A potentially fatal cranial pathology in a specimen of Tarchia.

Skulls of the Mongolian ankylosaurids Shamosaurus, Tarchia, and Saichania were scanned for information about their internal anatomy. Computed tomography (CT) imaging of the Tarchia skull revealed substantial internal anatomical differences from known Campanian North American taxa, particularly in the morphology of the airway. In addition, unexpected anomalies were detected within the airway and sinuses. The anomalies include multiple bilaterally distributed, variably sized hyperdense (mineralized) concretions within the airway and sinuses, the largest of which, positioned in the right nasal cavity medial to the supraorbitals, has an asymmetric ovoid shape that tapers caudally and which is partially encased within a hemispherical trabeculated osseous proliferation (sinus exostosis). Immediately adjacent to the exostosis is a subcircular transosseous defect in the prefrontal region of the skull roof that is partially filled with trabeculated ossified material with similar architectural features as the larger exostosis. Irregularities along the internal and external surfaces of the cranial vault may be associated. The radiologic features of the hemicircumferential exostosis suggest a chronic reactive osteoproliferation, possibly in response to an ongoing inflammatory reaction to primary sinus infection or, in combination with the unilateral transosseous defect, traumatically introduced infection with potentially fatal consequences. This report underscores the value of CT scanning of fossil vertebrate specimens, which in this case revealed large internal lesions of the skull that, at the time the scan was performed, were otherwise indiscernible.

Form and function in the avian pelvis.

The avian pelvis plays a critical role in the hindlimb function of birds, connecting the hindlimb and axial skeleton and serving as the major attachment site for proximal hindlimb musculature. To assess how diversification of locomotor modes in birds has impacted the evolution of avian pelvic morphology, we conducted a two-dimensional geometric morphometric analysis of bird pelves in dorsal and lateral views from 163 species (n = 261) across Aves. We investigated the relationships among pelvic shape and ecology, phylogeny, and allometry, and conducted disparity analyses to understand how pelvic morphospace has been explored through the diversification of Aves. We found that while phylogeny was correlated with shape, locomotor categories were significantly discriminated in morphospace in phylogenetically corrected analyses, as was pelvic size. Major shape trends across Aves distinguishing locomotor categories included the relative area of the preacetabular versus postacetabular ilium, how squat or narrow the pelvis is, and the extent of the caudal pelvic border. Birds adapted for hind limb-propelled swimming had particularly distinctive pelves, with narrow, elongated ilia likely useful for holding the hindlimbs close to the body midline and reducing drag. However, ecology and allometry only account for a small proportion of morphological variation, and in general locomotor groups overlapped substantially in morphospace. These results, alongside disparity through time analyses showing widespread convergence in pelvic morphology throughout the Cenozoic, suggest that avian lineages and ecotypes have extensively explored pelvic morphospace, perhaps aided by a loosening of evolutionary constraints following the evolution of forelimb-powered flight.

Alligator appendicular architecture across an ontogenetic niche shift.

A variety of species undergo ontogenetic niche shifts in either diet, habitat, or both. As a result, multiple ontogenetic stages are able to take advantage of different resources and live in sympatry without competing with one another. The American alligator (Alligator mississippiensis) begins to undergo an ontogenetic niche shift in both diet and habitat at a length of 1.2 m. They transition from a terrestrial wetland environment to a riverine environment and take advantage of different dietary resources. At 1.8 m, A. mississippiensis reaches sexual maturity. Ontogenetic shifts in habitat have the capacity to alter morphology, especially limb morphology, as different age classes traverse different ecological systems. We evaluated shape trends in the scapulae, humeri, ilia, and femora using geometric morphometrics to test whether there were punctuated changes in limb shape, shape disparity, and integration corresponding to either the ontogenetic habitat shift or onset of sexual maturity. We found size to strongly correlate with limb shape but found a continuous size gradient rather than punctuated changes in size. Furthermore, we found that adults (total length > 1.8 m) had significantly higher limb shape disparity than juveniles or subadults, likely related to ontogenetic decreases in limb use and a reduction in limb constraints. Finally, we found that the forelimb and hindlimb acted as a single integrated unit and that neither the forelimb nor hindlimb was significantly more integrated than the other. Therefore, the ontogenetic niche shift itself did not impact limb morphology in A. mississippiensis.

Early amphibians evolved distinct vertebrae for habitat invasions.

Living tetrapods owe their existence to a critical moment 360-340 million years ago when their ancestors walked on land. Vertebrae are central to locomotion, yet systematic testing of correlations between vertebral form and terrestriality and subsequent reinvasions of aquatic habitats is lacking, obscuring our understanding of movement capabilities in early tetrapods. Here, we quantified vertebral shape across a diverse group of Paleozoic amphibians (Temnospondyli) encompassing different habitats and nearly the full range of early tetrapod vertebral shapes. We demonstrate that temnospondyls were likely ancestrally terrestrial and had several early reinvasions of aquatic habitats. We find a greater diversity in temnospondyl vertebrae than previously known. We also overturn long-held hypotheses centered on weight-bearing, showing that neural arch features, including muscle attachment, were plastic across the water-land divide and do not provide a clear signal of habitat preferences. In contrast, intercentra traits were critical, with temnospondyls repeatedly converging on distinct forms in terrestrial and aquatic taxa, with little overlap between. Through our geometric morphometric study, we have been able to document associations between vertebral shape and environmental preferences in Paleozoic tetrapods and to reveal morphological constraints imposed by vertebrae to locomotion, independent of ancestry.

Paleobiology in the 21st Century.

The diversity of research performed by vertebrate paleobiologists is immense, including primary descriptions of newly discovered taxa, biomechanical analyses of extant animals, large-scale macroevolutionary analyses with hundreds of taxa, and many others. This special issue of The Anatomical Record highlights the scope of the research done by paleobiologists, with an emphasis on dinosaurs and their relatives, and is structured into three broad categories of research: (1) anatomical descriptions, (2) the biology of the dinosaurs and their relatives, and (3) macroevolutionary trends. Although these topics all fall within the purview of contemporary paleobiology, many of these topics have been studied in one form or another since the beginning of dinosaur paleobiology as a field. Rather than the topics themselves, what characterizes the modern renaissance of paleobiology is the employment of newly developed quantitative techniques to analyze the relationships between taxa and their evolutionary history. While primary anatomical descriptions remain the central pillar of dinosaur paleobiology, researchers now have a suitable baseline understanding of dinosaurian anatomy and their major evolutionary relationships. Using this baseline, they are able to ask more complex questions and refine their understanding of dinosaurian evolution. The aim of this issue is to exemplify the range of topics examined within dinosaur paleobiology and to look forward to the future of paleobiology. Anat Rec, 303:645-648, 2020. © 2020 American Association for Anatomy.

New Dromaeosaurid Dinosaur (Theropoda, Dromaeosauridae) from New Mexico and Biodiversity of Dromaeosaurids at the end of the Cretaceous.

Dromaeosaurids (Theropoda: Dromaeosauridae), a group of dynamic, swift predators, have a sparse fossil record, particularly at the time of their extinction near the Cretaceous-Paleogene boundary. Here we report on a new dromaeosaurid, Dineobellator notohesperus, gen. and sp. nov., consisting of a partial skeleton from the Upper Cretaceous (Maastrichtian) of New Mexico, the first diagnostic dromaeosaurid to be recovered from the latest Cretaceous of the southern United States (southern Laramidia). The holotype includes elements of the skull, axial, and appendicular skeleton. The specimen reveals a host of morphologies that shed light on new behavioral attributes for these feathered dinosaurs. Unique features on its forelimbs suggest greater strength capabilities in flexion than the normal dromaeosaurid condition, in conjunction with a relatively tighter grip strength in the manual claws. Aspects of the caudal vertebrae suggest greater movement near the tail base, aiding in agility and predation. Phylogenetic analysis places Dineobellator within Velociraptorinae. Its phylogenetic position, along with that of other Maastrichtian taxa (Acheroraptor and Dakotaraptor), suggests dromaeosaurids were still diversifying at the end of the Cretaceous. Furthermore, its recovery as a second North American Maastrichtian velociraptorine suggests vicariance of North American velociraptorines after a dispersal event during the Campanian-Maastrichtian from Asia. Features of Dineobellator also imply that dromaeosaurids were active predators that occupied discrete ecological niches while living in the shadow of Tyrannosaurus rex, until the end of the dinosaurs' reign.

Filling in Gaps in the Ceratopsid Histologic Database: Histology of Two Basal Centrosaurines and an Assessment of the Utility of Rib Histology in the Ceratopsidae.

Bone histology grants substantial insight into the growth and biology of fossil vertebrates. Many of the major non-avian dinosaurian clades have been extensively sampled for bone histologic data allowing reconstruction of their growth as well as the assessment of the evolution of growth changes along phylogenies. However, horned ceratopsians are poorly represented in paleohistologic studies. Further, the ceratopsian taxa that have been examined are unevenly sampled phylogenetically with very basal forms and highly derived forms making up the majority of studied taxa. In order to rectify this, we have histologically sampled Avaceratops from Montana and Yehuecauhceratops from northern Mexico to assess how mid-sized basal centrosaurines grew relative to more basal and derived forms. Based on results from these taxa, basal centrosaurines present a mosaic of growth characters intermediate between those seen in basal ceratopsians and more derived centrosaurines. Further, Yehuecauhceratops has many lines of arrested growth preserved, suggesting that the large number of lines of arrested growth found in a high-latitude Pachyrhinosaurus specimen may be a result of phylogeny rather than geography. Since lines of arrested growth are not preserved in long bones of many ceratopsians, especially chasmosaurines, we also histologically sampled ribs of Avaceratops and Pachyrhinosaurus. However, the largest ribs were highly remodeled obscuring lines of arrested growth, making it unlikely that rib histology will clarify growth trends in ceratopsians. These centrosaurines add to the growing ceratopsian histological database and demonstrate that basal centrosaurines grew in a manner intermediate between non-ceratopsid taxa and derived centrosaurines. Anat Rec, 303:935-948, 2020. © 2019 Wiley Periodicals, Inc.

Quantifying shape and ecology in avian pedal claws: The relationship between the bony core and keratinous sheath.

Terrestrial tetrapods use their claws to interact with their environments in a plethora of ways. Birds in particular have developed a diversity of claw shapes since they are often not bound to terrestrial locomotion and have heterogeneous body masses ranging several orders of magnitude. Numerous previous studies have hypothesized a connection between pedal claw shape and ecological mode in birds, yet have generated conflicting results, spanning from clear ecological groupings based on claw shape to a complete overlap of ecological modes. The majority of these studies have relied on traditional morphometric arc measurements of keratinous sheaths and have variably accounted for likely confounding factors such as body mass and phylogenetic relatedness. To better address the hypothesized relationship between ecology and claw shape in birds, we collected 580 radiographs allowing visualization of the bony core and keratinous sheath shape in 21 avian orders. Geometric morphometrics was used to quantify bony core and keratinous sheath shape and was compared to results using traditional arc measurements. Neither approach significantly separates bird claws into coarse ecological categories after integrating body size and phylogenetic relatedness; however, some separation between ecological groups is evident and we find a gradual shift from the claw shape of ground-dwelling birds to those of predatory birds. Further, the bony claw core and keratinous sheath are significantly correlated, and the degree of functional integration does not differ across ecological groups. Therefore, it is likely possible to compare fossil bony cores with extant keratinous sheaths after applying corrections. Finally, traditional metrics and geometric morphometric shape are significantly, yet loosely correlated. Based on these results, future workers are encouraged to use geometric morphometric approaches to study claw geometry and account for confounding factors such as body size, phylogeny, and individual variation prior to predicting ecology in fossil taxa.

A Centrosaurine (Dinosauria: Ceratopsia) from the Aguja Formation (Late Campanian) of Northern Coahuila, Mexico.

While centrosaurines and ceratopsids in general are abundant in the Late Campanian of northern Laramidia, they are much less commonly found in southern Laramidia. This has supported hypotheses of dinosaur provinciality and endemism in the Late Cretaceous with the delineation of at least two separate faunal zones, north and south Laramidia. There have been 12 genera of centrosaurines recognized from northern Laramidia while two genera, Diabloceratops and Nasutoceratops, have been named from southern Laramidia. We present an osteological description and taphonomic outline for a new centrosaurine ceratopsid from the Aguja Formation of northern Coahuila, Mexico that is not currently diagnosable to the generic level, but likely represents a new taxon. Further, we have included three-dimensional surface scans of all material attributed to this animal. Considering the large number of centrosaurines from northern Laramidia, it is likely that cladistic analyses are biased towards this faunal zone. New findings of southern centrosaurines are needed to correct this bias. This discovery expands the range of centrosaurines south to Coahuila, Mexico and adds new information to better characterize the morphology and taxonomy of centrosaurines from southern Laramidia and their evolution in comparison to their northern counterparts.