The osteology of the Late Triassic reptile Scleromochlus taylori from µCT data.

Scleromochlus taylori is one of the most enigmatic members of the herpetofauna from the Lossiemouth Sandstone Formation (Upper Triassic) of Elgin (Moray, Scotland). For many years it was thought to be closely related to pterosaurs and dinosaurs, but the anatomy of this animal is difficult to interpret because of the notoriously poor preservation of the six available specimens, which comprise void space in the sandstone after the bones were destroyed by diagenesis. Historically, these fossils have been studied using physical molds, which provide only incomplete, and potentially distorted, information. Due to these uncertainties, interpretations of the anatomy, phylogenetic relationships, and paleobiology of Scleromochlus taylori have remained contentious. Here, we use microcomputed tomographic (µCT) techniques to redescribe and illustrate the osteology of Scleromochlus in detail, building upon a short redescription of keystone features of the anatomy that we recently published. We digitally visualize, describe, and figure previously inaccessible-and thus unaltered-portions of its skeleton, as well as providing new observations on the exposed parts of each specimen. This work reveals many novel features of the skull, mandible, trunk, tail, girdles, forelimb, and hindlimb (particularly of the manus, femur, and pes), demonstrating that historic molding techniques failed, in some cases, to accurately capture the anatomy of Scleromochlus. Our review sheds light on some of the most controversial aspects of Scleromochlus morphology showing that this taxon retains plesiomorphic features of Avemetatarsalia in the postcranial skeleton, alongside a suite of synapomorphies diagnostic of pterosauromorphs (the broad clade of pterosaurs and taxa more closely related to them than dinosaurs), particularly one subgroup, the lagerpetids. Consistent with recent work, our updated phylogenetic analyses (Maximum Parsimony and Bayesian Inference) demonstrate that Scleromochlus taylori is an avemetatarsalian archosaur that is recovered firmly in an early diverging position within Pterosauromorpha, as a member of Lagerpetidae, thus shedding important information on the origin of pterosaurs, the first group of vertebrates to evolve powered flight.

The biogeographic history of neosuchian crocodiles and the impact of saltwater tolerance variability.

Extant neosuchian crocodiles are represented by only 24 taxa that are confined to the tropics and subtropics. However, at other intervals during their 200 Myr evolutionary history the clade reached considerably higher levels of species-richness, matched by more widespread distributions. Neosuchians have occupied numerous habitats and niches, ranging from dwarf riverine forms to large marine predators. Despite numerous previous studies, several unsolved questions remain with respect to their biogeographic history, including the geographical origins of major groups, e.g. Eusuchia and Neosuchia itself. We carried out the most comprehensive biogeographic analysis of Neosuchia to date, based on a multivariate K-means clustering approach followed by the application of two ancestral area estimation methods (BioGeoBEARS and Bayesian ancestral location estimation) applied to two recently published phylogenies. Our results place the origin of Neosuchia in northwestern Pangaea, with subsequent radiations into Gondwana. Eusuchia probably emerged in the European archipelago during the Late Jurassic/Early Cretaceous, followed by dispersals to the North American and Asian landmasses. We show that putative transoceanic dispersal events are statistically significantly less likely to happen in alligatoroids. This finding is consistent with the saltwater intolerant physiology of extant alligatoroids, bolstering inferences of such intolerance in their ancestral lineages.

Origins of slow growth on the crocodilian stem lineage.

Crocodilians grow slowly and have low metabolic rates similar to other living reptiles, but palaeohistology indicates that they evolved from an ancestor with higher growth rates.1,2,3,4,5 It remains unclear when slow growth appeared in the clade due to the sparse data on key divergences among early Mesozoic members of their stem lineage. We present new osteohistological data from a broad sample of early crocodylomorphs, evaluated in a phylogenetic context alongside other pseudosuchians. We find that the transition to slow-growing bone types during mid-late ontogeny occurred around the origin of Crocodylomorpha during the Late Triassic. Earlier-diverging pseudosuchians had high maximum growth rates, as indicated by the presence of woven bone during middle and (sometimes) late ontogeny.6,7,8,9 Large-bodied pseudosuchians in particular exhibit some of the fastest-growing bone types, giving evidence for prolonged, rapid growth. By contrast, early-branching crocodylomorphs, including a new large-bodied taxon, had slow maximum rates of bone deposition, as evidenced by the presence of predominantly parallel-fibered or lamellar bone tissue during middle-late ontogeny. Late Triassic crocodylomorphs show skeletal anatomy consistent with "active" terrestrial habits,10,11,12 and their slow growth rates reject hypotheses linking this transition with sedentary, semiaquatic lifestyles or sprawling posture. Faster-growing pseudosuchian lineages go extinct in the Triassic, whereas slow-growing crocodylomorphs do not. This contrasts with the Jurassic radiation of fast-growing dinosaurs on the bird-stem lineage,13 suggesting that the End-Triassic mass extinction initiated a divergent distribution of growth strategies that persist in present-day archosaurs.

Multiple pathways to herbivory underpinned deep divergences in ornithischian evolution.

The extent to which evolution is deterministic is a key question in biology,1,2,3,4,5,6,7,8,9 with intensive debate on how adaptation6,10,11,12,13 and constraints14,15,16 might canalize solutions to ecological challenges.4,5,6 Alternatively, unique adaptations1,9,17 and phylogenetic contingency1,3,18 may render evolution fundamentally unpredictable.3 Information from the fossil record is critical to this debate,1,2,11 but performance data for extinct taxa are limited.7 This knowledge gap is significant, as general morphology may be a poor predictor of biomechanical performance.17,19,20 High-fiber herbivory originated multiple times within ornithischian dinosaurs,21 making them an ideal clade for investigating evolutionary responses to similar ecological pressures.22 However, previous biomechanical modeling studies on ornithischian crania17,23,24,25 have not compared early-diverging taxa spanning independent acquisitions of herbivory. Here, we perform finite-element analysis on the skull of five early-diverging members of the major ornithischian clades to characterize morphofunctional pathways to herbivory. Results reveal limited functional convergence among ornithischian clades, with each instead achieving comparable performance, in terms of reconstructed patterns and magnitudes of functionally induced stress, through different adaptations of the feeding apparatus. Thyreophorans compensated for plesiomorphic low performance through increased absolute size, heterodontosaurids expanded jaw adductor muscle volume, ornithopods increased jaw system efficiency, and ceratopsians combined these approaches. These distinct solutions to the challenges of herbivory within Ornithischia underpinned the success of this diverse clade. Furthermore, the resolution of multiple solutions to equivalent problems within a single clade through macroevolutionary time demonstrates that phenotypic evolution is not necessarily predictable, instead arising from the interplay of adaptation, innovation, contingency, and constraints.1,2,3,7,8,9,18.

100 million years of turtle paleoniche dynamics enable the prediction of latitudinal range shifts in a warming world.

Past responses to environmental change provide vital baseline data for estimating the potential resilience of extant taxa to future change. Here, we investigate the latitudinal range contraction that terrestrial and freshwater turtles (Testudinata) experienced from the Late Cretaceous to the Paleogene (100.5-23.03 mya) in response to major climatic changes. We apply ecological niche modeling (ENM) to reconstruct turtle niches, using ancient and modern distribution data, paleogeographic reconstructions, and the HadCM3L climate model to quantify their range shifts in the Cretaceous and late Eocene. We then use the insights provided by these models to infer their probable ecological responses to future climate scenarios at different representative concentration pathways (RCPs 4.5 and 8.5 for 2100), which project globally increased temperatures and spreading arid biomes at lower to mid-latitudes. We show that turtle ranges are predicted to expand poleward in the Northern Hemisphere, with decreased habitat suitability at lower latitudes, inverting a trend of latitudinal range contraction that has been prevalent since the Eocene. Trionychids and freshwater turtles can more easily track their niches than Testudinidae and other terrestrial groups. However, habitat destruction and fragmentation at higher latitudes will probably reduce the capability of turtles and tortoises to cope with future climate changes.

Multi-proxy dentition analyses reveal niche partitioning between sympatric herbivorous dinosaurs.

Dentitions of the sympatric herbivorous dinosaurs Hungarosaurus (Ankylosauria, Nodosauridae) and Mochlodon (Ornithopoda, Rhabdodontidae) (Santonian, Hungary) were analysed to investigate their dietary ecology, using several complementary methods-orientation patch count, tooth replacement rate, macrowear, tooth wear rate, traditional microwear, and dental microwear texture analysis (DMTA). Tooth formation time is similar in Hungarosaurus and Mochlodon, and traditional and DMTA microwear features suggest low-browsing habits for both taxa, consistent with their inferred stances and body sizes. However, Mochlodon possesses a novel adaptation for increasing dental durability: the dentine on the working side of the crown is double the thickness of that on the balancing side. Moreover, crown morphology, enamel thickness, macrowear orientation, and wear rate differ greatly between the two taxa. Consequently, these sympatric herbivores probably exploited plants of different toughness, implying dietary selectivity and niche partitioning. Hungarosaurus is inferred to have eaten softer vegetation, whereas Mochlodon likely fed on tougher material. Compared to the much heavier, quadrupedal Hungarosaurus, the bipedal Mochlodon wore down more than twice as much of its crown volume during the functional life of the tooth. This heavy tooth wear might correlate with more intensive food processing and, in turn, could reflect differences in the metabolic requirements of these animals.

Scleromochlus and the early evolution of Pterosauromorpha.

Pterosaurs, the first vertebrates to evolve powered flight, were key components of Mesozoic terrestrial ecosystems from their sudden appearance in the Late Triassic until their demise at the end of the Cretaceous1-6. However, the origin and early evolution of pterosaurs are poorly understood owing to a substantial stratigraphic and morphological gap between these reptiles and their closest relatives6, Lagerpetidae7. Scleromochlus taylori, a tiny reptile from the early Late Triassic of Scotland discovered over a century ago, was hypothesized to be a key taxon closely related to pterosaurs8, but its poor preservation has limited previous studies and resulted in controversy over its phylogenetic position, with some even doubting its identification as an archosaur9. Here we use microcomputed tomographic scans to provide the first accurate whole-skeletal reconstruction and a revised diagnosis of Scleromochlus, revealing new anatomical details that conclusively identify it as a close pterosaur relative1 within Pterosauromorpha (the lagerpetid + pterosaur clade). Scleromochlus is anatomically more similar to lagerpetids than to pterosaurs and retains numerous features that were probably present in very early diverging members of Avemetatarsalia (bird-line archosaurs). These results support the hypothesis that the first flying reptiles evolved from tiny, probably facultatively bipedal, cursorial ancestors1.

Interelemental osteohistological variation in Massospondylus carinatus and its implications for locomotion.

Massospondylus carinatus Owen, 1854 is an iconic basal sauropodomorph dinosaur from the Early Jurassic of southern Africa. Over 200 specimens have been referred to this taxon, spanning the entire ontogenetic series from embryo to adult. Consequently, it provides an ideal sample for investigating dinosaur developmental biology, including growth patterns and growth rates, through osteohistological analysis. Massospondylus carinatus was the first early-branching sauropodomorph dinosaur for which a femoral growth series was sampled. Since then, growth series of other non-avian dinosaur taxa have shown that growth plasticity, interelemental variation, and ontogenetic locomotory shifts can complicate our understanding of growth curves and patterns. To investigate these questions further, it is necessary to sample multiple skeletal elements from multiple individuals across a large range of sizes, something that is often hindered by the incompleteness of the fossil record. Here, we conducted a broad, multielement osteohistological study of long bones (excluding metapodials) from 27 specimens of Massospondylus carinatus that span its ontogenetic series. Our study reveals substantial variations in growth history. A cyclical woven-parallel complex is the predominant bone tissue pattern during early and mid-ontogeny, which transitions to slower forming parallel-fibred bone during very late ontogeny. The bone tissue is interrupted by irregularly spaced cyclical growth marks (CGMs) including lines of arrested growth indicating temporary cessations in growth. These CGMs show that the previously recorded femoral growth plasticity is also visible in other long bones, with a poor correlation between body size (measured by midshaft circumference) and CGM numbers. Furthermore, we found that the growth trajectory for an individual can vary depending on which limb element is studied. This makes the establishment of an accurate growth curve and determination of the onset of reproductive maturity difficult for this taxon. Finally, we found no evidence of differential growth rates in forelimb vs hindlimb samples from the same individual, providing further evidence falsifying hypothesised ontogenetic postural shifts in Massospondylus carinatus.

Inner ear biomechanics reveals a Late Triassic origin for mammalian endothermy.

Endothermy underpins the ecological dominance of mammals and birds in diverse environmental settings1,2. However, it is unclear when this crucial feature emerged during mammalian evolutionary history, as most of the fossil evidence is ambiguous3-17. Here we show that this key evolutionary transition can be investigated using the morphology of the endolymph-filled semicircular ducts of the inner ear, which monitor head rotations and are essential for motor coordination, navigation and spatial awareness18-22. Increased body temperatures during the ectotherm-endotherm transition of mammal ancestors would decrease endolymph viscosity, negatively affecting semicircular duct biomechanics23,24, while simultaneously increasing behavioural activity25,26 probably required improved performance27. Morphological changes to the membranous ducts and enclosing bony canals would have been necessary to maintain optimal functionality during this transition. To track these morphofunctional changes in 56 extinct synapsid species, we developed the thermo-motility index, a proxy based on bony canal morphology. The results suggest that endothermy evolved abruptly during the Late Triassic period in Mammaliamorpha, correlated with a sharp increase in body temperature (5-9 °C) and an expansion of aerobic and anaerobic capacities. Contrary to previous suggestions3-14, all stem mammaliamorphs were most probably ectotherms. Endothermy, as a crucial physiological characteristic, joins other distinctive mammalian features that arose during this period of climatic instability28.

Unravelling the structural variation of lizard osteoderms.

Vertebrate skin is a remarkable organ that supports and protects the body. It consists of two layers, the epidermis and the underlying dermis. In some tetrapods, the dermis includes mineralised organs known as osteoderms (OD). Lizards, with over 7,000 species, show the greatest diversity in OD morphology and distribution, yet we barely understand what drives this diversity. This multiscale analysis of five species of lizards, whose lineages diverged ∼100-150 million years ago, compared the micro- and macrostructure, material properties, and bending rigidity of their ODs, and examined the underlying bones of the skull roof and jaw (including teeth when possible). Unsurprisingly, OD shape, taken alone, impacts bending rigidity, with the ODs of Corucia zebrata being most flexible and those of Timon lepidus being most rigid. Macroscopic variation is also reflected in microstructural diversity, with differences in tissue composition and arrangement. However, the properties of the core bony tissues, in both ODs and cranial bones, were found to be similar across taxa, although the hard, capping tissue on the ODs of Heloderma and Pseudopus had material properties similar to those of tooth enamel. The results offer evidence on the functional adaptations of cranial ODs, but questions remain regarding the factors driving their diversity. STATEMENT OF SIGNIFICANCE: Understanding nature has always been a significant source of inspiration for various areas of the physical and biological sciences. Here we unravelled a novel biomineralization, i.e. calcified tissue, OD, forming within the skin of lizards which show significant diversity across the group. A range of techniques were used to provide an insight into these exceptionally diverse natural structures, in an integrated, whole system fashion. Our results offer some suggestions into the functional and biomechanical adaptations of OD and their hierarchical structure. This knowledge can provide a potential source of inspiration for biomimetic and bioinspired designs, applicable to the manufacturing of light-weight, damage-tolerant and multifunctional materials for areas such as tissue engineering.

A new early branching armored dinosaur from the Lower Jurassic of southwestern China.

The early evolutionary history of the armored dinosaurs (Thyreophora) is obscured by their patchily distributed fossil record and by conflicting views on the relationships of Early Jurassic taxa. Here, we describe an early diverging thyreophoran from the Lower Jurassic Fengjiahe Formation of Yunnan Province, China, on the basis of an associated partial skeleton that includes skull, axial, limb, and armor elements. It can be diagnosed as a new taxon based on numerous cranial and postcranial autapomorphies and is further distinguished from all other thyreophorans by a unique combination of character states. Although the robust postcranium is similar to that of more deeply nested ankylosaurs and stegosaurs, phylogenetic analysis recovers it as either the sister taxon of Emausaurus or of the clade Scelidosaurus+ Eurypoda. This new taxon, Yuxisaurus kopchicki, represents the first valid thyreophoran dinosaur to be described from the Early Jurassic of Asia and confirms the rapid geographic spread and diversification of the clade after its first appearance in the Hettangian. Its heavy build and distinctive armor also hint at previously unrealized morphological diversity early in the clade's history.

From the plated Stegosaurus to the tank-like Ankylosaurus, armoured dinosaurs are some of the most extraordinary creatures to have roamed the earth. Fossils from this group are abundant from the Late Jurassic period, 155 million years ago, up until the end of the age of the dinosaurs. However, only a few fossils exist from the early part of the Jurassic, making it difficult to understand how these fantastic beasts came to be. More early fossils could help to fill in gaps about armoured dinosaur biology and evolution. Yao et al. describe the anatomy of a new armoured dinosaur, baptized Yuxisaurus, which was found in rocks of Early Jurassic age in southwestern China. Covered in sharp spines, this medium-sized animal was much sturdier and stockier than its immediate relatives, suggesting that the ancestors of Stegosaurus and Ankylosaurus had a wider variety of body forms than once thought. Its presence in China also shows that armoured dinosaurs spread across the world early in their history. Yuxisaurus could help researchers to understand how million years of evolution produced the armoured species we are more familiar with today. As more fossils may emerge from the rocks of southwestern China, it could become possible to further piece together early dinosaur evolution.

Cranial functional morphology of the pseudosuchian Effigia and implications for its ecological role in the Triassic.

Pseudosuchians, archosaurian reptiles more closely related to crocodylians than to birds, exhibited high morphological diversity during the Triassic with numerous examples of morphological convergence described between Triassic pseudosuchians and post-Triassic dinosaurs. One example is the shuvosaurid Effigia okeeffeae which exhibits an "ostrich-like" bauplan comprising a gracile skeleton with edentulous jaws and large orbits, similar to ornithomimid dinosaurs and extant palaeognaths. This bauplan is regarded as an adaptation for herbivory, but this hypothesis assumes morphological convergence confers functional convergence, and has received little explicit testing. Here, we restore the skull morphology of Effigia, perform myological reconstructions, and apply finite element analysis to quantitatively investigate skull function. We also perform finite element analysis on the crania of the ornithomimid dinosaur Ornithomimus edmontonicus, the extant palaeognath Struthio camelus and the extant pseudosuchian Alligator mississippiensis to assess the degree of functional convergence with a taxon that exhibit "ostrich-like" bauplans and its closest extant relatives. We find that Effigia possesses a mosaic of mechanically strong and weak features, including a weak mandible that likely restricted feeding to the anterior portion of the jaws. We find limited functional convergence with Ornithomimus and Struthio and limited evidence of phylogenetic constraints with extant pseudosuchians. We infer that Effigia was a specialist herbivore that likely fed on softer plant material, a niche unique among the study taxa and potentially among contemporaneous Triassic herbivores. This study increases the known functional diversity of pseudosuchians and highlights that superficial morphological similarity between unrelated taxa does not always imply functional and ecological convergence.

Bizarre dermal armour suggests the first African ankylosaur.

Ankylosauria is a diverse clade of armoured dinosaurs whose members were important constituents of many Cretaceous faunas. Phylogenetic analyses imply that the clade diverged from its sister taxon, Stegosauria, during the late Early Jurassic, but the fossil records of both clades are sparse until the Late Jurassic (~150 million years ago). Moreover, Ankylosauria is almost entirely restricted to former Laurasian continents, with only a single valid Gondwanan taxon. Spicomellus afer gen. et sp. nov. appears to represent the earliest-known ankylosaur and the first to be named from Africa, from the Middle Jurassic (Bathonian-Callovian) of Morocco, filling an important gap in dinosaur evolution. The specimen consists of a rib with spiked dermal armour fused to its dorsal surface, an unprecedented morphology among extinct and extant vertebrates. The specimen reveals an unrealized morphological diversity of armoured dinosaurs during their early evolution, and implies the presence of an important but undiscovered Gondwanan fossil record.

Evolution: The Two Faces of Plant-Eating Dinosaurs.

Plant-eating dinosaurs evolved varied feeding strategies. A new study demonstrates convergent evolution of their skulls and teeth towards two distinct functional optima, one resembling advanced mammalian herbivory and the other echoing herbivory in birds and other reptiles.

Dinosaur diversification rates were not in decline prior to the K-Pg boundary.

Determining the tempo and mode of non-avian dinosaur extinction is one of the most contentious issues in palaeobiology. Extensive disagreements remain over whether their extinction was catastrophic and geologically instantaneous or the culmination of long-term evolutionary trends. These conflicts have arisen due to numerous hierarchical sampling biases in the fossil record and differences in analytical methodology, with some studies identifying long-term declines in dinosaur richness prior to the Cretaceous-Palaeogene (K-Pg) boundary and others proposing continued diversification. Here, we use Bayesian phylogenetic generalized linear mixed models to assess the fit of 12 dinosaur phylogenies to three speciation models (null, slowdown to asymptote, downturn). We do not find strong support for the downturn model in our analyses, which suggests that dinosaur speciation rates were not in terminal decline prior to the K-Pg boundary and that the clade was still capable of generating new taxa. Nevertheless, we advocate caution in interpreting the results of such models, as they may not accurately reflect the complexities of the underlying data. Indeed, current phylogenetic methods may not provide the best test for hypotheses of dinosaur extinction; the collection of more dinosaur occurrence data will be essential to test these ideas further.

Reconstructing the archosaur radiation using a Middle Triassic archosauriform tooth assemblage from Tanzania.

Following the Permo-Triassic mass extinction, Archosauriformes-the clade that includes crocodylians, birds, and their extinct relatives outside crown Archosauria-rapidly diversified into many distinct lineages, became distributed globally, and, by the Late Triassic, filled a wide array of resource zones. Current scenarios of archosauriform evolution are ambiguous with respect to whether their taxonomic diversification in the Early-Middle Triassic coincided with the initial evolution of dietary specializations that were present by the Late Triassic or if their ecological disparity arose sometime after lineage diversification. Late Triassic archosauriform dietary specialization is recorded by morphological divergence from the plesiomorphic archosauriform tooth condition (laterally-compressed crowns with serrated carinae and a generally triangular lateral profile). Unfortunately, the roots of this diversification are poorly documented, with few known Early--Middle Triassic tooth assemblages, limiting characterizations of morphological diversity during this critical, early period in archosaur evolution. Recent fieldwork (2007-2017) in the Middle Triassic Manda Beds of the Ruhuhu Basin, Tanzania, recovered a tooth assemblage that provides a window into this poorly sampled interval. To investigate the taxonomic composition of that collection, we built a dataset of continuous quantitative and discrete morphological characters based on in situ teeth of known taxonomic status (e.g., Nundasuchus, Parringtonia: N = 65) and a sample of isolated teeth (N = 31). Using crown heights from known taxa to predict tooth base ratio (= base length/width), we created a quantitative morphospace for the tooth assemblage. The majority of isolated, unassigned teeth fall within a region of morphospace shared by several taxa from the Manda Beds (e.g., Nundasuchus, Parringtonia); two isolated teeth fall exclusively within a "Pallisteria" morphospace. A non-metric multidimensional scaling ordination (N = 67) of 11 binary characters reduced overlap between species. The majority of the isolated teeth from the Manda assemblage fall within the Nundasuchus morphospace. This indicates these teeth are plesiomorphic for archosauriforms as Nundasuchus exhibits the predicted plesiomorphic condition of archosauriform teeth. Our model shows that the conservative tooth morphologies of archosauriforms can be differentiated and assigned to species and/or genus, rendering the model useful for identifying isolated teeth. The large overlap in tooth shape among the species present and their overall similarity indicates that dietary specialization lagged behind species diversification in archosauriforms from the Manda Beds, a pattern predicted by Simpson's "adaptive zones" model. Although applied to a single geographic region, our methods offer a promising means to reconstruct ecological radiations and are readily transferable across a broad range of vertebrate taxa throughout Earth history.

Ngwevu intloko: a new early sauropodomorph dinosaur from the Lower Jurassic Elliot Formation of South Africa and comments on cranial ontogeny in Massospondylus carinatus.

Our knowledge of Early Jurassic palaeobiodiversity in the upper Elliot Formation of South Africa has increased markedly in recent years with the discovery of new fossils, re-assessments of previously collected material and a better understanding of Stormberg Group stratigraphy. Here, Ngwevu intloko, a new genus of upper Elliot basal sauropodomorph is named on the basis of a complete skull and partial skeleton (BP/1/4779) previously assigned to Massospondylus carinatus. It can be distinguished from all other basal sauropodomorphs by a combination of 16 cranial and six postcranial characters. The new species is compared to a small ontogenetic series of M. carinatus as well as to a range of closely related taxa. Taphonomic deformation, sexual dimorphism and ontogeny are rejected as possible explanations for the morphological differences present between BP/1/4779 and other taxa. Osteohistological examination reveals that BP/1/4779 had nearly reached adult size at the time of its death at a minimum age of 10 years.