Search

Eosuchus lerichei is a gavialoid crocodylian from late Paleocene marine deposits of northwestern Europe, known from a skull and lower jaws, as well as postcrania. Its sister taxon relationship with the approximately contemporaneous species Eosuchus minor from the east coast of the USA has been explained through transoceanic dispersal, indicating a capability for salt excretion that is absent in extant gavialoids. However, there is currently no anatomical evidence to support marine adaptation in extinct gavialoids. Furthermore, the placement of Eosuchus within Gavialoidea is labile, with some analyses supporting affinities with the Late Cretaceous to early Paleogene "thoracosaurs." Here we present novel data on the internal and external anatomy of the skull of E. lerichei that enables a revised diagnosis, with 6 autapormorphies identified for the genus and 10 features that enable differentiation of the species from Eosuchus minor. Our phylogenetic analyses recover Eosuchus as an early diverging gavialid gavialoid that is not part of the "thoracosaur" group. In addition to thickened semi-circular canal walls of the endosseous labyrinth and paratympanic sinus reduction, we identify potential osteological correlates for salt glands in the internal surface of the prefrontal and lacrimal bones of E. lerichei. These salt glands potentially provide anatomical evidence for the capability of transoceanic dispersal within Eosuchus, and we also identify them in the Late Cretaceous "thoracosaur" Portugalosuchus. Given that the earliest diverging and stratigraphically oldest gavialoids either have evidence for a nasal salt gland and/or have been recovered from marine deposits, this suggests the capacity for salt excretion might be ancestral for Gavialoidea. Mapping osteological and geological evidence for marine adaptation onto a phylogeny indicates that there was probably more than one independent loss/reduction in the capacity for salt excretion in gavialoids., (© 2024 The Author(s). The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Dinosaurs dominated Mesozoic terrestrial ecosystems for ∼160 million years, but their biogeographic origin remains poorly understood. The earliest unequivocal dinosaur fossils appear in the Carnian (∼230 Ma) of southern South America and Africa, leading most authors to propose southwestern Gondwana as the likely center of origin. However, the high taxonomic and morphological diversity of these earliest assemblages suggests a more ancient evolutionary history that is currently unsampled. Phylogenetic uncertainty at the base of Dinosauria, combined with the subsequent appearance of dinosaurs throughout Laurasia in their early evolutionary history, further complicates this picture. Here, we estimate the distribution of early dinosaurs and their archosaurian relatives under a phylogenetic maximum likelihood framework, testing alternative topological arrangements and incorporating potential abiotic barriers to dispersal into our biogeographic models. For the first time, we include spatiotemporal sampling heterogeneity in these models, which frequently supports a low-latitude Gondwanan origin for dinosaurs. These results are best supported when silesaurids are constrained as early-diverging ornithischians, which is likely because this topology accounts for the otherwise substantial ornithischian ghost lineage, explaining the group's absence from the fossil record prior to the Early Jurassic. Our results suggest that the archosaur radiation also took place within low-latitude Gondwana following the end-Permian extinction before lineages dispersed across Pangaea into ecologically and climatically distinct provinces during the Late Triassic. Mesozoic terrestrial vertebrates are under-sampled at low paleolatitudes, and our findings suggest that heterogeneous sampling has hitherto obscured the true paleobiogeographic origin of dinosaurs and their kin., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2025 The Authors. Published by Elsevier Inc. All rights reserved.)

Argochampsa krebsi is a gavialoid crocodylian from the early Paleogene of North Africa. Based on its recovered phylogenetic relationship with South American species, it has been inferred to have been capable of transoceanic dispersal, but potential anatomical correlates for a marine lifestyle have yet to be identified. Based on CT scans of a mostly complete and well-preserved skull, we reconstruct the endocranial anatomy of Argochampsa and compare it to that of other gavialoids. We demonstrate that Argochampsa possesses concave depressions on the internal surface of the prefrontals and lacrimals, which have been inferred to represent osteological correlates for salt glands in unequivocally marine metriorhynchoid thalattosuchian crocodyliforms. The presence of these salt glands suggests that Argochampsa likely frequented pelagic environments and provides additional support for the capability of transoceanic dispersal within Gavialoidea. We also newly interpret osteological correlates for salt glands in the Miocene north African gavialoid Sutekhsuchus dowsoni, providing further support that saltwater tolerance was widespread and possibly ancestral in Gavialoidea, given that they have been previously reported in the Late Cretaceous-early Paleogene species Eosuchus lerichei and Portugalosuchus azenhae. In addition to these gavialoids, as well as metriorhynchids, we also identify these osteological salt gland correlates in the Paleocene northwest African dyrosaurid Rhabdognathus aslerensis, which represents another crocodyliform lineage thought to be capable of transoceanic dispersal. Given that dyrosaurids, gavialoids, and metriorhynchoids are distantly related lineages, the evolution of salt glands is likely a convergent ecological adaptation to the occupation of pelagic environments. Nevertheless, we demonstrate limited evaluation of the presence of these osteological correlates across Crocodyliformes, including within most extant species, such that it remains possible that they are much more widespread., (© 2024 The Author(s). Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

Much of our view on Mesozoic dinosaur diversity is obscured by biases in the fossil record. In particular, spatiotemporal sampling heterogeneity affects identification of the timing and geographical location of radiations, the recognition of the latitudinal diversity gradient, as well as interpretation of purported extinctions, faunal turnovers and their drivers, including the Early Jurassic Jenkyns Event and across the Jurassic/Cretaceous boundary. The current distribution of sampling means it is impossible to robustly determine whether these 'events' were globally synchronous and geologically instantaneous or spatiotemporally staggered. Accounting for sampling heterogeneity is also paramount to reconciling notable differences in results based on sampling-standardized dinosaur species richness versus reconstructions of diversification rates, particularly with regards to the lead-up to the Cretaceous/Paleogene mass extinction. Incorporation of a greater proportion of stratigraphically well-resolved dinosaurs into analyses is also imperative and must include the substantial Mesozoic radiation of birds. Given the relative rarity of temporally successive, well-sampled spatial windows, it remains possible that dinosaur species richness and diversification rate showed little change after the clade's initial radiation until the Cretaceous/Paleogene boundary. However, better understanding of underlying sampling, combined with a holistic approach to reconstructing dinosaur diversity and diversification, is an important step in testing this hypothesis.

The Barremian-aged Wessex Formation of the Isle of Wight, UK, offers a globally significant glimpse into the sauropod dinosaur faunas of the early Cretaceous. These deposits have yielded specimens of several neosauropod lineages, such as rebbachisaurids, titanosauriforms (including some of the earliest titanosaur remains), and possible flagellicaudatans. Here, we report an undescribed sauropod partial hindlimb from the Wessex Formation (NHMUK PV R16500) and analyse its phylogenetic affinities. This hindlimb preserves the left tibia, astragalus and pes, lacking only a few phalanges. NHMUK PV R16500 can be diagnosed based on two autapomorphies: an unusually high distal end to midshaft transverse width ratio in metatarsals III and IV, and the presence of small bump-like projections located in the centre of the proximal articular surfaces of the unguals of pedal digits I and II. The phylogenetic affinities of NHMUK PV R16500 are uncertain: although our analyses recover it as an early-branching somphospondylan, a single character change moves it to close to Flagellicaudata when extended implied weighting is applied. The possibility of flagellicaudatan affinities for NHMUK PV R16500 implies a potential ghost lineage that survived the Jurassic/Cretaceous boundary; however, we present evidence that the somphospondylan position is more probable and should be preferred., Competing Interests: We declare we have no competing interests., (© 2024 The Authors.)

The bone-eating worm Osedax is a speciose and globally distributed clade, primarily found on whale carcasses in marine environments. The earliest fossil evidence for Osedax borings was previously described in plesiosaur and sea turtle bones from the mid-Cretaceous of the United Kingdom, representing the only unequivocal pre-Oligocene occurrences. Confirming through CT scanning, we present new evidence of Osedax borings in three plesiosaur specimens and, for the first time, identify borings in two mosasaur specimens. All specimens are from the Late Cretaceous: one from the Cenomanian of the United Kingdom, two from the Campanian of the southeastern United States, and two from the Maastrichtian of Belgium. This extends the geographic range of Osedax in the Cretaceous to both sides of the northern Atlantic Ocean. The bones contain five borehole morphotypes, potentially created by different species of Osedax , with the Cenomanian specimen containing three morphotypes within a single tooth. This combined evidence of heightened species diversity by the Cenomanian and broad geographic range by the Campanian potentially indicates an earlier origin and diversification for this clade than previously hypothesized. Preservational biases indicate that Osedax was probably even more widely distributed and speciose in the Cretaceous than apparent in the fossil record.

Skeletal remains of sauropod dinosaurs have been known from Australia for over 100 years. Unfortunately, the classification of the majority of these specimens to species level has historically been impeded by their incompleteness. This has begun to change in the last 15 years, primarily through the discovery and description of several partial skeletons from the Cenomanian-lower Turonian (lower Upper Cretaceous) Winton Formation in central Queensland, with four species erected to date: Australotitan cooperensis , Diamantinasaurus matildae , Savannasaurus elliottorum , and Wintonotitan wattsi . The first three of these appear to form a clade (Diamantinasauria) of early diverging titanosaurs (or close relatives of titanosaurs), whereas Wintonotitan wattsi is typically recovered as a distantly related non-titanosaurian somphospondylan. Through the use of 3D scanning, we digitised numerous specimens of Winton Formation sauropods, facilitating enhanced comparison between type and referred specimens, and heretofore undescribed specimens. We present new anatomical information on the holotype specimen of Diamantinasaurus matildae , and describe new remains pertaining to twelve sauropod individuals. Firsthand observations and digital analysis enabled previously proposed autapomorphic features of all four named Winton Formation sauropod species to be identified in the newly described specimens, with some specimens exhibiting putative autapomorphies of more than one species, prompting a reassessment of their taxonomic validity. Supported by a specimen-level phylogenetic analysis, we suggest that Australotitan cooperensis is probably a junior synonym of Diamantinasaurus matildae , but conservatively regard it herein as an indeterminate diamantinasaurian, meaning that the Winton Formation sauropod fauna now comprises three (rather than four) valid diamantinasaurian species: Diamantinasaurus matildae , Savannasaurus elliottorum , and Wintonotitan wattsi , with the latter robustly supported as a member of the clade for the first time. We refer some of the newly described specimens to these three species and provide revised diagnoses, with some previously proposed autapomorphies now regarded as diamantinasaurian synapomorphies. Our newly presented anatomical data and critical reappraisal of the Winton Formation sauropods facilitates a more comprehensive understanding of the mid-Cretaceous sauropod palaeobiota of central Queensland., Competing Interests: The authors declare that they have no competing interests., (© 2024 Beeston et al.)

Whereas living representatives of Pseudosuchia, crocodylians, number fewer than 30 species, more than 700 pseudosuchian species are known from their 250-million-year fossil record, displaying far greater ecomorphological diversity than their extant counterparts. With a new time-calibrated tree of >500 species, we use a phylogenetic framework to reveal that pseudosuchian evolutionary history and diversification dynamics were directly shaped by the interplay of abiotic and biotic processes over hundreds of millions of years, supported by information theory analyses. Speciation, but not extinction, is correlated with higher temperatures in terrestrial and marine lineages, with high sea level associated with heightened extinction in non-marine taxa. Low lineage diversity and increased speciation in non-marine species is consistent with opportunities for niche-filling, whereas increased competition may have led to elevated extinction rates. In marine lineages, competition via increased lineage diversity appears to have driven both speciation and extinction. Decoupling speciation and extinction, in combination with ecological partitioning, reveals a more complex picture of pseudosuchian evolution than previously understood. As the number of species threatened with extinction by anthropogenic climate change continues to rise, the fossil record provides a unique window into the drivers that led to clade success and those that may ultimately lead to extinction., (© 2023. The Author(s).)

The interrelationships of the extant crocodylians Gavialis gangeticus and Tomistoma schlegelii have been historically disputed. Whereas molecular analyses indicate a sister taxon relationship between these two gavialoid species, morphological datasets typically place Gavialis as the outgroup to all other extant crocodylians. Recent morphological-based phylogenetic analyses have begun to resolve this discrepancy, recovering Gavialis as the closest living relative of Tomistoma; however, several stratigraphically early fossil taxa are recovered as closer to Gavialis than Tomistoma, resulting in anomalously early divergence timings. As such, additional morphological data might be required to resolve these remaining discrepancies. 'Tomistoma' dowsoni is an extinct species of gavialoid from the Miocene of North Africa. Utilising CT scans of a near-complete, referred skull, we reconstruct the neuroanatomy and neurosensory apparatus of 'Tomistoma' dowsoni. Based on qualitative and quantitative morphometric comparisons with other crocodyliforms, the neuroanatomy of 'Tomistoma' dowsoni is characterised by an intermediate morphology between the two extant gavialoids, more closely resembling Gavialis. This mirrors the results of recent studies based on the external anatomy of these three species and other fossil gavialoids. Several neuroanatomical features of these species appear to reflect ecological and/or phylogenetic signals. For example, the 'simple' morphology of their neurosensory apparatus is broadly similar to that of other long and narrow-snouted (longirostrine), aquatic crocodyliforms. A dorsoventrally short, anteroposteriorly long endosseous labyrinth is also associated with longirostry. These features indicate that snout and skull morphology, which are themselves partly constrained by ecology, exert an influence on neuroanatomical morphology, as has also been recognised in birds and turtles. Conversely, the presence of a pterygoid bulla in Gavialis and several extinct gavialoids, and its absence in Tomistoma schlegelii, could be interpreted as a phylogenetic signal of crocodylians more closely related to Gavialis than to Tomistoma. Evaluation of additional fossil gavialoids will be needed to further test whether these and other neuroanatomical features primarily reflect a phylogenetic or ecological signal. By incorporating such previously inaccessible information of extinct and extant gavialoids into phylogenetic and macroecological studies, we can potentially further constrain the clade's interrelationships, as well as evaluate the timing and ecological association of the evolution of these neuroanatomical features. Finally, our study supports recent phylogenetic analyses that place 'Tomistoma' dowsoni as being phylogenetically closer to Gavialis gangeticus than to Tomistoma schlegelii, indicating the necessity of a taxonomic revision of this fossil species., (© 2023 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

Titanosaurian sauropod dinosaurs were diverse and abundant throughout the Cretaceous, with a global distribution. However, few titanosaurian taxa are represented by multiple skeletons, let alone skulls. Diamantinasaurus matildae , from the lower Upper Cretaceous Winton Formation of Queensland, Australia, was heretofore represented by three specimens, including one that preserves a braincase and several other cranial elements. Herein, we describe a fourth specimen of Diamantinasaurus matildae that preserves a more complete skull-including numerous cranial elements not previously known for this taxon-as well as a partial postcranial skeleton. The skull of Diamantinasaurus matildae shows many similarities to that of the coeval Sarmientosaurus musacchioi from Argentina (e.g. quadratojugal with posterior tongue-like process; braincase with more than one ossified exit for cranial nerve V; compressed-cone-chisel-like teeth), providing further support for the inclusion of both taxa within the clade Diamantinasauria. The replacement teeth within the premaxilla of the new specimen are morphologically congruent with teeth previously attributed to Diamantinasaurus matildae , and Diamantinasauria more broadly, corroborating those referrals. Plesiomorphic characters of the new specimen include a sacrum comprising five vertebrae (also newly demonstrated in the holotype of Diamantinasaurus matildae ), rather than the six or more that typify other titanosaurs. However, we demonstrate that there have been a number of independent acquisitions of a six-vertebrae sacrum among Somphospondyli and/or that there have been numerous reversals to a five-vertebrae sacrum, suggesting that sacral count is relatively plastic. Other newly identified plesiomorphic features include: the overall skull shape, which is more similar to brachiosaurids than 'derived' titanosaurs; anterior caudal centra that are amphicoelous, rather than procoelous; and a pedal phalangeal formula estimated as 2-2-3-2-0. These features are consistent with either an early-branching position within Titanosauria, or a position just outside the titanosaurian radiation, for Diamantinasauria, as indicated by alternative character weighting approaches applied in our phylogenetic analyses, and help to shed light on the early assembly of titanosaurian anatomy that has until now been obscured by a poor fossil record., Competing Interests: We declare we have no competing interests., (© 2023 The Authors.)

The numerous explanations for why Earth's biodiversity is concentrated at low latitudes fail to explain variation in the strength and even direction of the gradient through deep time. Consequently, we do not know if today's gradient is representative of what might be expected on other planets or is merely an idiosyncrasy of Earth's history. We propose a hierarchy of factors driving the latitudinal distribution of diversity: (i) over geologically long time spans, diversity is largely predicted by climate; (ii) when climatic gradients are shallow, diversity tracks habitat area; and (iii) historical contingencies linked to niche conservatism have geologically short-term, transient influence at most. Thus, latitudinal diversity gradients, although variable in strength and direction, are largely predictable on our planet and possibly others., Competing Interests: Declaration of interests The authors declare no conflicts of interest., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

It has long been debated why groups such as non-avian dinosaurs became extinct whereas mammals and other lineages survived the Cretaceous/Paleogene mass extinction 66 million years ago. We used Markov networks, ecological niche partitioning, and Earth System models to reconstruct North American food webs and simulate ecospace occupancy before and after the extinction event. We find a shift in latest Cretaceous dinosaur faunas, as medium-sized species counterbalanced a loss of megaherbivores, but dinosaur niches were otherwise stable and static, potentially contributing to their demise. Smaller vertebrates, including mammals, followed a consistent trajectory of increasing trophic impact and relaxation of niche limits beginning in the latest Cretaceous and continuing after the mass extinction. Mammals did not simply proliferate after the extinction event; rather, their earlier ecological diversification might have helped them survive.

The disjunct geographical range of many lineages of caudates points to a complex evolutionary and biogeographic history that cannot be disentangled by only considering the present-day distribution of salamander biodiversity. Here, we provide a critical reappraisal of the published fossil record of caudates from the Palearctic and quantitatively evaluate the quality of the group's fossil record. Stem-Urodela and Karauridae were widespread in the Palearctic in the Middle Jurassic, suggesting an earlier, unsampled diversification for this group. Cryptobranchidae reached Europe no later than the Oligocene, but this clade was subsequently extirpated from this continent, as well as from western and central Asia. The relatively recent appearance of hynobiids in the fossil record (Early Miocene) is most likely an artefact of a taphonomic bias against the preservation of high-mountain, stream-type environments which early members likely inhabited. Salamandroids first appear in Europe, expanding into Asia by the Miocene. The apparently enigmatic and disjunct distribution of extant caudate lineages is therefore explained by a wider past geographical range, as testified by the fossil record, which was fragmented during the late Cenozoic by a combination of tectonic (i.e. the uplift of the Tibetan Plateau) and climatic drivers, resulting in regional extirpations., (© 2022 The Authors.)

The Upper Cretaceous Winton Formation of Queensland, Australia, has produced several partial sauropod skeletons, but cranial remains-including teeth-remain rare. Herein, we present the first description of sauropod teeth from this formation, based on specimens from three separate sites. An isolated tooth and a dentary fragment from the Diamantinasaurus matildae type locality are considered to be referable to that titanosaurian taxon. A single tooth from the D. matildae referred specimen site is similarly regarded as being part of that individual. Seventeen teeth from a new site that are morphologically uniform, and similar to the teeth from the two Diamantinasaurus sites, are assigned to Diamantinasauria. All sauropod teeth recovered from the Winton Formation to date are compressed-cone-chisel-shaped, have low slenderness index values (2.00-2.88), are lingually curved at their apices, mesiodistally convex on their lingual surfaces, and lack prominent carinae and denticles. They are markedly different from the chisel-like teeth of derived titanosaurs, more closely resembling the teeth of early branching members of the titanosauriform radiation. This provides further support for a 'basal' titanosaurian position for Diamantinasauria. Scanning electron microscope microwear analysis of the wear facets of several teeth reveals more scratches than pits, implying that diamantinasaurians were mid-height (1-10 m) feeders. With a view to assessing the spatio-temporal distribution of sauropod tooth morphotypes before and after deposition of the Winton Formation, we provide a comprehensive continent-by-continent review of the early titanosauriform global record (Early to early Late Cretaceous). This indicates that throughout the Early-early Late Cretaceous, sauropod faunas transitioned from being quite diverse at higher phylogenetic levels and encompassing a range of tooth morphologies at the start of the Berriasian, to faunas comprising solely titanosaurs with limited dental variability by the end-Turonian. Furthermore, this review highlights the different ways in which this transition unfolded on each continent, including the earliest records of titanosaurs with narrow-crowned teeth on each continent., (© 2022 The Authors.)

Today, warm-water coral reefs are limited to tropical-to-subtropical latitudes. These diverse ecosystems extended further poleward in the geological past, but the mechanisms driving these past distributions remain uncertain. Here, we test the role of climate and palaeogeography in shaping the distribution of coral reefs over geological timescales. To do so, we combine habitat suitability modelling, Earth System modelling and the ~247-million-year geological record of scleractinian coral reefs. A broader latitudinal distribution of climatically suitable habitat persisted throughout much of the Mesozoic-early Paleogene due to an expanded tropical belt and more equable distribution of shallow marine substrate. The earliest Cretaceous might be an exception, with reduced shallow marine substrate during a 'cold-snap' interval. Climatically suitable habitat area became increasingly skewed towards the tropics from the late Paleogene, likely steepening the latitudinal biodiversity gradient of reef-associated taxa. This was driven by global cooling and increases in tropical shallow marine substrate resulting from the tectonic evolution of the Indo-Australian Archipelago. Although our results suggest global warming might permit long-term poleward range expansions, coral reef ecosystems are unlikely to keep pace with the rapid rate of anthropogenic climate change., (© 2022. The Author(s).)

Dinosaurs dominated Mesozoic terrestrial ecosystems globally. However, whereas a pole-to-pole geographic distribution characterized ornithischians and theropods, sauropods were restricted to lower latitudes. Here, we evaluate the role of climate in shaping these biogeographic patterns through the Jurassic-Cretaceous (201-66 mya), combining dinosaur fossil occurrences, past climate data from Earth System models, and habitat suitability modeling. Results show that, uniquely among dinosaurs, sauropods occupied climatic niches characterized by high temperatures and strongly bounded by minimum cold temperatures. This constrained the distribution and dispersal pathways of sauropods to tropical areas, excluding them from latitudinal extremes, especially in the Northern Hemisphere. The greater availability of suitable habitat in the southern continents, particularly in the Late Cretaceous, might be key to explaining the high diversity of sauropods there, relative to northern landmasses. Given that ornithischians and theropods show a flattened or bimodal latitudinal biodiversity gradient, with peaks at higher latitudes, the closer correspondence of sauropods to a subtropical concentration could hint at fundamental thermophysiological differences to the other two clades., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Notosuchians are an extinct clade of terrestrial crocodyliforms with a particularly rich record in the late Early to Late Cretaceous (approx. 130-66 Ma) of Gondwana. Although much of this diversity comes from South America, Africa and Indo-Madagascar have also yielded numerous notosuchian remains. Three notosuchian species are currently recognized from the early Late Cretaceous (approx. 100 Ma) Kem Kem Group of Morocco, including the peirosaurid Hamadasuchus rebouli . Here, we describe two new specimens that demonstrate the presence of at least a fourth notosuchian species in this fauna. Antaeusuchus taouzensis n. gen. n. sp. is incorporated into one of the largest notosuchian-focused character-taxon matrices yet to be compiled, comprising 443 characters scored for 63 notosuchian species, with an increased sampling of African and peirosaurid species. Parsimony analyses run under equal and extended implied weighting consistently recover Antaeusuchus as a peirosaurid notosuchian, supported by the presence of two distinct waves on the dorsal dentary surface, a surangular which laterally overlaps the dentary above the mandibular fenestra, and a relatively broad mandibular symphysis. Within Peirosauridae, Antaeusuchus is recovered as the sister taxon of Hamadasuchus . However, it differs from Hamadasuchus with respect to several features, including the ornamentation of the lateral surface of the mandible, the angle of divergence of the mandibular rami, the texture of tooth enamel and the shape of the teeth, supporting their generic distinction. We present a critical reappraisal of the non-South American Gondwanan notosuchian record, which spans the Middle Jurassic-late Eocene. This review, as well as our phylogenetic analyses, indicate the existence of at least three approximately contemporaneous peirosaurid lineages within the Kem Kem Group, alongside other notosuchians, and support the peirosaurid affinities of the 'trematochampsid' Miadanasuchus oblita from the Maastrichtian of Madagascar. Furthermore, the Cretaceous record demonstrates the presence of multiple lineages of approximately contemporaneous notosuchians in several African and Madagascan faunas, and supports previous suggestions regarding an undocumented pre-Aptian radiation of Notosuchia. By contrast, the post-Cretaceous record is depauperate, comprising rare occurrences of sebecosuchians in north Africa prior to their extirpation., (© 2021 The Authors.)

First appearing in the latest Cretaceous, Crocodylia is a clade of semi-aquatic, predatory reptiles, defined by the last common ancestor of extant alligators, caimans, crocodiles, and gharials. Despite large strides in resolving crocodylian interrelationships over the last three decades, several outstanding problems persist in crocodylian systematics. Most notably, there has been persistent discordance between morphological and molecular datasets surrounding the affinities of the extant gharials, Gavialis gangeticus and Tomistoma schlegelii . Whereas molecular data consistently support a sister taxon relationship, in which they are more closely related to crocodylids than to alligatorids, morphological data indicate that Gavialis is the sister taxon to all other extant crocodylians. Here we present a new morphological dataset for Crocodylia based on a critical reappraisal of published crocodylian character data matrices and extensive firsthand observations of a global sample of crocodylians. This comprises the most taxonomically comprehensive crocodylian dataset to date (144 OTUs scored for 330 characters) and includes a new, illustrated character list with modifications to the construction and scoring of characters, and 46 novel characters. Under a maximum parsimony framework, our analyses robustly recover Gavialis as more closely related to Tomistoma than to other extant crocodylians for the first time based on morphology alone. This result is recovered regardless of the weighting strategy and treatment of quantitative characters. However, analyses using continuous characters and extended implied weighting (with high k -values) produced the most resolved, well-supported, and stratigraphically congruent topologies overall. Resolution of the gharial problem reveals that: (1) several gavialoids lack plesiomorphic features that formerly drew them towards the stem of Crocodylia; and (2) more widespread similarities occur between species traditionally divided into tomistomines and gavialoids, with these interpreted here as homology rather than homoplasy. There remains significant temporal incongruence regarding the inferred divergence timing of the extant gharials, indicating that several putative gavialids ('thoracosaurs') are incorrectly placed and require future re-appraisal. New alligatoroid interrelationships include: (1) support for a North American origin of Caimaninae in the latest Cretaceous; (2) the recovery of the early Paleogene South American taxon Eocaiman as a 'basal' alligatoroid; and (3) the paraphyly of the Cenozoic European taxon Diplocynodon . Among crocodyloids, notable results include modifications to the taxonomic content of Mekosuchinae, including biogeographic affinities of this clade with latest Cretaceous-early Paleogene Asian crocodyloids. In light of our new results, we provide a comprehensive review of the evolutionary and biogeographic history of Crocodylia, which included multiple instances of transoceanic and continental dispersal., Competing Interests: The authors declare that they have no competing interests., (© 2021 Rio and Mannion.)

It has often been suggested that the productivity of an ecosystem affects the number of species that it can support. Despite decades of study, the nature, extent, and underlying mechanisms of this relationship are unclear. One suggested mechanism is the "more individuals" hypothesis (MIH). This proposes that productivity controls the number of individuals in the ecosystem, and that more individuals can be divided into a greater number of species before their population size is sufficiently small for each to be at substantial risk of extinction. Here, we test this hypothesis using REvoSim: an individual-based eco-evolutionary system that simulates the evolution and speciation of populations over geological time, allowing phenomena occurring over timescales that cannot be easily observed in the real world to be evaluated. The individual-based nature of this system allows us to remove assumptions about the nature of speciation and extinction that previous models have had to make. Many of the predictions of the MIH are supported in our simulations: Rare species are more likely to undergo extinction than common species, and species richness scales with productivity. However, we also find support for relationships that contradict the predictions of the strict MIH: species population size scales with productivity, and species extinction risk is better predicted by relative than absolute species population size, apparently due to increased competition when total community abundance is higher. Furthermore, we show that the scaling of species richness with productivity depends upon the ability of species to partition niche space. Consequently, we suggest that the MIH is applicable only to ecosystems in which niche partitioning has not been halted by species saturation. Some hypotheses regarding patterns of biodiversity implicitly or explicitly overlook niche theory in favor of neutral explanations, as has historically been the case with the MIH. Our simulations demonstrate that niche theory exerts a control on the applicability of the MIH and thus needs to be accounted for in macroecology., Competing Interests: The authors declare no conflict of interest., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Sauropod dinosaurs were an abundant and diverse component of the Upper Jurassic Morrison Formation of the USA, with 24 currently recognized species. However, some authors consider this high diversity to have been ecologically unviable and the validity of some species has been questioned, with suggestions that they represent growth series (ontogimorphs) of other species. Under this scenario, high sauropod diversity in the Late Jurassic of North America is greatly overestimated. One putative ontogimorph is the enigmatic diplodocoid Amphicoelias altus , which has been suggested to be synonymous with Diplodocus . Given that Amphicoelias was named first, it has priority and thus Diplodocus would become its junior synonym. Here, we provide a detailed re-description of A. altus in which we restrict it to the holotype individual and support its validity, based on three autapomorphies. Constraint analyses demonstrate that its phylogenetic position within Diplodocoidea is labile, but it seems unlikely that Amphicoelias is synonymous with Diplodocus . As such, our re-evaluation also leads us to retain Diplodocus as a distinct genus. There is no evidence to support the view that any of the currently recognized Morrison sauropod species are ontogimorphs. Available data indicate that sauropod anatomy did not dramatically alter once individuals approached maturity. Furthermore, subadult sauropod individuals are not prone to stemward slippage in phylogenetic analyses, casting doubt on the possibility that their taxonomic affinities are substantially misinterpreted. An anatomical feature can have both an ontogenetic and phylogenetic signature, but the former does not outweigh the latter when other characters overwhelmingly support the affinities of a taxon. Many Morrison Formation sauropods were spatio-temporally and/or ecologically separated from one another. Combined with the biases that cloud our reading of the fossil record, we contend that the number of sauropod dinosaur species in the Morrison Formation is currently likely to be underestimated, not overestimated., (© 2021 The Authors.)

There is significant geographic variation in species richness. However, the nature of the underlying relationships, such as that between species richness and environmental stability, remains unclear. The stability-time hypothesis suggests that environmental instability reduces species richness by suppressing speciation and increasing extinction risk. By contrast, the patch-mosaic hypothesis suggests that small-scale environmental instability can increase species richness by providing a steady supply of non-equilibrium environments. Although these hypotheses are often applied to different time scales, their core mechanisms are in conflict. Reconciling these apparently competing hypotheses is key to understanding how environmental conditions shape the distribution of biodiversity. Here, we use REvoSim, an individual-based, eco-evolutionary system, to model the evolution of sessile organisms in environments with varying magnitudes and scales of environmental instability. We demonstrate that when environments have substantial permanent heterogeneity, a high level of localized environmental instability reduces biodiversity, whereas in environments lacking permanent heterogeneity, high levels of localized instability increase biodiversity. By contrast, broad-scale environmental instability, acting on the same time scale, invariably reduces biodiversity. Our results provide a new view of the biodiversity-disturbance relationship that reconciles contrasting hypotheses within a single model and implies constraints on the environmental conditions under which those hypotheses apply. These constraints can inform attempts to conserve adaptive potential in different environments during the current biodiversity crisis.

The latitudinal biodiversity gradient (LBG), in which species richness decreases from tropical to polar regions, is a pervasive pattern of the modern biosphere. Although the distribution of fossil occurrences suggests this pattern has varied through deep time, the recognition of palaeobiogeographic patterns is hampered by geological and anthropogenic biases. In particular, spatial sampling heterogeneity has the capacity to impact upon the reconstruction of deep time LBGs. Here we use a simulation framework to test the detectability of three different types of LBG (flat, unimodal and bimodal) over the last 300 Myr. We show that heterogeneity in spatial sampling significantly impacts upon the detectability of genuine LBGs, with known biodiversity patterns regularly obscured after applying the spatial sampling window of fossil collections. Sampling-standardization aids the reconstruction of relative biodiversity gradients, but cannot account for artefactual absences introduced by geological and anthropogenic biases. Therefore, we argue that some previous studies might have failed to recover the 'true' LBG type owing to incomplete and heterogeneous sampling, particularly between 200 and 20 Ma. Furthermore, these issues also have the potential to bias global estimates of past biodiversity, as well as inhibit the recognition of extinction and radiation events.

Competing Interests: The author declares no competing interest.

The Cretaceous/Paleogene mass extinction, 66 Ma, included the demise of non-avian dinosaurs. Intense debate has focused on the relative roles of Deccan volcanism and the Chicxulub asteroid impact as kill mechanisms for this event. Here, we combine fossil-occurrence data with paleoclimate and habitat suitability models to evaluate dinosaur habitability in the wake of various asteroid impact and Deccan volcanism scenarios. Asteroid impact models generate a prolonged cold winter that suppresses potential global dinosaur habitats. Conversely, long-term forcing from Deccan volcanism (carbon dioxide [CO 2 ]-induced warming) leads to increased habitat suitability. Short-term (aerosol cooling) volcanism still allows equatorial habitability. These results support the asteroid impact as the main driver of the non-avian dinosaur extinction. By contrast, induced warming from volcanism mitigated the most extreme effects of asteroid impact, potentially reducing the extinction severity., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

There is no consensus about how terrestrial biodiversity was assembled through deep time, and in particular whether it has risen exponentially over the Phanerozoic. Using a database of 60 859 fossil occurrences, we show that the spatial extent of the worldwide terrestrial tetrapod fossil record itself expands exponentially through the Phanerozoic. Changes in spatial sampling explain up to 67% of the change in known fossil species counts, and these changes are decoupled from variation in habitable land area that existed through time. Spatial sampling therefore represents a real and profound sampling bias that cannot be explained as redundancy. To address this bias, we estimate terrestrial tetrapod diversity for palaeogeographical regions of approximately equal size. We find that regional-scale diversity was constrained over timespans of tens to hundreds of millions of years, and similar patterns are recovered for major subgroups, such as dinosaurs, mammals and squamates. Although the Cretaceous/Palaeogene mass extinction catalysed an abrupt two- to three-fold increase in regional diversity 66 million years ago, no further increases occurred, and recent levels of regional diversity do not exceed those of the Palaeogene. These results parallel those recovered in analyses of local community-level richness. Taken together, our findings strongly contradict past studies that suggested unbounded diversity increases at local and regional scales over the last 100 million years.

Titanosaurs were a globally distributed clade of Cretaceous sauropods. Historically regarded as a primarily Gondwanan radiation, there is a growing number of Eurasian taxa, with several putative titanosaurs contemporaneous with, or even pre-dating, the oldest known Southern Hemisphere remains. The early Late Cretaceous Jinhua Formation, in Zhejiang Province, China, has yielded two putative titanosaurs, Jiangshanosaurus lixianensis and Dongyangosaurus sinensis . Here, we provide a detailed re-description and diagnosis of Jiangshanosaurus , as well as new anatomical information on Dongyangosaurus . Previously, a 'derived' titanosaurian placement for Jiangshanosaurus was primarily based on the presence of procoelous anterior caudal centra. We show that this taxon had amphicoelous anterior-middle caudal centra. Its only titanosaurian synapomorphy is that the dorsal margins of the scapula and coracoid are approximately level with one another. Dongyangosaurus can clearly be differentiated from Jiangshanosaurus , and displays features that indicate a closer relationship to the titanosaur radiation. Revised scores for both taxa are incorporated into an expanded phylogenetic data matrix, comprising 124 taxa scored for 548 characters. Under equal weights parsimony, Jiangshanosaurus is recovered as a member of the non-titanosaurian East Asian somphospondylan clade Euhelopodidae, and Dongyangosaurus lies just outside of Titanosauria. However, when extended implied weighting is applied, both taxa are placed within Titanosauria. Most other 'middle' Cretaceous East Asian sauropods are probably non-titanosaurian somphospondylans, but at least Xianshanosaurus appears to belong to the titanosaur radiation. Our analyses also recover the Early Cretaceous European sauropod Normanniasaurus genceyi as a 'derived' titanosaur, clustering with Gondwanan taxa. These results provide further support for a widespread diversification of titanosaurs by at least the Early Cretaceous., Competing Interests: We declare we have no competing interests., (© 2019 The Authors.)

Reef corals are currently undergoing climatically driven poleward range expansions, with some evidence for equatorial range retractions. Predicting their response to future climate scenarios is critical to their conservation, but ecological models are based only on short-term observations. The fossil record provides the only empirical evidence for the long-term response of organisms under perturbed climate states. The palaeontological record from the Last Interglacial (LIG; 125 000 years ago), a time of global warming, suggests that reef corals experienced poleward range shifts and an equatorial decline relative to their modern distribution. However, this record is spatio-temporally biased, and existing methods cannot account for data absence. Here, we use ecological niche modelling to estimate reef corals' realized niche and LIG distribution, based on modern and fossil occurrences. We then make inferences about modelled habitability under two future climate change scenarios (RCP4.5 and RCP8.5). Reef coral ranges during the LIG were comparable to the present, with no prominent equatorial decrease in habitability. Reef corals are likely to experience poleward range expansion and large equatorial declines under RCP4.5 and RCP8.5. However, this range expansion is probably optimistic in the face of anthropogenic climate change. Incorporation of fossil data in niche models improves forecasts of biodiversity responses under global climatic change., Competing Interests: We declare we have no competing interests.

We examine how the history of Phanerozoic marine biodiversity relates to environmental change. Our focus is on North America, which has a relatively densely sampled history. By transforming time series into the time-frequency domain using wavelets, histories of biodiversity are shown to be similar to sea level, temperature and oceanic chemistry at multiple timescales. Fluctuations in sea level play an important role in driving Phanerozoic biodiversity at timescales >50 Myr, and during finite intervals at shorter periods. Subsampled and transformed marine genera time series reinforce the idea that Permian-Triassic, Triassic-Jurassic, and Cretaceous-Paleogene mass extinctions were geologically rapid, whereas the Ordovician-Silurian and Late Devonian 'events' were longer lived. High cross wavelet power indicates that biodiversity is most similar to environmental variables (sea level, plate fragmentation, δ 18 O, δ 13 C, δ 34 S and 87 Sr/ 86 Sr) at periods >200 Myr, when they are broadly in phase (i.e. no time lag). They are also similar at shorter periods for finite durations of time (e.g. during some mass extinctions). These results suggest that long timescale processes (e.g. plate kinematics) are the primary drivers of biodiversity, whilst processes with significant variability at shorter periods (e.g. glacio-eustasy, continental uplift and erosion, volcanism, asteroid impact) play a moderating role. Wavelet transforms are a useful approach for isolating information about times and frequencies of biological activity and commonalities with environmental variables.

The fossil record provides one of the strongest tests of the hypothesis that diversity within local communities is constrained over geological timescales. Constraints to diversity are particularly controversial in modern terrestrial ecosystems, yet long-term patterns are poorly understood. Here we document patterns of local richness in Phanerozoic terrestrial tetrapods using a global data set comprising 145,332 taxon occurrences from 27,531 collections. We show that the local richness of non-flying terrestrial tetrapods has risen asymptotically since their initial colonization of land, increasing at most threefold over the last 300 million years. Statistical comparisons support phase-shift models, with most increases in local richness occurring: (1) during the colonization of land by vertebrates, concluding by the late Carboniferous; and (2) across the Cretaceous/Paleogene boundary. Individual groups, such as mammals, lepidosaurs and dinosaurs also experienced early increases followed by periods of stasis often lasting tens of millions of years. Mammal local richness abruptly tripled across the Cretaceous/Paleogene boundary, but did not increase over the next 66 million years. These patterns are consistent with the hypothesis that diversity is constrained at the local-community scale.

In the lead-up to the Cretaceous/Paleogene mass extinction, dinosaur diversity is argued to have been either in long-term decline, or thriving until their sudden demise. The latest Cretaceous (Campanian-Maastrichtian [83-66 Ma]) of North America provides the best record to address this debate, but even here diversity reconstructions are biased by uneven sampling. Here we combine fossil occurrences with climatic and environmental modelling to quantify latest Cretaceous North American dinosaur habitat. Ecological niche modelling shows a Campanian-to-Maastrichtian habitability decrease in areas with present-day rock-outcrop. However, a continent-wide projection demonstrates habitat stability, or even a Campanian-to-Maastrichtian increase, that is not preserved. This reduction of the spatial sampling window resulted from formation of the proto-Rocky Mountains and sea-level regression. We suggest that Maastrichtian North American dinosaur diversity is therefore likely to be underestimated, with the apparent decline a product of sampling bias, and not due to a climatically-driven decrease in habitability as previously hypothesised.

The Jurassic/Cretaceous (J/K) boundary, 145 million years ago, has long been recognised as an extinction event or faunal turnover for sauropod dinosaurs, with many 'basal' lineages disappearing. However, recently, a number of 'extinct' groups have been recognised in the Early Cretaceous, including diplodocids in Gondwana, and non-titanosauriform macronarians in Laurasia. Turiasauria, a clade of non-neosauropod eusauropods, was originally thought to have been restricted to the Late Jurassic of western Europe. However, its distribution has recently been extended to the Late Jurassic of Tanzania ( Tendaguria tanzaniensis ), as well as to the Early Cretaceous of the USA ( Mierasaurus bobyoungi and Moabosaurus utahensis ), demonstrating the survival of another 'basal' clade across the J/K boundary. Teeth from the Middle Jurassic-Early Cretaceous of western Europe and North Africa have also tentatively been attributed to turiasaurs, whilst recent phylogenetic analyses recovered Late Jurassic taxa from Argentina and China as further members of Turiasauria. Here, an anterior dorsal centrum and neural arch (both NHMUK 1871) from the Early Cretaceous Wealden Supergroup of the UK are described for the first time. NHMUK 1871 shares several synapomorphies with Turiasauria, especially the turiasaurs Moabosaurus and Tendaguria , including: (1) a strongly dorsoventrally compressed centrum; (2) the retention of prominent epipophyses; and (3) an extremely low, non-bifid neural spine. NHMUK 1871 therefore represents the first postcranial evidence for Turiasauria from European deposits of Early Cretaceous age. Although turiasaurs show clear heterodont dentition, only broad, characteristically 'heart'-shaped teeth can currently be attributed to Turiasauria with confidence. As such, several putative turiasaur occurrences based on isolated teeth from Europe, as well as the Middle Jurassic and Early Cretaceous of Africa, cannot be confidently referred to Turiasauria. Unequivocal evidence for turiasaurs is therefore restricted to the late Middle Jurassic-Early Cretaceous of western Europe, the Late Jurassic of Tanzania, and the late Early Cretaceous of the USA, although remains from elsewhere might ultimately demonstrate that the group had a near-global distribution., Competing Interests: The author declares that they have no competing interests.

The fragmentation of the supercontinent Pangaea has been suggested to have had a profound impact on Mesozoic terrestrial vertebrate distributions. One current paradigm is that geographic isolation produced an endemic biota in East Asia during the Jurassic, while simultaneously preventing diplodocoid sauropod dinosaurs and several other tetrapod groups from reaching this region. Here we report the discovery of the earliest diplodocoid, and the first from East Asia, to our knowledge, based on fossil material comprising multiple individuals and most parts of the skeleton of an early Middle Jurassic dicraeosaurid. The new discovery challenges conventional biogeographical ideas, and suggests that dispersal into East Asia occurred much earlier than expected. Moreover, the age of this new taxon indicates that many advanced sauropod lineages originated at least 15 million years earlier than previously realised, achieving a global distribution while Pangaea was still a coherent landmass.

Brachiosauridae is a clade of titanosauriform sauropod dinosaurs that includes the well-known Late Jurassic taxa Brachiosaurus and Giraffatitan . However, there is disagreement over the brachiosaurid affinities of most other taxa, and little consensus regarding the clade's composition or inter-relationships. An unnamed partial sauropod skeleton was collected from middle-late Oxfordian (early Late Jurassic) deposits in Damparis, in the Jura department of eastern France, in 1934. Since its brief description in 1943, this specimen has been informally known in the literature as the 'Damparis sauropod' and 'French Bothriospondylus ', and has been considered a brachiosaurid by most authors. If correctly identified, this would make the specimen the earliest known titanosauriform. Coupled with its relatively complete nature and the rarity of Oxfordian sauropod remains in general, this is an important specimen for understanding the early evolution of Titanosauriformes. Full preparation and description of this specimen, known from teeth, vertebrae and most of the appendicular skeleton of a single individual, recognises it as a distinct taxon: Vouivria damparisensis gen. et sp. nov. Phylogenetic analysis of a data matrix comprising 77 taxa (including all putative brachiosaurids) scored for 416 characters recovers a fairly well resolved Brachiosauridae. Vouivria is a basal brachiosaurid, confirming its status as the stratigraphically oldest known titanosauriform. Brachiosauridae consists of a paraphyletic array of Late Jurassic forms, with Europasaurus , Vouivria and Brachiosaurus recovered as successively more nested genera that lie outside of a clade comprising ( Giraffatitan + Sonorasaurus ) + ( Lusotitan + ( Cedarosaurus + Venenosaurus )). Abydosaurus forms an unresolved polytomy with the latter five taxa. The Early Cretaceous South American sauropod Padillasaurus was previously regarded as a brachiosaurid, but is here placed within Somphospondyli. A recent study contended that a number of characters used in a previous iteration of this data matrix are 'biologically related', and thus should be excluded from phylogenetic analysis. We demonstrate that almost all of these characters show variation between taxa, and implementation of sensitivity analyses, in which these characters are excluded, has no effect on tree topology or resolution. We argue that where there is morphological variation, this should be captured, rather than ignored. Unambiguous brachiosaurid remains are known only from the USA, western Europe and Africa, and the clade spanned the Late Jurassic through to the late Albian/early Cenomanian, with the last known occurrences all from the USA. Regardless of whether their absence from the Cretaceous of Europe, as well as other regions entirely, reflects regional extinctions and genuine absences, or sampling artefacts, brachiosaurids appear to have become globally extinct by the earliest Late Cretaceous., Competing Interests: The authors declare there are no competing interests.

The Late Jurassic to Early Cretaceous interval represents a time of environmental upheaval and cataclysmic events, combined with disruptions to terrestrial and marine ecosystems. Historically, the Jurassic/Cretaceous (J/K) boundary was classified as one of eight mass extinctions. However, more recent research has largely overturned this view, revealing a much more complex pattern of biotic and abiotic dynamics than has previously been appreciated. Here, we present a synthesis of our current knowledge of Late Jurassic-Early Cretaceous events, focusing particularly on events closest to the J/K boundary. We find evidence for a combination of short-term catastrophic events, large-scale tectonic processes and environmental perturbations, and major clade interactions that led to a seemingly dramatic faunal and ecological turnover in both the marine and terrestrial realms. This is coupled with a great reduction in global biodiversity which might in part be explained by poor sampling. Very few groups appear to have been entirely resilient to this J/K boundary 'event', which hints at a 'cascade model' of ecosystem changes driving faunal dynamics. Within terrestrial ecosystems, larger, more-specialised organisms, such as saurischian dinosaurs, appear to have suffered the most. Medium-sized tetanuran theropods declined, and were replaced by larger-bodied groups, and basal eusauropods were replaced by neosauropod faunas. The ascent of paravian theropods is emphasised by escalated competition with contemporary pterosaur groups, culminating in the explosive radiation of birds, although the timing of this is obfuscated by biases in sampling. Smaller, more ecologically diverse terrestrial non-archosaurs, such as lissamphibians and mammaliaforms, were comparatively resilient to extinctions, instead documenting the origination of many extant groups around the J/K boundary. In the marine realm, extinctions were focused on low-latitude, shallow marine shelf-dwelling faunas, corresponding to a significant eustatic sea-level fall in the latest Jurassic. More mobile and ecologically plastic marine groups, such as ichthyosaurs, survived the boundary relatively unscathed. High rates of extinction and turnover in other macropredaceous marine groups, including plesiosaurs, are accompanied by the origin of most major lineages of extant sharks. Groups which occupied both marine and terrestrial ecosystems, including crocodylomorphs, document a selective extinction in shallow marine forms, whereas turtles appear to have diversified. These patterns suggest that different extinction selectivity and ecological processes were operating between marine and terrestrial ecosystems, which were ultimately important in determining the fates of many key groups, as well as the origins of many major extant lineages. We identify a series of potential abiotic candidates for driving these patterns, including multiple bolide impacts, several episodes of flood basalt eruptions, dramatic climate change, and major disruptions to oceanic systems. The J/K transition therefore, although not a mass extinction, represents an important transitional period in the co-evolutionary history of life on Earth., (© 2016 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Australian dinosaurs have played a rare but controversial role in the debate surrounding the effect of Gondwanan break-up on Cretaceous dinosaur distribution. Major spatiotemporal gaps in the Gondwanan Cretaceous fossil record, coupled with taxon incompleteness, have hindered research on this effect, especially in Australia. Here we report on two new sauropod specimens from the early Late Cretaceous of Queensland, Australia, that have important implications for Cretaceous dinosaur palaeobiogeography. Savannasaurus elliottorum gen. et sp. nov. comprises one of the most complete Cretaceous sauropod skeletons ever found in Australia, whereas a new specimen of Diamantinasaurus matildae includes the first ever cranial remains of an Australian sauropod. The results of a new phylogenetic analysis, in which both Savannasaurus and Diamantinasaurus are recovered within Titanosauria, were used as the basis for a quantitative palaeobiogeographical analysis of macronarian sauropods. Titanosaurs achieved a worldwide distribution by at least 125 million years ago, suggesting that mid-Cretaceous Australian sauropods represent remnants of clades which were widespread during the Early Cretaceous. These lineages would have entered Australasia via dispersal from South America, presumably across Antarctica. High latitude sauropod dispersal might have been facilitated by Albian-Turonian warming that lifted a palaeoclimatic dispersal barrier between Antarctica and South America.

The Early Jurassic of China has long been recognized for its diverse array of sauropodomorph dinosaurs. However, the contribution of this record to our understanding of early sauropod evolution is complicated by a dearth of information on important transitional taxa. We present a revision of the poorly known taxon Sanpasaurus yaoi Young, 1944 from the late Early Jurassic Ziliujing Formation of Sichuan Province, southwest China. Initially described as the remains of an ornithopod ornithischian, we demonstrate that the material catalogued as IVPP V156 is unambiguously referable to Sauropoda. Although represented by multiple individuals of equivocal association, Sanpasaurus is nonetheless diagnosable with respect to an autapomorphic feature of the holotypic dorsal vertebral series. Additional material thought to be collected from the type locality is tentatively referred to Sanpasaurus . If correctly attributed, a second autapomorphy is present in a referred humerus. The presence of a dorsoventrally compressed pedal ungual in Sanpasaurus is of particular interest, with taxa possessing this typically 'vulcanodontid' character exhibiting a much broader geographic distribution than previously thought. Furthermore, the association of this trait with other features of Sanpasaurus that are broadly characteristic of basal eusauropods underscores the mosaic nature of the early sauropod-eusauropod transition. Our revision of Sanpasaurus has palaeobiogeographic implications for Early Jurassic sauropods, with evidence that the group maintained a cosmopolitan Pangaean distribution., Competing Interests: The authors declare that they have no competing interests.

Reconstructing deep time trends in biodiversity remains a central goal for palaeobiologists, but our understanding of the magnitude and tempo of extinctions and radiations is confounded by uneven sampling of the fossil record. In particular, the Jurassic/Cretaceous (J/K) boundary, 145 million years ago, remains poorly understood, despite an apparent minor extinction and the radiation of numerous important clades. Here we apply a rigorous subsampling approach to a comprehensive tetrapod fossil occurrence data set to assess the group's macroevolutionary dynamics through the J/K transition. Although much of the signal is exclusively European, almost every higher tetrapod group was affected by a substantial decline across the boundary, culminating in the extinction of several important clades and the ecological release and radiation of numerous modern tetrapod groups. Variation in eustatic sea level was the primary driver of these patterns, controlling biodiversity through availability of shallow marine environments and via allopatric speciation on land.

The colossal size and body plan of sauropod dinosaurs are unparalleled in terrestrial vertebrates. However, to date, there have been only limited attempts to examine temporal and phylogenetic patterns in the sauropod bauplan. Here, we combine three-dimensional computational models with phylogenetic reconstructions to quantify the evolution of whole-body shape and body segment properties across the sauropod radiation. Limitations associated with the absence of soft tissue preservation in fossils result in large error bars about mean absolute body shape predictions. However, applying any consistent skeleton : body volume ratio to all taxa does yield changes in body shape that appear concurrent with major macroevolutionary events in sauropod history. A caudad shift in centre-of-mass (CoM) in Middle Triassic Saurischia, associated with the evolution of bipedalism in various dinosaur lineages, was reversed in Late Triassic sauropodomorphs. A craniad CoM shift coincided with the evolution of quadrupedalism in the Late Triassic, followed by a more striking craniad shift in Late Jurassic-Cretaceous titanosauriforms, which included the largest sauropods. These craniad CoM shifts are strongly correlated with neck enlargement, a key innovation in sauropod evolution and pivotal to their gigantism. By creating a much larger feeding envelope, neck elongation is thought to have increased feeding efficiency and opened up trophic niches that were inaccessible to other herbivores. However, we find that relative neck size and CoM position are not strongly correlated with inferred feeding habits. Instead the craniad CoM positions of titanosauriforms appear closely linked with locomotion and environmental distributions, potentially contributing to the continued success of this group until the end-Cretaceous, with all other sauropods having gone extinct by the early Late Cretaceous.