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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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Invertebrate bioerosion on fossil bone can contribute to reconstructions of benthic taxonomic assemblages and inform us about oxygenation levels, water depth and exposure time on the seafloor prior to burial. However, these traces are not commonly described in the fossil record. To date, there have been only 13 published studies describing a total of 15 instances of invertebrate bioerosion on marine reptile fossil bones from the Mesozoic globally. We surveyed the collections of several UK museums with substantial occurrences of Mesozoic marine reptiles for evidence of invertebrate bioerosion. Here, we document 153 specimens exhibiting 171 newly recorded instances of invertebrate bioerosion on Jurassic and Cretaceous marine reptile bones. Several major bioeroding taxonomic groups are identified. Within the geological strata of the United Kingdom, there is a higher prevalence of bioerosion in the Cretaceous relative to the Jurassic, despite greater sampling of specimens from the Jurassic. Although biotic turnover and food web restructuring might have played a role, potentially pertaining to heightened productivity during the later stages of the Mesozoic Marine Revolution, we consider it more likely that this temporal change corresponds to differences in depositional environment and taphonomic history between the sampled rock units. In particular, the Cretaceous deposits are characterized by heightened oxygenation levels relative to their Jurassic counterparts, as well as reworking, which would have allowed two phases of bioerosion. A spatiotemporally broader dataset on invertebrate bioerosion on vertebrate bone will be important in further testing this and other hypotheses. [ABSTRACT FROM AUTHOR]

Aim: Understanding connections between environment and biodiversity is crucial for conservation, identifying causes of ecosystem stress, and predicting population responses to changing environments. Explaining biodiversity requires an understanding of how species richness and environment covary across scales. Here, we identify scales and locations at which biodiversity is generated and correlates with environment. Location: Full latitudinal range per continent. Time Period: Present day. Major Taxa Studied: Terrestrial vertebrates: all mammals, carnivorans, bats, songbirds, hummingbirds, amphibians. Methods: We describe the use of wavelet power spectra, cross‐power and coherence for identifying scale‐dependent trends across Earth's surface. Spectra reveal scale‐ and location‐dependent coherence between species richness and topography (E), mean annual precipitation (Pn), temperature (Tm) and annual temperature range (ΔT). Results: >97% of species richness of taxa studied is generated at large scales, that is, wavelengths ≳103 km, with 30%–69% generated at scales ≳104 km. At these scales, richness tends to be highly coherent and anti‐correlated with E and ΔT, and positively correlated with Pn and Tm. Coherence between carnivoran richness and ΔT is low across scales, implying insensitivity to seasonal temperature variations. Conversely, amphibian richness is strongly anti‐correlated with ΔT at large scales. At scales ≲103 km, examined taxa, except carnivorans, show highest richness within the tropics. Terrestrial plateaux exhibit high coherence between carnivorans and E at scales ∼103 km, consistent with contribution of large‐scale tectonic processes to biodiversity. Results are similar across different continents and for global latitudinal averages. Spectral admittance permits derivation of rules‐of‐thumb relating long‐wavelength environmental and species richness trends. Main Conclusions: Sensitivities of mammal, bird and amphibian populations to environment are highly scale dependent. At large scales, carnivoran richness is largely independent of temperature and precipitation, whereas amphibian richness correlates strongly with precipitation and temperature, and anti‐correlates with temperature range. These results pave the way for spectral‐based calibration of models that predict biodiversity response to climate change scenarios. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Although sauropod dinosaur bones are the most abundant vertebrate fossils found in the Upper Cretaceous Winton Formation of northeast Australia, only subadult and adult specimens have been described to date. Herein, we describe the first juvenile sauropod from Australia, derived from the Winton Formation (Cenomanian–lower Turonian). The preserved material belongs to a single individual and is sufficiently diagnostic to classify as a juvenile Diamantinasaurus matildae—the third specimen to be referred to the species. It also enables the identification of a new local autapomorphy for Diamantinasaurus: a distinct tuberosity on the medial surface of the scapula, posterior to the junction of the acromion and the distal blade. Nevertheless, several morphological changes are observable between the juvenile and the two adult skeletons of Diamantinasaurus matildae. These include less well-defined or entirely absent muscle attachment sites on the juvenile bones relative to the heavily scarred and rugose adult specimens. Overlapping elements between the juvenile and the two adult skeletons indicate allometric changes for Diamantinasaurus matildae throughout ontogeny, with limb bones growing at a more rapid proportional rate than other skeletal elements. Finally, we review the global record of juvenile sauropod remains, demonstrating that the growth patterns of sauropods vary greatly between taxa. Although titanosaurs display a range of isometry and allometry in the growth of individual bones, it appears that allometric growth was the primary pattern for this group. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Hudiesaurus sinojapanorum is a Late Jurassic sauropod from northwestern China that was erected on the basis of a cervicodorsal vertebra, four teeth, and a nearly complete forelimb. However, re-evaluation of this material, and comparisons with other taxa, indicate that there are few grounds for regarding these specimens as congeneric. Consequently, although we retain the vertebra as the holotype specimen of Hudiesaurus, the forelimb is assigned to a new taxon—Rhomaleopakhus turpanensis, gen. et sp. nov. The teeth previously referred to Hudiesaurus are poorly preserved but resemble those of several other 'core Mamenchisaurus-like taxa' (CMTs) from East Asia, such as Mamenchisaurus sinocanadorum. Phylogenetic analyses confirm that Hudiesaurus is a CMT and the sister taxon of Xinjiangtitan. Despite some uniquely shared features, their large size, and close geographic provenance, Hudiesaurus and Xinjiangtitan are retained as distinct genera based on their stratigraphic separation and numerous anatomical differences. Rhomaleopakhus is also shown to be a CMT in all analyses, being most closely related to Chuanjiesaurus and Analong. We link the convergent evolution of robust antebrachia and an enlarged olecranon in CMTs, titanosaurs, and some ornithischians (e.g., ceratopsids) to a more flexed orientation of the forearm, an enhanced role for the forelimb in locomotion, and an anterior shift in the whole-body center of mass. CMTs and titanosaurs potentially converged on a feeding strategy in which the ability to increase browse height via bipedal rearing was sacrificed in return for more efficient locomotion that improved travel between patchily distributed food sources. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The titanosaurian sauropod dinosaur Diamantinasaurus matildae is represented by two individuals from the Cenomanian–lower Turonian 'upper' Winton Formation of central Queensland, north-eastern Australia. The type specimen has been described in detail, whereas the referred specimen, which includes several elements not present in the type series (partial skull, atlas, axis and postaxial cervical vertebrae), has only been described briefly. Herein, we provide a comprehensive description of this referred specimen, including a thorough assessment of the external and internal anatomy of the braincase, and identify several new autapomorphies of D. matildae. Via an expanded data matrix consisting of 125 taxa scored for 552 characters, we recover a close, well-supported relationship between Diamantinasaurus and its contemporary, Savannasaurus elliottorum. Unlike previous iterations of this data matrix, under a parsimony framework we consistently recover Diamantinasaurus and Savannasaurus as early-diverging members of Titanosauria using both equal weighting and extended implied weighting, with the overall topology largely consistent between analyses. We erect a new clade, named Diamantinasauria herein, that also includes the contemporaneous Sarmientosaurus musacchioi from southern Argentina, which shares several cranial features with the referred Diamantinasaurus specimen. Thus, Diamantinasauria is represented in the mid-Cretaceous of both South America and Australia, supporting the hypothesis that some titanosaurians, in addition to megaraptoran theropods and possibly some ornithopods, were able to disperse between these two continents via Antarctica. Conversely, there is no evidence for rebbachisaurids in Australia, which might indicate that they were unable to expand into high latitudes before their extinction in the Cenomanian–Turonian. Likewise, there is no evidence for titanosaurs with procoelous caudal vertebrae in the mid-Cretaceous Australian record, despite scarce but compelling evidence for their presence in both Antarctica and New Zealand during the Campanian–Maastrichtian. These later titanosaurs presumably dispersed into these landmasses from South America before the Campanian (~85 Mya), when seafloor spreading between Zealandia and Australia commenced. Although Australian mid-Cretaceous dinosaur faunas appear to be cosmopolitan at higher taxonomic levels, closer affinities with South America at finer scales are becoming better supported for sauropods, theropods and ornithopods. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Mammals are the only surviving group of Cynodontia, a synapsid clade that first appeared in the fossil record in the late Permian, ~260 million years ago. Using three metrics that capture skeletal completeness, we quantify the quality of the early cynodont fossil record in time and space to evaluate the impact of sampling and preservational biases on our understanding of the group's evolutionary history. There is no consistent global sampling signal for early cynodonts. Completeness of the cynodont fossil record increases across the Permian–Triassic boundary, peaking in the Early to early Late Triassic. This peak is dominated by specimens from southern Africa and South America, where a highly seasonal climate probably favoured preservation. Completeness is generally lower thereafter, correlating with a shift from a Gondwanan to a predominantly Laurasian fossil record. Phylogenetic and stratigraphic congruence in early cynodonts is high, although their fossil record exhibits less skeletal completeness overall than other tetrapod clades, including the contemporaneous anomodont synapsids. This discrepancy could be due to differences in the diagnosability of their fossils, especially for small‐bodied species. Establishing the timing and assembly of derived ('mammalian') anatomical features in Cynodontia is obscured by sampling. Two of the major nodes at which acquisition of mammalian features is concentrated (Cynodontia and Mammaliamorpha) suffer from lengthy intervals of poor sampling before becoming abundant parts of tetrapod faunas. Low completeness in these intervals limits our ability to determine when certain 'key' mammalian characteristics evolved, or to identify the selective pressures that might have driven their origins. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Once a much more globally widespread crocodylian clade, Tomistominae is today represented by just one species, Tomistoma schlegelii (the false gharial), restricted to south-east Asia. Although tomistomine fossil occurrences are recognized from the early Eocene (∼55 Ma) onwards, their remains are often incomplete, making appropriate taxonomic classification within the group problematic. This is especially pertinent to several taxa from the Miocene of Europe, which were historically erected from fragmentary remains. Here we re-examine and describe four approximately contemporaneous taxa from Malta and Italy to determine their taxonomy and phylogenetic affinities: Melitosaurus champsoides, Tomistoma calaritanum, Tomistoma gaudense and Tomistoma lyceense. We place them into a phylogenetic analysis for the first time, comprising 70 taxa scored for 244 characters, several of which are revised or novel, and apply a number of character weighting strategies. Whereas 'Tomistoma lyceense' is deemed to be an indeterminate tomistomine, a unique combination of features confirms Melitosaurus champsoides, Tomistoma calaritanum and Tomistoma gaudense as three distinct species. These three taxa are recovered as derived tomistomines, with characters such as a posterior maxillary process between the lacrimal and nasal, large supratemporal fenestrae that are wider than long, and the posteromedial alignment of the last three premaxillary teeth, suggesting a close relationship with the approximately contemporaneous European taxa, Tomistoma lusitanica and Gavialosuchus eggenburgensis. It is unlikely that any of these species belong to Tomistoma, with the possibility that they can all be classified under Melitosaurus and Gavialosuchus. However, we retain them in open nomenclature pending reassessment of the remaining European Miocene tomistomines. Our taxonomic and phylogenetic revision helps to elucidate past tomistomine diversity in the Miocene of the Mediterranean region, prior to the group's extirpation, and is an important first step in resolving the complicated history of European tomistomine systematics. [ABSTRACT FROM AUTHOR]

Spatiotemporal changes in fossil specimen completeness can bias our understanding of a group's evolutionary history. The quality of the sauropodomorph fossil record was assessed a decade ago, but the number of valid species has since increased by 60%, and 17% of the taxa from that study have since undergone taxonomic revision. Here, we assess how 10 years of additional research has changed our outlook on the group's fossil record. We quantified the completeness of all 307 sauropodomorph species currently considered valid using the skeletal completeness metric, which calculates the proportion of a complete skeleton preserved for each taxon. Taxonomic and stratigraphic age revisions, rather than new species, are the drivers of the most significant differences between the current results and those of the previous assessment. No statistical differences appeared when we use our new dataset to generate temporal completeness curves based solely on taxa known in 2009 or 1999. We now observe a severe drop in mean completeness values across the Jurassic–Cretaceous boundary that never recovers to pre‐Cretaceous levels. Explaining this pattern is difficult, as we find no convincing evidence that it is related to environmental preferences or body size changes. Instead, it might result from: (1) reduction of terrestrial fossil preservation space due to sea level rise; (2) ecological specificities and relatively high diagnosability of Cretaceous species; and/or (3) increased sampling of newly explored sites with many previously unknown taxa. Revisiting patterns in this manner allows us to test the longevity of conclusions made in previous quantitative studies. [ABSTRACT FROM AUTHOR]

Diplocynodon is a genus of basal alligatoroid comprising nine species, which spanned the late Palaeocene to middle Miocene of Europe. Despite recent revisions of most Diplocynodon species, one of the earliest named and most complete, Diplocynodon hantoniensis , has not been re-described for over 150 years. This species is known from the remains of numerous individuals from the Priabonian (late Eocene) Headon Hill Formation, which crops out at Hordle (Hordwell) Cliff in Hampshire, United Kingdom. Here we re-describe and diagnose Diplocynodon hantoniensis , providing the first detailed description of postcranial anatomy in Diplocynodon , and indeed any basal alligatoroid. Diplocynodon hantoniensis is diagnosed by four autapomorphies, including retention of the ectopterygoid–pterygoid flexure through ontogeny and a unique anterior process of the ectopterygoid adjacent to the posteriormost maxillary alveoli. A critical review of previously referred remains from elsewhere in Europe and the USA restricts Diplocynodon hantoniensis to the late Eocene of the UK. Through comparisons with extant crocodylians, the well-preserved postcranial skeleton enables the interpretation of numerous muscle attachments in the forelimbs and hindlimbs, providing a potentially rich source of character data for future phylogenetic analyses. Based on a comparison of humeral morphology between a large sample of crocodylian species, we outline two new morphological characters in the humerus. We include D. hantoniensis in a phylogenetic analysis, including all putative Diplocynodon species (103 taxa scored for 187 characters). We use four different character-weighting schemes: equal weighting, implied weighting (k value = 8) and extended implied weighting with k -values of 4 and 8. In general, these weighted analyses produce congruent results with the equal-weights analysis, and increase the resolution within Diplocynodon. We recover a monophyletic Diplocynodon in three of the four analyses. However, the fourth analysis, with the strongest downweighting of homoplastic characters and missing data (extended implied weighting with k = 4), recovers the Palaeocene Diplocynodon remensis outside Diplocynodon. Our comprehensive revision of one of the most completely known Diplocynodon species facilitates comparisons in the genus, as well as between other basal alligatoroids, and forms the basis for comparing postcranial anatomy in other fossil crocodylians. [ABSTRACT FROM AUTHOR]

The 24 extant crocodylian species are the remnants of a once much more diverse and widespread clade. Crocodylomorpha has an approximately 230 million year evolutionary history, punctuated by a series of radiations and extinctions. However, the group's fossil record is biased. Previous studies have reconstructed temporal patterns in subsampled crocodylomorph palaeobiodiversity, but have not explicitly examined variation in spatial sampling, nor the quality of this record. We compiled a dataset of all taxonomically diagnosable non‐marine crocodylomorph species (393). Based on the number of phylogenetic characters that can be scored for all published fossils of each species, we calculated a completeness value for each taxon. Mean average species completeness (56%) is largely consistent within subgroups and for different body size classes, suggesting no significant biases across the crocodylomorph tree. In general, average completeness values are highest in the Mesozoic, with an overall trend of decreasing completeness through time. Many extant taxa are identified in the fossil record from very incomplete remains, but this might be because their provenance closely matches the species' present‐day distribution, rather than through autapomorphies. Our understanding of nearly all crocodylomorph macroevolutionary 'events' is essentially driven by regional patterns, with no global sampling signal. Palaeotropical sampling is especially poor for most of the group's history. Spatiotemporal sampling bias impedes our understanding of several Mesozoic radiations, whereas molecular divergence times for Crocodylia are generally in close agreement with the fossil record. However, the latter might merely be fortuitous, i.e. divergences happened to occur during our ephemeral spatiotemporal sampling windows. [ABSTRACT FROM AUTHOR]

The Late Jurassic Tendaguru Formation of Tanzania, southeastern Africa, records a rich sauropod fauna, including the diplodocoids Dicraeosaurus and Tornieria, and the brachiosaurid titanosauriform Giraffatitan. However, the taxonomic affinities of other sympatric sauropod taxa are poorly understood. Here, we critically reassess and redescribe these problematic taxa, and present the largest phylogenetic analysis for sauropods (117 taxa scored for 542 characters) to explore their placement in Eusauropoda. Janenschia robusta has played a prominent role in discussions of titanosaur origins, with various authors referring at least some remains to Titanosauria, a clade otherwise known only from the Cretaceous. Redescription of the holotype of Janenschia, and all referable remains, supports its validity and placement as a non-neosauropod eusauropod. It forms a clade with Haestasaurus from the earliest Cretaceous of the UK, and the Middle/Late Jurassic Chinese sauropod Bellusaurus. Phylogenetic analysis and CT scans of the internal pneumatic tissue structure of Australodocus bohetii tentatively support a non-titanosaurian somphospondylan identification, making it the only known pre-Cretaceous representative of that clade. New information on the internal pneumatic tissue structure of the dorsal vertebrae of the enigmatic Tendaguria tanzaniensis, coupled with a full redescription, results in its novel placement as a turiasaur. Tendaguria is the sister taxon of Moabosaurus, from the Early Cretaceous of North America, and is the first turiasaur recognized from Gondwana. A previously referred caudal sequence cannot be assigned to Janenschia and displays several features that indicate a close relationship with Middle–Late Jurassic East Asian mamenchisaurids. It can be diagnosed by six autapomorphies, so we erect the new taxon Wamweracaudia keranjei gen. et sp. nov. The presence of a mamenchisaurid in the Late Jurassic of southern Gondwana indicates an earlier and more widespread diversification of this clade than previously realized, prior to the geographic isolation of East Asia. Our revised phylogenetic dataset sheds light on the evolutionary history of Eusauropoda, including supporting a basal diplodocoid placement for Haplocanthosaurus, and elucidating the interrelationships of rebbachisaurids. The Tendaguru Formation shares representatives of nearly all sauropod lineages with Middle Jurassic–earliest Cretaceous global faunas, but displays a greater range of diversity than any of those faunas considered individually. Biogeographic analysis indicates that the Tendaguru sauropod fauna was assembled as a result of three main phenomena during the late Early and/or Middle Jurassic: (1) invasions from Euramerica (brachiosaurids, turiasaurs); (2) endemism in west Gondwana (dicraeosaurids, diplodocids); and (3) regional extinctions that restricted the ranges of once widespread groups (mamenchisaurids, the Janenschia lineage). Multiple dispersals across the Central Gondwanan Desert are required to explain the distributions of Jurassic sauropods, suggesting that this geographic feature was at most a filter barrier that became easier to cross during the late Middle Jurassic. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The titanosaurian sauropod dinosaur Mendozasaurus neguyelap is represented by several partial skeletons from a single locality within the Coniacian (lower Upper Cretaceous) Sierra Barrosa Formation in the south of Mendoza Province, northern Neuquén Basin, Argentina. A detailed revision of Mendozasaurus, including previously undocumented remains from the holotype site, allows us to more firmly establish its position within Titanosauria, as well as enabling an emended diagnosis of this taxon. Autapomorphies include: (1) middle and posterior cervical vertebrae with tall and transversely expanded neural spines that are wider than the centra, formed laterally by spinodiapophyseal laminae that are not connected with the pre- or postzygapophyses; (2) anterior caudal vertebrae (excluding anteriormost) with ventrolateral ridge-like expansion of prezygapophyses; and (3) humerus with divided lateral distal condyle on anterior surface. New remains demonstrate that the presacral vertebrae of Mendozasaurus were not unusually short anteroposteriorly, with this compression instead resulting from taphonomic crushing. Comparative studies of articulated pedes of other taxa allow us to interpret that the pedal formula of Mendozasaurus was 2-2-2-2-0, based on disarticulated bones that form a right hind foot. Mendozasaurus was incorporated into an expanded version of a titanosauriform-focussed phylogenetic data matrix, along with several other contemporaneous South American titanosaurs. The resultant data matrix comprises 84 taxa scored for 423 characters, and our phylogenetic analysis recovers Mendozasaurus as the most basal member of a diverse Lognkosauria, including Futalognkosaurus and the gigantic titanosaurs Argentinosaurus, Notocolossus, Patagotitan and Puertasaurus. Lognkosauria forms a clade with Rinconsauria (Muyelensaurus + Rinconsaurus), with Epachthosaurus and Pitekunsaurus recovered at the base of this grouping. A basal lithostrotian position for this South American clade is well supported, contrasting with some analyses that have placed these taxa outside of Lithostrotia or closer to Saltasauridae. The sister clade to this South American group is composed of an array of near-global taxa and supports the hypothesis that most titanosaurian clades were widespread by the Early–middle Cretaceous. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Despite being globally widespread and abundant throughout much of the Mesozoic, the early record of sauropod dinosaur evolution is extremely poor. As such, any new remains can provide significant additions to our understanding of this important radiation. Here, we describe two sauropod middle cervical vertebrae from a new Early Jurassic locality in the Haute Moulouya Basin, Central High Atlas of Morocco. The possession of opisthocoelous centra, a well-developed system of centrodiapophyseal laminae, and the higher elevation of the postzygapophyses relative to the prezygapophyses, all provide strong support for a placement within Sauropoda. Absence of pneumaticity indicates non-neosauropod affinities, and several other features, including a tubercle on the dorsal margin of the prezygapophyses and an anteriorly slanting neural spine, suggest close relationships with various basal eusauropods, such as the Middle Jurassic taxa Jobaria tiguidensis and Patagosaurus fariasi. Phylogenetic analyses also support a position close to the base of Eusauropoda. The vertebrae differ from the only other Early Jurassic African sauropod dinosaurs preserving overlapping remains (the Moroccan Tazoudasaurus naimi and South African Pulanesaura eocollum), as well as stratigraphically younger taxa, although we refrain from erecting a new taxon due to the limited nature of the material. These new specimens represent one of the earliest eusauropod taxa and are an important additional data point for elucidating the early evolution of the clade. [ABSTRACT FROM AUTHOR]