New occurrences of the bone-eating worm Osedax from Late Cretaceous marine reptiles and implications for its biogeography and diversification.

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.

Neuroanatomy of the crocodylian Tomistoma dowsoni from the Miocene of North Africa provides insights into the evolutionary history of gavialoids.

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.

Asteroid impact, not volcanism, caused the end-Cretaceous dinosaur extinction.

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 [CO2]-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.

Evolutionary simulations clarify and reconcile biodiversity-disturbance models.

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.

Spatial sampling heterogeneity limits the detectability of deep time latitudinal biodiversity gradients.

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.

The apparent exponential radiation of Phanerozoic land vertebrates is an artefact of spatial sampling biases.

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.

Near-Stasis in the Long-Term Diversification of Mesozoic Tetrapods.

How did evolution generate the extraordinary diversity of vertebrates on land? Zero species are known prior to ~380 million years ago, and more than 30,000 are present today. An expansionist model suggests this was achieved by large and unbounded increases, leading to substantially greater diversity in the present than at any time in the geological past. This model contrasts starkly with empirical support for constrained diversification in marine animals, suggesting different macroevolutionary processes on land and in the sea. We quantify patterns of vertebrate standing diversity on land during the Mesozoic-early Paleogene interval, applying sample-standardization to a global fossil dataset containing 27,260 occurrences of 4,898 non-marine tetrapod species. Our results show a highly stable pattern of Mesozoic tetrapod diversity at regional and local levels, underpinned by a weakly positive, but near-zero, long-term net diversification rate over 190 million years. Species diversity of non-flying terrestrial tetrapods less than doubled over this interval, despite the origins of exceptionally diverse extant groups within mammals, squamates, amphibians, and dinosaurs. Therefore, although speciose groups of modern tetrapods have Mesozoic origins, rates of Mesozoic diversification inferred from the fossil record are slow compared to those inferred from molecular phylogenies. If high speciation rates did occur in the Mesozoic, then they seem to have been balanced by extinctions among older clades. An apparent 4-fold expansion of species richness after the Cretaceous/Paleogene (K/Pg) boundary deserves further examination in light of potential taxonomic biases, but is consistent with the hypothesis that global environmental disturbances such as mass extinction events can rapidly adjust limits to diversity by restructuring ecosystems, and suggests that the gradualistic evolutionary diversification of tetrapods was punctuated by brief but dramatic episodes of radiation.

Rates of dinosaur body mass evolution indicate 170 million years of sustained ecological innovation on the avian stem lineage.

Large-scale adaptive radiations might explain the runaway success of a minority of extant vertebrate clades. This hypothesis predicts, among other things, rapid rates of morphological evolution during the early history of major groups, as lineages invade disparate ecological niches. However, few studies of adaptive radiation have included deep time data, so the links between extant diversity and major extinct radiations are unclear. The intensively studied Mesozoic dinosaur record provides a model system for such investigation, representing an ecologically diverse group that dominated terrestrial ecosystems for 170 million years. Furthermore, with 10,000 species, extant dinosaurs (birds) are the most speciose living tetrapod clade. We assembled composite trees of 614-622 Mesozoic dinosaurs/birds, and a comprehensive body mass dataset using the scaling relationship of limb bone robustness. Maximum-likelihood modelling and the node height test reveal rapid evolutionary rates and a predominance of rapid shifts among size classes in early (Triassic) dinosaurs. This indicates an early burst niche-filling pattern and contrasts with previous studies that favoured gradualistic rates. Subsequently, rates declined in most lineages, which rarely exploited new ecological niches. However, feathered maniraptoran dinosaurs (including Mesozoic birds) sustained rapid evolution from at least the Middle Jurassic, suggesting that these taxa evaded the effects of niche saturation. This indicates that a long evolutionary history of continuing ecological innovation paved the way for a second great radiation of dinosaurs, in birds. We therefore demonstrate links between the predominantly extinct deep time adaptive radiation of non-avian dinosaurs and the phenomenal diversification of birds, via continuing rapid rates of evolution along the phylogenetic stem lineage. This raises the possibility that the uneven distribution of biodiversity results not just from large-scale extrapolation of the process of adaptive radiation in a few extant clades, but also from the maintenance of evolvability on vast time scales across the history of life, in key lineages.

Environmental drivers of crocodyliform extinction across the Jurassic/Cretaceous transition.

Crocodyliforms have a much richer evolutionary history than represented by their extant descendants, including several independent marine and terrestrial radiations during the Mesozoic. However, heterogeneous sampling of their fossil record has obscured their macroevolutionary dynamics, and obfuscated attempts to reconcile external drivers of these patterns. Here, we present a comprehensive analysis of crocodyliform biodiversity through the Jurassic/Cretaceous (J/K) transition using subsampling and phylogenetic approaches and apply maximum-likelihood methods to fit models of extrinsic variables to assess what mediated these patterns. A combination of fluctuations in sea-level and episodic perturbations to the carbon and sulfur cycles was primarily responsible for both a marine and non-marine crocodyliform biodiversity decline through the J/K boundary, primarily documented in Europe. This was tracked by high extinction rates at the boundary and suppressed origination rates throughout the Early Cretaceous. The diversification of Eusuchia and Notosuchia likely emanated from the easing of ecological pressure resulting from the biodiversity decline, which also culminated in the extinction of the marine thalattosuchians in the late Early Cretaceous. Through application of rigorous techniques for estimating biodiversity, our results demonstrate that it is possible to tease apart the complex array of controls on diversification patterns in major archosaur clades.

Decoupling speciation and extinction reveals both abiotic and biotic drivers shaped 250 million years of diversity in crocodile-line archosaurs.

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.

Reappraisal of sauropod dinosaur diversity in the Upper Cretaceous Winton Formation of Queensland, Australia, through 3D digitisation and description of new specimens.

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.

The hierarchy of factors predicting the latitudinal diversity gradient.

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.

A nearly complete skull of the sauropod dinosaur Diamantinasaurus matildae from the Upper Cretaceous Winton Formation of Australia and implications for the early evolution of titanosaurs.

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.

Multi-variate models are essential for understanding vertebrate diversification in deep time.

Statistical models are helping palaeontologists to elucidate the history of biodiversity. Sampling standardization has been extensively applied to remedy the effects of uneven sampling in large datasets of fossil invertebrates. However, many vertebrate datasets are smaller, and the issue of uneven sampling has commonly been ignored, or approached using pairwise comparisons with a numerical proxy for sampling effort. Although most authors find a strong correlation between palaeodiversity and sampling proxies, weak correlation is recorded in some datasets. This has led several authors to conclude that uneven sampling does not influence our view of vertebrate macroevolution. We demonstrate that multi-variate regression models incorporating a model of underlying biological diversification, as well as a sampling proxy, fit observed sauropodomorph dinosaur palaeodiversity best. This bivariate model is a better fit than separate univariate models, and illustrates that observed palaeodiversity is a composite pattern, representing a biological signal overprinted by variation in sampling effort. Multi-variate models and other approaches that consider sampling as an essential component of palaeodiversity are central to gaining a more complete understanding of deep time vertebrate diversification.

Climatic and tectonic drivers shaped the tropical distribution of coral reefs.

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.

Biogeographic history of Palearctic caudates revealed by a critical appraisal of their fossil record quality and spatio-temporal distribution.

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.

Climatic constraints on the biogeographic history of Mesozoic dinosaurs.

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.

Sauropod dinosaur teeth from the lower Upper Cretaceous Winton Formation of Queensland, Australia and the global record of early titanosauriforms.

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.

Shifts in food webs and niche stability shaped survivorship and extinction at the end-Cretaceous.

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.