgroundAustralia's dinosaurian fossil record is exceptionally poor compared to that of other similar-sized continents. Most taxa are known from fragmentary isolated remains with uncertain taxonomic and phylogenetic placement. A better understanding of the Australian dinosaurian record is crucial to understanding the global palaeobiogeography of dinosaurian groups, including groups previously considered to have had Gondwanan origins, such as the titanosaurs and carcharodontosaurids.Methodology/principal findingsWe describe three new dinosaurs from the late Early Cretaceous (latest Albian) Winton Formation of eastern Australia, including; Wintonotitan wattsi gen. et sp. nov., a basal titanosauriform; Diamantinasaurus matildae gen. et sp. nov., a derived lithostrotian titanosaur; and Australovenator wintonensis gen. et sp. nov., an allosauroid. We compare an isolated astragalus from the Early Cretaceous of southern Australia; formerly identified as Allosaurus sp., and conclude that it most-likely represents Australovenator sp.Conclusion/significanceThe occurrence of Australovenator from the Aptian to latest Albian confirms the presence in Australia of allosauroids basal to the Carcharodontosauridae. These new taxa, along with the fragmentary remains of other taxa, indicate a diverse Early Cretaceous sauropod and theropod fauna in Australia, including plesiomorphic forms (e.g. Wintonotitan and Australovenator) and more derived forms (e.g. Diamantinasaurus).

BACKGROUND: Major biological and cultural innovations in late Pliocene hominin evolution are frequently linked to the spread or fluctuating presence of C(4) grass in African ecosystems. Whereas the deep sea record of global climatic change provides indirect evidence for an increase in C(4) vegetation with a shift towards a cooler, drier and more variable global climatic regime beginning approximately 3 million years ago (Ma), evidence for grassland-dominated ecosystems in continental Africa and hominin activities within such ecosystems have been lacking. METHODOLOGY/PRINCIPAL FINDINGS: We report stable isotopic analyses of pedogenic carbonates and ungulate enamel, as well as faunal data from approximately 2.0 Ma archeological occurrences at Kanjera South, Kenya. These document repeated hominin activities within a grassland-dominated ecosystem. CONCLUSIONS/SIGNIFICANCE: These data demonstrate what hitherto had been speculated based on indirect evidence: that grassland-dominated ecosystems did in fact exist during the Plio-Pleistocene, and that early Homo was active in open settings. Comparison with other Oldowan occurrences indicates that by 2.0 Ma hominins, almost certainly of the genus Homo, used a broad spectrum of habitats in East Africa, from open grassland to riparian forest. This strongly contrasts with the habitat usage of Australopithecus, and may signal an important shift in hominin landscape usage.

BackgroundExtended neoteny and late stage allometric growth increase morphological disparity between growth stages in at least some dinosaurs. Coupled with relatively low dinosaur density in the Upper Cretaceous of North America, ontogenetic transformational representatives are often difficult to distinguish. For example, many hadrosaurids previously reported to represent relatively small lambeosaurine species were demonstrated to be juveniles of the larger taxa. Marginocephalians (pachycephalosaurids + ceratopsids) undergo comparable and extreme cranial morphological change during ontogeny.Methodology/principal findingsCranial histology, morphology and computer tomography reveal patterns of internal skull development that show the purported diagnostic characters for the pachycephalosaurids Dracorex hogwartsia and Stygimoloch spinifer are ontogenetically derived features. Coronal histological sections of the frontoparietal dome of an adult Pachycephalosaurus wyomingensis reveal a dense structure composed of metaplastic bone with a variety of extremely fibrous and acellular tissue. Coronal histological sections and computer tomography of a skull and frontoparietal dome of Stygimoloch spinifer reveal an open intrafrontal suture indicative of a subadult stage of development. These dinosaurs employed metaplasia to rapidly grow and change the size and shape of their horns, cranial ornaments and frontoparietal domes, resulting in extreme cranial alterations during late stages of growth. We propose that Dracorex hogwartsia, Stygimoloch spinifer and Pachycephalosaurus wyomingensis are the same taxon and represent an ontogenetic series united by shared morphology and increasing skull length.Conclusions/significanceDracorex hogwartsia (juvenile) and Stygimoloch spinifer (subadult) are reinterpreted as younger growth stages of Pachycephalosaurus wyomingensis (adult). This synonymy reduces the number of pachycephalosaurid taxa from the Upper Cretaceous of North America and demonstrates the importance of cranial ontogeny in evaluating dinosaur diversity and taxonomy. These growth stages reflect a continuum rather than specific developmental steps defined by "known" terminal morphologies.