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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 million years ago), combining dinosaur fossil occurrences, past climate data from Earth System models, and habitat suitability modelling. Results show that uniquely among dinosaurs, sauropods occupied climatic niches characterised 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.

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. Diversity within terrestrial tetrapod communities has risen by at most threefold over the last 300 million years, consistent with a constrained model of diversification.

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.

Vertebrate microfossil assemblages in a stratigraphic sequence often yield similar assortments of taxa but at different relative abundances, potentially indicative of marginal paleocommunity changes driven by paleoenvironmental change over time. For example, stratigraphically younger assemblages in the Dinosaur Park Formation (DPF) yield proportionally more aquatic taxa, consistent with marine transgression. However, individual deposits may have received specimens from multiple source paleocommunities over time, limiting our ability to confidently identify ecologically significant, paleocommunity differences through direct assemblage comparisons. We adapted a three-source, two-tracer Bayesian mixing model to quantify proportional contributions from different source habitats to DPF microfossil assemblages. Prior information about the compositions of separate, relatively unmixed terrestrial, freshwater, and marine assemblages from the Belly River Group allowed us to define expected taxon percent abundances for the end-member habitats likely contributing specimens to the mixed deposits. We compared the mixed assemblage and end-member distributions using 21 different combinations of vertebrate taxa. Chondrichthyan, dinosaur, and amphibian occurrence patterns ultimately allowed us to parse the contributions from the potential sources to 14 of the 15 mixed assemblages. The results confirmed a significant decline in terrestrial contributions at younger DPF sites, driven primarily by increased freshwater specimen inputs—not incursions from the adjacent marine paleocommunity. A rising base level likely increased lateral channel migration and the prevalence of freshwater habitats on the landscape, factors that contributed to increased paleocommunity mixing at younger channel deposit sites. Bayesian methods can account for source-mixing bias, which may be common in assemblages associated with major paleoenvironmental changes.

Until now, only one Cenozoic fossil mammal from the Chilean Precordillera (Arica and Parinacota Region) has been reported, Caraguatypotherium munozi (Mesotheriidae: Notoungulata). In this study, we describe a fourth specimen of C. munozi and a new armadillo species, Epipeltephilus caraguensis (Peltephilidae: Cingulata), both collected from a new site closer to the fossiliferous outcrops of the Caragua area (Serravallian – Tortonian). E. caraguensis differs from other members of the family in having: two sulci in the articular surface of the mobile osteoderm; having a tubular, rough and raised anterior edge; a conspicuous transverse depression; and four widely spaced foramina. This taxon represents the youngest known peltephilid from intermediate latitudes and indicates a wide geographic distribution (Patagonia to Central Andes) of the family just prior to its extinction. The new mesothere specimen is 19% larger than previous records. The revision of the dental features of C. munozi allowed the identification of an ambiguous trait in its original diagnosis, i.e. an enamel fracture was misinterpreted with the presence of a posterior sulcus on the talonid of the m3, suggesting that further taxonomic and systematic revision for the Caragua mesothere is necessary. Although the fossil record from the Caragua area is still scarce, mesotheriines seem to be abundant at this latitude, just as has been observed at several early to late Miocene sites such as Chucal (Chile), Cerdas and Nazareno (Bolivia), as well as in southern regions such as Arroyo Chasico and Mendoza (Argentina). The presence of a new peltephilid species in Caragua sustains the hypothesis of provincialism during the Miocene in intermediate latitudes. Our findings also provide further support for probable faunal movements between intermediate and higher latitudes rather than to lower ones.

Vertebrate microfossil bonebeds (VMBs)—localized concentrations of small resilient vertebrate hard parts—are commonly studied to recover otherwise rarely found small-bodied taxa, and to document relative taxonomic abundance and species richness in ancient vertebrate communities. Analyses of taphonomic comparability among VMBs have often found significant differences in size and shape distributions, and thus considered them to be non-isotaphonomic. Such outcomes of “strict” statistical tests of isotaphonomy suggest discouraging limits on the potential for broad, comparative paleoecological reconstruction using VMBs. Yet it is not surprising that sensitive statistical tests highlight variations among VMB sites, especially given the general lack of clarity with regard to the definition of “strict” isotaphonomic comparability. We rigorously sampled and compared six VMB localities representing two distinct paleoenvironments (channel and pond/lake) of the Upper Cretaceous Judith River Formation to evaluate biases related to sampling strategies and depositional context. Few defining distinctions in bioclast size and shape are evident in surface collections, and most site-to-site comparisons of sieved collections are indistinguishable (p≤0.003). These results provide a strong case for taphonomic equivalence among the majority of Judith River VMBs, and bode well for future studies of paleoecology, particularly in relation to investigations of faunal membership and community structure in Late Cretaceous wetland ecosystems. The taphonomic comparability of pond/lake and channel-hosted VMBs in the Judith River Formation is also consistent with a formative model that contends that channel-hosted VMBs were reworked from pre-existing pond/lake assemblages, and thus share taphonomic history.

Theropod dinosaurs from the Late Jurassic of Gondwana are still poorly known, with Elaphrosaurus bambergi Janensch, 1920, from the late Kimmeridgian of Tendaguru, Tanzania, being the only taxon represented by more than isolated remains from Africa. Having long been considered a coelurosaurian, more specifically an ornithomimosaur, Elaphrosaurus is currently regarded as a basal ceratosaur. Here, we revise the osteology and phylogenetic position of this important taxon. Elaphrosaurus shows many unusual osteological characters, including extremely elongated and constricted cervical vertebrae, an expansive shoulder girdle with strongly modified forelimbs, a relatively small ilium, and elongate hindlimbs with a very small ascending process of the astragalus that is fused to the tibia. We found this taxon to share many derived characters with noasaurids, such as: strongly elongate cervical and dorsal vertebrae;low, rectangular neural spines in the mid-caudal vertebrae;presence of only an anterior centrodiapophyseal lamina in anterior caudal vertebrae;presence of a wide, U-shaped notch between the glenoid and the anteroventral hook in the coracoid;a laterally flared postacetabular blade of the ilium;a flat anterior side of the distal tibia;and a reduced shaft of metatarsal II. Our analysis placed Elaphrosaurus within a dichotomous Noasauridae as part of a Jurassic subclade, here termed Elaphrosaurinae, that otherwise includes taxa from eastern Asia. These results underscore the long and complex evolutionary history of abelisauroids, which is still only beginning to be understood. (C) 2016 The Linnean Society of London

Like many long-standing dinosaur taxa, Brachiosaurus altithorax from the Upper Jurassic Morrison Formation of North America suffers from taxonomic issues stemming from a relatively incomplete holotype. Lack of anatomical overlap has precluded definitive referral of important specimens, including a mostly complete skull discovered in 1883. We redescribe this skull and some other significant brachiosaurid specimens based on new preparation and computed tomographic (CT) data. We argue that these elements are most parsimoniously referred to B. altithorax. Including these tentatively referred elements in a phylogenetic analysis does not alter the hypothesized relationships of B. altithorax, congruent with the presence of a single brachiosaurid taxon across western North America in the Late Jurassic. Based on CT data, we estimate that B. altithorax had a slower tooth replacement rate than Camarasaurus or the diplodocoids it shared Morrison Formation landscapes with, which suggests a difference in diet and/or feeding strategy. Anat Rec, 303:732-758, 2020. © 2019 Wiley Periodicals, Inc.

The ‘Kem Kem beds’ of Morocco have yielded abundant material of large-bodied theropod dinosaurs, but remains of small theropod taxa are rare. Here, we describe two femora that provide additional information on the diversity of small-bodied theropods in the Gondwanan mid-Cretaceous. An almost complete femur (ROM 64666) represents a noasaurid theropod based on the presence of an elongate anteromedial flange that arises from the distal shaft and terminates proximal to the distal end of the bone; osteohistological analysis indicates that it was from a juvenile individual. It is possible that this femur represents a juvenile Deltadromeus , which is recovered as a putative noasaurid in some phylogenetic analyses. Nevertheless, as the affinities of Deltadromeus are debated, this femur currently represents the first definitive record of a noasaurid from northern Africa and one of the few records of this clade from the early Late Cretaceous. Moreover, if this specimen is not a juvenile Deltadromeus then it probably represents a new addition to the Kem Kem theropod assemblage. A second partial femur (ROM 65779) can be identified only as an indeterminate averostran theropod. It is similar in size to ROM 64666, but the presence of an external fundamental system (EFS) indicates that it pertained to a small-sized adult individual. These observations indicate that the two femora described herein are from different theropod taxa, thereby demonstrating that at least one additional small-bodied theropod taxon was present in the Kem Kem fauna, adding to the already high theropod species-richness of this unit.

Non-avian dinosaurs went extinct 66 million years ago, geologically coincident with the impact of a large bolide (comet or asteroid) during an interval of massive volcanic eruptions and changes in temperature and sea level. There has long been fervent debate about how these events affected dinosaurs. We review a wealth of new data accumulated over the past two decades, provide updated and novel analyses of long-term dinosaur diversity trends during the latest Cretaceous, and discuss an emerging consensus on the extinction's tempo and causes. Little support exists for a global, long-term decline across non-avian dinosaur diversity prior to their extinction at the end of the Cretaceous. However, restructuring of latest Cretaceous dinosaur faunas in North America led to reduced diversity of large-bodied herbivores, perhaps making communities more susceptible to cascading extinctions. The abruptness of the dinosaur extinction suggests a key role for the bolide impact, although the coarseness of the fossil record makes testing the effects of Deccan volcanism difficult.

Summary 1. Body mass is strongly related to both physiological and ecological properties of living organisms. As a result, generating robust, broadly applicable models for estimating body mass in the fossil record provides the opportunity to reconstruct palaeobiology and investigate evolutionary ecology on a large temporal scale. 2. A recent study provided strong evidence that the minimum circumference of stylopodial elements (humerus and femur) is conservatively associated with body mass in living quadrupeds. Unfortunately, this model is not directly applicable to extinct bipeds, such as non-avian dinosaurs. 3. This study presents a new equation that mathematically corrects the quadruped equ ation for use in bipeds. It is derived from the systemic difference in the circumference-to-area scaling relationship of two circles (hypothetical quadruped) and one circle (hypothetical biped), which represent the cross-section of the main weight-bearing limb bones. 4. When applied to a newly constructed data set of femoral circumferences and body masses in living birds, the new equation reveals errors that are significantly lower than other published equations, but significantly higher than the error inherent in the avian data set. Such errors, however, are expected given the unique overall femoral circumference–body mass scaling relationship found in birds. 5. Body mass estimates for a sample of bipedal dinosaurs using the new model are consistent with recent estimates based on volumetric life reconstructions, but, in contrast, this equation is simpler to use, with the concomitant potential to provide a wider set of body mass estimates for extinct bipeds. 6. Although it is evident that no one estimation model is flawless, the combined use of the corrected quadrupedal equations and the previously published quadrupedal equation offer a consistent approach with which to estimate body masses in both quadrupeds and bipeds. These models have implications for conducting large-scale macroevolutionary analyses of body size throughout the evolutionary history of terrestrial vertebrates, and, in particular, across major changes in body plan, such as the evolution of bipedality in archosaurs and quadrupedality in

The vertebrate fauna of the Cloverly Formation has been studied for more than 75 years, but remains poorly sampled and incompletely understood. We undertook an extensive survey of the formation that resulted in the discovery of several new, highly productive vertebrate microfossil bonebeds (VMBs). Comprehensive sampling of these and other sites has nearly doubled the known vertebrate diversity of the Cloverly Formation. In addition to the comparatively well-known dinosaurs, this augmented faunal list includes hybodontoid sharks, numerous bony fishes, three lissamphibian lineages, lizards, multiple crocodylians, and several new mammal occurrences. The known Cloverly vertebrate fauna now more closely resembles those of other late Early Cretaceous formations in North America, indicating broad similarities across wide geographic areas at this time. In addition, this work underscores the important role VMBs can play in areas previously studied primarily through surface prospecting and quarrying, espec...

Data, whether images, measurements, counts, occurrences, or character codings, are a cornerstone of vertebrate paleontology. Every published paper, master's thesis, and doctoral dissertation relies on these data to document patterns and processes in evolution, ecology, taphonomy, geography, geologic time, and functional morphology, to name just a few. In turn, the vertebrate paleontology community relies on published data in order to reproduce and verify others' work, as well as to expand upon published analyses in new ways without having to reconstitute data sets that have been used by earlier authors and to accurately preserve data for future generations of researchers. Here, we review several databases that are of interest to vertebrate paleontologists and strongly advocate for more deposition of basic research data in publicly accessible databases by vertebrate paleontologists.

Dinosaurs provide excellent opportunities to examine the impact of sampling biases on the palaeodiversity of terrestrial organisms. The stratigraphical and geographical ranges of 847 dinosaurian species are analysed for palaeodiversity patterns and compared to several sampling metrics. The observed diversity of dinosaurs, Theropoda, Sauropodomorpha and Ornithischia, are positively correlated with sampling at global and regional scales. Sampling metrics for the same region correlate with each other, suggesting that different metrics often capture the same signal. Regional sampling metrics perform well as explanations for regional diversity patterns, but correlations with global diversity are weaker. Residual diversity estimates indicate that sauropodomorphs diversified during the Late Triassic, but major increases in the diversity of theropods and ornithischians did not occur until the Early Jurassic. Diversity increased during the Jurassic, but many groups underwent extinction during the Late Jurassic or at the Jurassic/Cretaceous boundary. Although a recovery occurred during the Cretaceous, only sauropodomorphs display a long-term upward trend. The Campanian-Maastrichtian diversity 'peak' is largely a sampling artefact. There is little evidence for a gradualistic decrease in diversity prior to the end-Cretaceous mass extinction (except for ornithischians), and when such decreases do occur they are small relative to those experienced earlier in dinosaur evolution. © The Geological Society of London 2011.

Pneumatic (air-filled) postcranial bones are unique to birds among extant tetrapods. Unambiguous skeletal correlates of postcranial pneumaticity first appeared in the Late Triassic (approximately 210 million years ago), when they evolved independently in several groups of bird-line archosaurs (ornithodirans). These include the theropod dinosaurs (of which birds are extant representatives), the pterosaurs, and sauropodomorph dinosaurs. Postulated functions of skeletal pneumatisation include weight reduction in large-bodied or flying taxa, and density reduction resulting in energetic savings during foraging and locomotion. However, the influence of these hypotheses on the early evolution of pneumaticity has not been studied in detail previously. We review recent work on the significance of pneumaticity for understanding the biology of extinct ornithodirans, and present detailed new data on the proportion of the skeleton that was pneumatised in 131 non-avian theropods and Archaeopteryx. This includes all taxa known from significant postcranial remains. Pneumaticity of the cervical and anterior dorsal vertebrae occurred early in theropod evolution. This ‘common pattern’ was conserved on the line leading to birds, and is likely present in Archaeopteryx. Increases in skeletal pneumaticity occurred independently in as many as 12 lineages, highlighting a remarkably high number of parallel acquisitions of a bird-like feature among non-avian theropods. Using a quantitative comparative framework, we show that evolutionary increases in skeletal pneumaticity are significantly concentrated in lineages with large body size, suggesting that mass reduction in response to gravitational constraints at large body sizes influenced the early evolution of pneumaticity. However, the body size threshold for extensive pneumatisation is lower in theropod lineages more closely related to birds (maniraptorans). Thus, relaxation of the relationship between body size and pneumatisation preceded the origin of birds and cannot be explained as an adaptation for flight. We hypothesise that skeletal density modulation in small, non-volant, maniraptorans resulted in energetic savings as part of a multi-system response to increased metabolic demands. Acquisition of extensive postcranial pneumaticity in small-bodied maniraptorans may indicate avian-like high-performance endothermy.

Non-avian theropod dinosaurs attained large body sizes, monopolising terrestrial apex predator niches in the Jurassic-Cretaceous. From the Middle Jurassic onwards, Allosauroidea and Megalosauroidea comprised almost all large-bodied predators for 85 million years. Despite their enormous success, however, they are usually considered absent from terminal Cretaceous ecosystems, replaced by tyrannosaurids and abelisaurids. We demonstrate that the problematic allosauroids Aerosteon, Australovenator, Fukuiraptor and Neovenator form a previously unrecognised but ecologically diverse and globally distributed clade (Neovenatoridae, new clade) with the hitherto enigmatic theropods Chilantaisaurus, Megaraptor and the Maastrichtian Orkoraptor. This refutes the notion that allosauroid extinction pre-dated the end of the Mesozoic. Neovenatoridae includes a derived group (Megaraptora, new clade) that developed long, raptorial forelimbs, cursorial hind limbs, appendicular pneumaticity and small size, features acquired convergently in bird-line theropods. Neovenatorids thus occupied a 14-fold adult size range from 175 kg (Fukuiraptor) to approximately 2,500 kg (Chilantaisaurus). Recognition of this major allosauroid radiation has implications for Gondwanan paleobiogeography: The distribution of early Cretaceous allosauroids does not strongly support the vicariant hypothesis of southern dinosaur evolution or any particular continental breakup sequence or dispersal scenario. Instead, clades were nearly cosmopolitan in their early history, and later distributions are explained by sampling failure or local extinction.

Vertebrate microfossil assemblages in terrestrial formations are a promising source of data on the structure of fossil metacommunities. However, the degree to which these deposits capture true, metacommunity-level samples is unknown. Individual deposits may be biased in ways that limit their utility for intra- and inter-formation comparisons. This study describes the composition of ten vertebrate microfossil assemblages collected from the Lower Cretaceous Cloverly Formation in Montana and Wyoming, U.S.A., and evaluates whether the assemblages are sufficiently similar to suggest the presence of a single Cloverly metacommunity, or ‘paleocommunity type.’ The assemblages appear to be biased by factors related to the preferential incorporation and preservation of different taxa and skeletal element types, which compound with decreasing locality sample size. Less productive localities lack whole taxonomic groups, especially small, fully terrestrial vertebrates. Only the two vertebrate microfossil bonebeds approach the known formation richness. High individual tooth counts overrepresent particular species, especially the crocodylians. Despite these biases, the multiple assemblages nevertheless yield statistically similar taxon abundance rank orders, suggesting the presence of a single, formation-wide paleo-metacommunity, which bears resemblance to a viable metacommunity. The aggregate assemblage exhibits an ‘Eltonian pyramid’ trophic hierarchy for both terrestrial and aquatic taxa. Comparing the multiple assemblages synoptically also reveals possible landscape-scale abundance patterns for particular species. Taken together, microvertebrate assemblages offer insight into regional paleo-metacommunities and provide samples for inter-formational comparisons at this level. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Carrano, M. T., M. P. J. Oreska, and R. Lockwood. 2016. Vertebrate paleontology of the Cloverly Formation (Lower Cretaceous), II: Paleoecology. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2015.1071265.

Abelisaurid theropods are common members of Cretaceous Gondwanan faunas and are characterized by a bizarre, highly reduced forelimb. Unfortunately, forelimb elements are rarely preserved and thus the basic structure of the abelisaurid forelimb remains poorly understood. Until recently, the Upper Cretaceous Maevarano Formation of northwestern Madagascar has produced numerous exceptional specimens of the abelisaurid theropod Majungasaurus crenatissimus but comparatively little forelimb material. A recently discovered articulated skeleton of Majungasaurus preserves a virtually complete pectoral girdle and forelimb, which, along with additional isolated forelimb elements, affords important new insights into the structure of these elements. New specimens of the scapulocoracoid and humerus allow more detailed description of their morphology, and antebrachial and manual elements are described for the first time. The radius and ulna are approximately one-quarter the length of the humerus and both have ex...

Aim Modern biodiversity peaks in the tropics and declines poleward, a pattern that is potentially driven by climate. Although this latitudinal biodiversity gradient (LBG) also characterizes the marine invertebrate fossil record, distributions of ancient terrestrial faunas are poorly understood. This study utilizes data on the dinosaur fossil record to examine spatial patterns in terrestrial biodiversity throughout the Mesozoic. Location We compiled data on fossil occurrences across the globe. Methods We compiled a comprehensive dataset of Mesozoic dinosaur genera (738), including birds. Following the utilization of sampling standardization techniques to mediate for the uneven sampling of the fossil record, we constructed latitudinal patterns of biodiversity from this dataset. Results The dominant group of Mesozoic terrestrial vertebrates did not conform to the modern LBG. Instead, dinosaur diversity was highest at temperate palaeolatitudes throughout the 160 million year span of dinosaurian evolutionary history. Latitudinal diversity correlates strongly with the distribution of land area. Late Cretaceous sauropods and ornithischians exhibit disparate LBGs. Main conclusions The continuity of the palaeotemperate peak in dinosaur diversity indicates a diminished role for climate on the Mesozoic LBG; instead, dinosaur diversity may have been driven by the amount of land area among latitudinal belts. There is no evidence that the tropics acted as a cradle for dinosaur diversity. Geographical partitioning among major clades of herbivorous dinosaurs in the Late Cretaceous may result from the advanced stages of continental fragmentation and/or differing responses to increasing latitudinal climatic zonation. Our results suggest that the modern-day LBG on land was only established 30 million years ago, following a significant post-Eocene recalibration, potentially related to increased seasonality.

The Cretaceous dinosaur fauna of Indo-Pakistan has remained poorly understood because of a lack of asso- ciated and articulated remains, proliferation of named spe- cies, and an incomplete understanding of the dinosaur clades present (e.g. abelisaurid theropods; titanosaur sauro- pods). Continued work on existing collections, and new discoveries of dinosaur material from India, Pakistan and elsewhere in Gondwana, has begun to resolve the composi- tion and affinities of Indo-Pakistani dinosaurs. Here, we provide archival evidence that documents associations between postcranial remains of a sauropod collected from Chhota Simla, India by C. A. Matley in the 1930s and later described as 'Titanosaurus sp.' This partial skeleton, which represents only the fifth such documented association from Indo-Pakistan, is referable to Jainosaurus cf. septentrionalis and provides a fuller understanding of its anatomy and phylogenetic affinities.

Smithsonian Contributions to Paleobiology, no. 95: viii + 53 pages, 26 figures, 3 tables. New materials of Masiakasaurus knopfleri Sampson, Carrano, and Forster, 2001, and implications for the morphology of the Noasauridae (Theropoda: Ceratosauria).

The fossil record is our primary window onto the diversification of ancient life, but there are widespread concerns that sampling biases may distort observed palaeodiversity counts. Such concerns have been reinforced by numerous studies that found correlations between measures of sampling intensity and observed diversity. However, correlation does not necessarily mean that sampling controls observed diversity: an alternative view is that both sampling and diversity may be driven by some common factor (e.g. variation in continental flooding driven by sea level). The latter is known as the ‘common cause’ hypothesis. Here, we present quantitative analyses of the relationships between dinosaur diversity, sampling of the dinosaur fossil record, and changes in continental flooding and sea level, providing new insights into terrestrial common cause. Although raw data show significant correlations between continental flooding/sea level and both observed diversity and sampling, these correlations do not survive detrending or removal of short-term autocorrelation. By contrast, the strong correlation between diversity and sampling is robust to various data transformations. Correlations between continental flooding/sea level and taxic diversity/sampling result from a shared upward trend in all data series, and short-term changes in continental flooding/sea level and diversity/sampling do not correlate. The hypothesis that global dinosaur diversity is tied to sea-level fluctuations is poorly supported, and terrestrial common cause is unsubstantiated as currently conceived. Instead, we consider variation in sampling to be the preferred null hypothesis for short-term diversity variation in the Mesozoic terrestrial realm.

Synopsis Recent discoveries and analyses have drawn increased attention to Ceratosauria, a taxonomically and morphologically diverse group of basal theropods. By the time of its first appearance in the Late Jurassic, the group was probably globally distributed. This pattern eventually gave way to a primarily Gondwanan distribution by the Late Cretaceous. Ceratosaurs are one of several focal groups for studies of Cretaceous palaeobiogeography and their often bizarre morphological developments highlight their distinctiveness. Unfortunately, lack of phylogenetic resolution, shifting views of which taxa fall within Ceratosauria and minimal overlap in coverage between systematic studies, have made it difficult to explicate any of these important evolutionary patterns. Although many taxa are fragmentary, an increase in new, more complete forms has clarified much of ceratosaur anatomy, allowed the identification of additional materials and increased our ability to compare specimens and taxa. We studied nearly 40...

We review the historical sequence of discoveries of Majungasaurus crenatissimus, an abelisaurid theropod from the Upper Cretaceous (Maastrichtian) Maevarano Formation, Mahajanga Basin, northwestern Madagascar. For almost a century, beginning in 1895, periodic expeditions conducted by French, Japanese, and Malagasy teams yielded fragmentary and isolated remains of a medium-sized theropod dinosaur from the Maevarano Formation. These materials were first assigned to Megalosaurus crenatissimus but later to Dryptosaurus crenatissimus, then Majungasaurus crenatissimus, and, most recently, Majungatholus atopus. The taxon was variously considered to pertain to Tyrannosauridae, “Megalosauridae,” and Abelisauridae. Recent excavations undertaken by the Mahajanga Basin Project (1993 to present) have resulted in abundant and spectacular remains of this theropod. The new material allows us to establish Majungasaurus crenatissimus as the valid name for this theropod and to conclusively place it within Abelisaur...

The appendicular skeleton of the abelisaurid theropod Majungasaurus crenatissimus (Deperet, 1896) Lavocat, 1955 is described for the first time. The available materials include an incomplete pectoral girdle and forelimb, along with the ilium and a nearly complete hind limb. These materials display a number of ceratosaur, abelisauroid, and abelisaurid synapomorphies, supporting the phylogenetic placement of Majungasaurus based previously on cranial anatomy. As in Ceratosaurus and Carnotaurus, the scapular blade is relatively wide and has a pronounced dorsal lip over the glenoid. The humerus is short and bears a globular head, but is more slender than in Carnotaurus. The ilium has a preacetabular hook, a strong supraacetabular crest, a notched posterior margin, and peg-and-socket articulations with both the pubis and ischium. Hind limb elements are proportionally stocky, as in some other abelisaurids. The femur lacks a trochanteric shelf, the tibia has a greatly enlarged cnemial crest, and the fibu...

Fossils provide the principal basis for temporal calibrations, which are critical to the accuracy of divergence dating analyses. Translating fossil data into minimum and maximum bounds for calibrations is the most important, and often least appreciated, step of divergence dating. Properly justified calibrations require the synthesis of phylogenetic, paleontological, and geological evidence and can be difficult for non- specialists to formulate. The dynamic nature of the fossil record (e.g., new discoveries, taxonomic revisions, updates of global or local stratigraphy) requires that calibration data be updated continually lest they become obsolete. Here, we announce the Fossil Calibration Database (http://fossilcalibrations.org), a new open- access resource providing vetted fossil calibrations to the scientific community. Calibrations accessioned into this database are based on individual fossil specimens and follow best practices for phylogenetic justification and geochronological constraint. The associated Fossil Calibration Series, a calibration-themed publication series at Palaeontologia Electronica, will serve as one key pipeline for peer-reviewed calibrations to enter the database.

Hypotheses suggest that structural integrity of vertebrate bones is maintained by controlling bone strain magnitude via adaptive modelling in response to mechanical stimuli. Increased tissue-level strain magnitude and rate have both been identified as potent stimuli leading to increased bone formation. Mechanotransduction models hypothesize that osteocytes sense bone deformation by detecting fluid flow-induced drag in the bone's lacunar–canalicular porosity. This model suggests that the osteocyte's intracellular response depends on fluid-flow rate, a product of bone strain rate and gradient, but does not provide a mechanism for detection of strain magnitude. Such a mechanism is necessary for bone modelling to adapt to loads, because strain magnitude is an important determinant of skeletal fracture. Using strain gauge data from the limb bones of amphibians, reptiles, birds and mammals, we identified strong correlations between strain rate and magnitude across clades employing diverse locomotor styles and degrees of rhythmicity. The breadth of our sample suggests that this pattern is likely to be a common feature of tetrapod bone loading. Moreover, finding that bone strain magnitude is encoded in strain rate at the tissue level is consistent with the hypothesis that it might be encoded in fluid-flow rate at the cellular level, facilitating bone adaptation via mechanotransduction.

The fossil record of crocodylians and their relatives (pseudosuchians) reveals a rich evolutionary history, prompting questions about causes of long-term decline to their present-day low biodiversity. We analyse climatic drivers of subsampled pseudosuchian biodiversity over their 250 million year history, using a comprehensive new data set. Biodiversity and environmental changes correlate strongly, with long-term decline of terrestrial taxa driven by decreasing temperatures in northern temperate regions, and biodiversity decreases at lower latitudes matching patterns of increasing aridification. However, there is no relationship between temperature and biodiversity for marine pseudosuchians, with sea-level change and post-extinction opportunism demonstrated to be more important drivers. A ‘modern-type' latitudinal biodiversity gradient might have existed throughout pseudosuchian history, and range expansion towards the poles occurred during warm intervals. Although their fossil record suggests that current global warming might promote long-term increases in crocodylian biodiversity and geographic range, the 'balancing forces' of anthropogenic environmental degradation complicate future predictions., Crocodylians and their relatives have a rich evolutionary history. Here the authors show long-term decline of terrestrial crocodylians driven by decreasing temperatures but no relationship between temperature and biodiversity for marine crocodylians over their 250 million year history.

Quarry 9 is among the richest microvertebrate localities in the Morrison Formation, having thus far produced the remains of dozens of Late Jurassic taxa. Because this lenticular claystone deposit records such a high diversity of contemporaneous species, it provides an exceptionally detailed view of their paleoecology and local paleoenvironment. In this study, we reexamined the entire Quarry 9 collection, totaling more than 3700 specimens, and developed a revised faunal list that was used to determine taxonomic and ecological diversities. Comprehensive abundance data were collected as well, revealing significant discrepancies between the most diverse and most abundant groups. Amphibious taxa (crocodilians and turtles) were very abundant, and seemed to fill an important ecological role as “connectors” between the terrestrial and aquatic food webs. In contrast, small terrestrial taxa (small theropods, mammals, and small reptiles) were very diverse, highlighting their central placement within the terrestrial food web. Lithologically and sedimentologically, the deposition of Quarry 9 occurred in a low-energy pond or swamp, an interpretation supported by the available taphonomic data.

Here we redescribe the holotype and only specimen of Segisaurus halli, a small Early Jurassic dinosaur and the only theropod known from the Navajo Sandstone. Our study highlights several important ...

This paper investigates trends in the evolution of body size and shape in the Plesiosauria, a diverse clade of Mesozoic marine reptiles. Using measures from well-preserved plesiosaur specimens, we document and interpret evolutionary patterns in relative head size, body size, and locomotor variables. Size increase is a significant trend in the clade as a whole, and in constituent clades. The trend in relative head size is of variance increase; observed head sizes are both smaller and larger than ancestral values. In the locomotor system, changes in propodial and girdle proportions appear concomitant with body size increase and are interpreted as allometric responses to the physical constraints of large body size. Other trends in the locomotor system are significantly correlated with both body size and relative head size. These locomotor trends evolved convergently in several clades of plesiosaurs, and may have had an ecomorphological basis, although data are lacking to constrain speculation on this point. The evolution of the locomotor system in plesiosaurs sheds new light on the response of aquatic tetrapods to the physical constraints of foraging at large body size.

We describe the osteology of the new small theropod dinosaur Masiakasaurus knopfleri, from the Late Cretaceous Maevarano Formation of northwestern Madagascar. Approximately 40% of the skeleton is known, including parts of the jaws, axial column, forelimb, pelvic girdle, and hind limb. The jaws of Masiakasaurus are remarkably derived, bearing a heterodont, procumbent dentition that is unknown elsewhere among dinosaurs. The vertebrae are similar to those of abelisauroids in the reduction of the neural spine, lack of pleurocoelous fossae on the centrum, and extensively pneumatized neural arch. The limb skeleton is relatively gracile and bears numerous abelisauroid synapomorphies, including a rounded humeral head, peg-and-socket iliac-pubic articulation, prominent femoral medial epicondyle, expanded tibial cnemial crest, and double-grooved pedal unguals. The femora and tibiae show evidence of dimorphism. More specific features shared between Masiakasaurus, the Argentine Noasaurus, and the Indian Laev...

Current theories of locomotor biomechanics are based largely on observations of extant terrestrial mammals. However, extant mammals are limited with respect to certain aspects of terrestrial locomotion, and this constraint has confounded interpretations of limb design in this group. Dinosaurs include a wide array of large and small bipeds, record several transitions between bipedalism and quadrupedalism, and show a unique covariation of cursoriality and body size. Thus, the wider applications of biomechanical theories may be tested by applying them to the limb bones of dinosaurs. A broad examination of hindlimb and forelimb bone scaling patterns in dinosaurs and mammals reveals several general similarities that provide insight into the general constraints acting on terrestrial locomotion, particularly in animals with parasagittally oriented limbs. Most limb bones scale with negative allometry in these two groups, and larger taxa tend to scale more negatively than smaller forms. However, the strongly negative scaling of large mammals is mostly restricted to ungulates, whose unusually short femora may be the result of constraints of cursoriality at large body sizes. Bipedal and quadrupedal dinosaurs scale very similarly, with most differences apparently resulting from size rather than posture. Bone curvature tends to decrease with increasing body size, while femoral midshaft eccentricity tends to increase. Femoral midshaft eccentricity is explained as a general adaptation to mediolateral bending on parasagittal limb bones. These trends are more pronounced in dinosaurs than mammals; additional morphological constraints present in the dinosaurian hindlimb may contribute to this distinction.

(2001). A new fossil frog from the Upper Cretaceous Judith River Formation of Montana. Journal of Vertebrate Paleontology: Vol. 21, No. 1, pp. 190-194.

Vertebrate tracks are a unique, abundant source of fossil data that supplements the skeletal record in many ways. However, the utility of ichnofossil data depends on how specifically the authors of tracks can be identified. Despite this fact, there is little consensus about how to identify potential trackmakers, and existing methods differ in their bases, assumptions, and corresponding implications.In this paper we support the proposal that trackmakers should be identified primarily by skeletal structures that are both preserved in the ichnofossils and synapomorphies of some body-fossil clade. This synapomorphy-based technique enables certain taxa to be positively identified as candidate trackmakers and others to be excluded from consideration. In addition, the diagnostic level of the synapomorphy (i.e., to a higher or lower level) corresponds to that of the trackmaker. Additional features, such as body size and provenance, can be used in association with synapomorphies as additional differentiae of trackmaker identity.Trackway analyses are dependent on the level of trackmaker diagnosis, but not all analyses require the same diagnostic specificity. Palichnostratigraphic correlations to the stage level are shown to require at least a genus-level identification of a trackmaker, whereas studies of vertebrate distributions (i.e., origins, extinctions, ranges) accommodate much coarser designations. Anachronistic occurrences of trace and body fossils result in range extensions for either the skeletal taxon or the feature in question. For example, the temporal distribution of theropods can be extended on the basis of the footprint record, resulting in an earlier estimated divergence time for Dinosauria.

In this paper, I survey hindlimb and pelvic anatomy across non-avian dinosaurs and analyze these within a cladistic framework to quantify patterns of change within the locomotor apparatus. Specifically, I attempt to identify where homoplasy constitutes parallelism and may thereby be used to infer similar selective pressures on hindlimb function. Traditional methods of discrete character optimization are used along with two methods for evaluating changes in continuous characters in a phylogenetic context (squared-change parsimony and clade rank correlation). Resultant patterns are evaluated in light of the biomechanics of locomotion and the relationship between form and function in extant terrestrial vertebrates.Although non-avian dinosaurian locomotor morphology is strikingly uniform, these analyses reveal the repeated derivations of several morphological features that have potential relevance for hindlimb locomotor function. Anterior and posterior iliac expansion, a medially oriented femoral head, and an elevated femoral lesser trochanter each evolved independently multiple times within Dinosauria. These changes probably reflect enlargement of several hindlimb muscles as well as a general switch in their predominant function from abduction-adduction (characteristic of “sprawling” limb postures) to protraction-retraction (characteristic of parasagittal, or “erect,” limb postures). Several “avian” characteristics are shared with more basal theropods, and many were acquired convergently in other dinosaurian lineages. The evolution of the avian hindlimb therefore represents a cumulative acquisition of characters, many of which were quite far removed in time and function from the origin of flight.

Theropod hind limb disparity revisited: comments on Gatesy and Middleton (1997) Matthew T. Carrano a b & Christian A. Sidor a a Department of Organismal Biology and Anatomy , University of Chicago , 1027 East 57th Street, Chicago, Illinois, 60637 b Department of Anatomical Sciences , Health Sciences Center T-8, State University of New York at Stony Brook , Stony Brook, New York, 11794 Published online: 24 Aug 2010.

Extant birds represent the only diverse living bipeds, and can be informative for investigations into the life-history parameters of their extinct dinosaurian relatives. However, morphological changes that occurred during early avian evolution, including the unique adoption of a nearly horizontal femoral orientation associated with a shift in center of mass (CM), suggest that caution is warranted in the use of birds as analogs for nonavian dinosaur locomotion. In this study, we fitted a group of white leghorn chickens (Gallus gallus) with a weight suspended posterior to the hip in order to examine the effects on loading and morphology. This caused a CM shift that necessitated a change in femoral posture (by 35 degrees towards the horizontal, P < 0.001), and resulted in reorientation of the ground reaction force (GRF) vector relative to the femur (from 41 degrees to 82 degrees, P < 0.001). Despite similar strain magnitudes, an overall increase in torsion relative to bending (from 1.70 to 1.95 times bending, P < 0.001) was observed, which was weakly associated with a tendency for increased femoral cross-sectional dimensions (P = 0.1). We suggest that a relative increase in torsion is consistent with a change in femoral posture towards the horizontal, since this change increases the degree to which the bone axis and the GRF vector produce mediolateral long-axis rotation of the bone. These results support the hypothesis that a postural change during early avian evolution could underlie the allometric differences seen between bird and nonavian dinosaur femora by requiring more robust femoral dimensions in birds due to an increase in torsion.