Validating osteological correlates for the hepatic piston in the American alligator (Alligator mississippiensis).

Unlike the majority of sauropsids, which breathe primarily through costal and abdominal muscle contractions, extant crocodilians have evolved the hepatic piston pump, a unique additional ventilatory mechanism powered by the diaphragmaticus muscle. This muscle originates from the bony pelvis, wrapping around the abdominal viscera, extending cranially to the liver. The liver then attaches to the caudal margin of the lungs, resulting in a sub-fusiform morphology for the entire "pulmo-hepatic-diaphragmatic" structure. When the diaphragmaticus muscle contracts during inspiration, the liver is pulled caudally, lowering pressure in the thoracolumbar cavity, and inflating the lungs. It has been established that the hepatic piston pump requires the liver to be displaced to ventilate the lungs, but it has not been determined if the lungs are freely mobile or if the pleural tissues stretch ventrally. It has been hypothesized that the lungs are able to slide craniocaudally with the liver due to the smooth internal ceiling of the thoracolumbar cavity. We assess this through ultrasound video and demonstrate quantitatively and qualitatively that the pulmonary tissues are sliding craniocaudally across the interior thoracolumbar ceiling in actively ventilating live juvenile, sub-adult, and adult individuals (n = 7) of the American alligator (Alligator mississippiensis) during both natural and induced ventilation. The hepatic piston is a novel ventilatory mechanism with a relatively unknown evolutionary history. Questions related to when and under what conditions the hepatic piston first evolved have previously been left unanswered due to a lack fossilized evidence for its presence or absence. By functionally correlating specific characters in the axial skeleton to the hepatic piston, these osteological correlates can be applied to fossil taxa to reconstruct the evolution of the hepatic piston in extinct crocodylomorph archosaurs.

Cochlea development shapes bat sensory system evolution.

Sensory organs must develop alongside the skull within which they are largely encased, and this relationship can manifest as the skull constraining the organs, organs constraining the skull, or organs constraining one another in relative size. How this interplay between sensory organs and the developing skull plays out during the evolution of sensory diversity; however, remains unknown. Here, we examine the developmental sequence of the cochlea, the organ responsible for hearing and echolocation, in species with distinct diet and echolocation types within the ecologically diverse bat super-family Noctilionoidea. We found the size and shape of the cochlea largely correlates with skull size, with exceptions of Pteronotus parnellii, whose high duty cycle echolocation (nearly constant emission of sound pulses during their echolocation process allowing for detailed information gathering, also called constant frequency echolocation) corresponds to a larger cochlear and basal turn, and Monophyllus redmani, a small-bodied nectarivorous bat, for which interactions with other sensory organs restrict cochlea size. Our findings support the existence of developmental constraints, suggesting that both developmental and anatomical factors may act synergistically during the development of sensory systems in noctilionoid bats.

Perspectives on lung visualization: Three-dimensional anatomical modeling of computed and micro-computed tomographic data in comparative evolutionary morphology and medicine with applications for COVID-19.

The vertebrate respiratory system is challenging to study. The complex relationship between the lungs and adjacent tissues, the vast structural diversity of the respiratory system both within individuals and between taxa, its mobility (or immobility) and distensibility, and the difficulty of quantifying and visualizing functionally important internal negative spaces have all impeded descriptive, functional, and comparative research. As a result, there is a relative paucity of three-dimensional anatomical information on this organ system in all vertebrate groups (including humans) relative to other regions of the body. We present some of the challenges associated with evaluating and visualizing the vertebrate respiratory system using computed and micro-computed tomography and its subsequent digital segmentation. We discuss common mistakes to avoid when imaging deceased and live specimens and various methods for merging manual and threshold-based segmentation approaches to visualize pulmonary tissues across a broad range of vertebrate taxa, with a particular focus on sauropsids (reptiles and birds). We also address some of the recent work in comparative evolutionary morphology and medicine that have used these techniques to visualize respiratory tissues. Finally, we provide a clinical study on COVID-19 in humans in which we apply modeling methods to visualize and quantify pulmonary infection in the lungs of human patients.

Disentangling Mechanical and Sensory Modules in the Radiation of Noctilionoid Bats.

AbstractWith diverse mechanical and sensory functions, the vertebrate cranium is a complex anatomical structure whose shifts between modularity and integration, especially in mechanical function, have been implicated in adaptive diversification. Yet how mechanical and sensory systems and their functions coevolve, as well as how their interrelationship contributes to phenotypic disparity, remain largely unexplored. To examine the modularity, integration, and evolutionary rates of sensory and mechanical structures within the head, we analyzed hard and soft tissue scans from ecologically diverse bats in the superfamily Noctilionoidea, a clade that ranges from insectivores and carnivores to frugivores and nectarivores. We identified eight regions that evolved in a coordinated fashion, thus recognizable as evolutionary modules: five associated with bite force and three linked to olfactory, visual, and auditory systems. Interrelationships among these modules differ between Neotropical leaf-nosed bats (family Phyllostomidae) and other noctilionoids. Consistent with the hypothesis that dietary transitions begin with changes in the capacity to detect novel food items followed by adaptations to process them, peak rates of sensory module evolution predate those of some mechanical modules. We propose that the coevolution of structures influencing bite force, olfaction, vision, and hearing constituted a structural opportunity that allowed the phyllostomid ancestor to take advantage of existing ecological opportunities and contributed to the clade's remarkable radiation.

Projected northward shifts in eastern red-backed salamanders due to changing climate.

Many species' distributions are being impacted by the acceleration of climate change. Amphibians in particular serve numerous ecosystem functions and are useful indicators of environmental change. Understanding how their distributions have been impacted by climate change and will continue to be impacted is thus important to overall ecosystem health. Plethodon cinereus (Eastern Red-Backed Salamander) is a widespread species of lungless salamander (Plethodontidae) that ranges across northeastern North America. To better understand future potential lungless salamander range shifts, we quantify environmental favorability, the likelihood of membership in a set of sites where environmental conditions are favorable for a species, for P. cinereus in multiple time periods, and examine shifts in the species' distribution. First, utilizing a large data set of georeferenced records, we assessed which bioclimatic variables were associated with environmental favorability in P. cinereus. We then used species distribution modeling for two time periods (1961-1980 and 2001-2020) to determine whether there was a regional shift in environmental favorability in the past 60 years. Models were then used to project future distributions under eight climate change scenarios to quantify potential range shifts. Shifts were assessed using fuzzy logic, avoiding thresholds that oversimplify model predictions into artificial binary outputs. We found that P. cinereus presence is strongly associated with environmental stability. There has been a substantial northward shift in environmental favorability for P. cinereus between 1961-1980 and 2001-2020. This shift is predicted to continue by 2070, with larger shifts under higher greenhouse gas emission scenarios. As climate change accelerates, it is differentially impacting species but has especially strong impacts on dispersal-limited species. Our results show substantial northward shifts in climatic favorability in the last 60 years for P. cinereus, which are likely to be exacerbated by ongoing climate change. Since P. cinereus is dispersal-limited, these models may imply local extirpations along the southern modern range with limited northward dispersal. Continued monitoring of amphibians in the field will reveal microclimatic effects associated with climate change and the accuracy of the model predictions presented here.

Dots on a screen: The past, present, and future of morphometrics in the study of nonavian dinosaurs.

Using morphometrics to study nonavian dinosaur fossils is a practice that predates the origin of the word "dinosaur." By the 1970s, linear morphometrics had become established as a valuable tool for analyzing intra- and interspecific variation in nonavian dinosaurs. With the advent of more recent techniques such as geometric morphometrics and more advanced statistical approaches, morphometric analyses of nonavian dinosaurs have proliferated, granting unprecedented insight into many aspects of their biology and evolution. I outline the past, present, and future of morphometrics as applied to the study of nonavian dinosaurs zeroing in on five aspects of nonavian dinosaur paleobiology where morphometrics has been widely utilized to advance our knowledge: systematics, sexual dimorphism, locomotion, macroevolution, and trackways. Morphometric methods are especially susceptible to taphonomic distortion. As such, the impact of taphonomic distortion on original fossil shape is discussed as are current and future methods for quantifying and accounting for distortion with the goal of reducing the taphonomic noise to biological signal ratio. Finally, the future of morphometrics in nonavian dinosaur paleobiology is discussed as paleobiologists move into a "virtual paleobiology" framework, whereby digital renditions of fossils are captured via methods such as photogrammetry and computed tomography. These primary data form the basis for three-dimensional (3D) geometric morphometric analyses along with a slew of other forms of analyses. These 3D specimen data form part of the extended specimen and help to democratize paleobiology, unlocking the specimen from the physical museum and making the specimen available to researchers across the world.

Convergent evolution of quadrupedality in ornithischian dinosaurs was achieved through disparate forelimb muscle mechanics.

The secondary evolution of quadrupedality from bipedal ancestry is a rare evolutionary transition in tetrapods yet occurred convergently at least three times within ornithischian dinosaurs. Despite convergently evolving quadrupedal gait, ornithischians exhibited variable anatomy, particularly in the forelimbs, which underwent a major functional change from assisting in foraging and feeding in bipeds to becoming principal weight-bearing components of the locomotor system in quadrupeds. Here, we use three-dimensional multi-body dynamics models to demonstrate quantitatively that different quadrupedal ornithischian clades evolved distinct forelimb musculature, particularly around the shoulder. We find that major differences in glenohumeral abduction-adduction and long axis rotation muscle leverages were key drivers of mechanical disparity, thereby refuting previous hypotheses about functional convergence in major clades. Elbow muscle leverages were also disparate across the major ornithischian lineages, although high elbow extension muscle leverages were convergent between most quadrupeds. Unlike in ornithischian hind limbs, where differences are more closely tied to functional similarity than phylogenetic relatedness, mechanical disparity in ornithischian forelimbs appears to have been shaped primarily by phylogenetic constraints. Differences in ancestral bipedal taxa within each clade may have resulted in disparate ecomorphological constraints on the evolutionary pathways driving divergence in their quadrupedal descendants.

Ecological constraints on highly evolvable olfactory receptor genes and morphology in neotropical bats.

Although evolvability of genes and traits may promote specialization during species diversification, how ecology subsequently restricts such variation remains unclear. Chemosensation requires animals to decipher a complex chemical background to locate fitness-related resources, and thus the underlying genomic architecture and morphology must cope with constant exposure to a changing odorant landscape; detecting adaptation amidst extensive chemosensory diversity is an open challenge. In phyllostomid bats, an ecologically diverse clade that evolved plant visiting from a presumed insectivorous ancestor, the evolution of novel food detection mechanisms is suggested to be a key innovation, as plant-visiting species rely strongly on olfaction, supplementarily using echolocation. If this is true, exceptional variation in underlying olfactory genes and phenotypes may have preceded dietary diversification. We compared olfactory receptor (OR) genes sequenced from olfactory epithelium transcriptomes and olfactory epithelium surface area of bats with differing diets. Surprisingly, although OR evolution rates were quite variable and generally high, they are largely independent of diet. Olfactory epithelial surface area, however, is relatively larger in plant-visiting bats and there is an inverse relationship between OR evolution rates and surface area. Relatively larger surface areas suggest greater reliance on olfactory detection and stronger constraint on maintaining an already diverse OR repertoire. Instead of the typical case in which specialization and elaboration are coupled with rapid diversification of associated genes, here the relevant genes are already evolving so quickly that increased reliance on smell has led to stabilizing selection, presumably to maintain the ability to consistently discriminate among specific odorants-a potential ecological constraint on sensory evolution.

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.

Architecture of the bronchial tree in Cuvier's dwarf caiman (Paleosuchus palpebrosus).

We imaged the lungs of five Cuvier's dwarf caiman (Paleosuchus palpebrosus) via computed tomography (CT) and micro-computed tomography (µCT) and compared these data to the lungs of the American alligator (Alligator mississippiensis). These data demonstrate anatomical commonalities between the lungs of P. palpebrosus and A. mississippiensis, and a few notable differences. The structural similarities are (a) a proximally narrow, distally widened, hook-shaped primary bronchus; (b) a cervical ventral bronchus that branches of the primary bronchus and immediately makes a hairpin turn toward the apex of the lung; (c) a sequential series of dorsobronchi arising from the primary bronchus caudal to the cervical ventral bronchus; (d) intraspecifically highly variable medial sequence of secondary airways; (e) sac-like laterobronchi; and (f) grossly dead-ended caudal group bronchi in the caudal and ventral aspects of the lung. The primary differences between the two taxa are in the overall number of large bronchi (fewer in P. palpebrosus), and the number of branches that contribute to the cardiac regions. Imaging data of both a live and deceased specimen under varying states (postprandial, fasting, total lung capacity, open to atmosphere) indicate that the caudal margin and position of the lungs shift craniocaudally relative to the vertebral column. These imaging data suggest that the smooth thoracic ceiling may be correlated to visceral movement during ventilation, but this hypothesis warrants validation. These results provide the scaffolding for future comparisons between crocodilians, for generating preliminary reconstructions of the ancestral crocodilian bronchial tree, and establishing new hypotheses of bronchial homology across Archosauria.

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.

Inter- and intraspecific variation in the Artibeus species complex demonstrates size and shape partitioning among species.

Neotropical leaf-nosed bats (family Phyllostomidae) are one of the most diverse mammalian families and Artibeus spp. is one of the most speciose phyllostomid genera. In spite of their species diversity, previous work on Artibeus crania using linear morphometrics has uncovered limited interspecific variation. This dearth of shape variation suggests that differences in cranial morphology are not contributing to niche partitioning across species, many of which are often found in sympatry. Using two-dimensional geometric morphometric methods on crania from eleven species from the Artibeus species complex, the current study demonstrates substantial cranial interspecific variation, sexual size and shape dimorphism, and intraspecific geographic variation. The majority of species were shown to have a unique size and shape, which suggests that each species may be taking advantage of slightly different ecological resources. Further, both sexual size and shape dimorphism were significant in the Artibeus species complex. Male and female Artibeus are known to have sex specific foraging strategies, with males eating near their roosts and females feeding further from their roosts. The presence of cranial sexual dimorphism in the Artibeus species complex, combined with previous work showing that different fruit size and hardness is correlated with different cranial shapes in phyllostomids, indicates that the males and females may be utilizing different food resources, leading to divergent cranial morphotypes. Additional field studies will be required to confirm this emergent hypothesis. Finally, significant geographical shape variation was found in a large intraspecific sample of Artibeus lituratus crania. However, this variation was not correlated with latitude and instead may be linked to local environmental factors. Additional work on ecology and behavior in the Artibeus species complex underlying the morphological variation uncovered in this study will allow for a better understanding of how the group has reached its present diversity.

Find the food first: An omnivorous sensory morphotype predates biomechanical specialization for plant based diets in phyllostomid bats.

The role of mechanical morphologies in the exploitation of novel niche space is well characterized; however, the role of sensory structures in unlocking new niches is less clear. Here, we investigate the relationship between the evolution of sensory structures and diet during the radiation of noctilionoid bats. With a broad range of foraging ecologies and a well-supported phylogeny, noctilionoids constitute an ideal group for studying this relationship. We used diffusible iodine-based contrast enhanced computed tomography scans of 44 noctilionoid species to analyze relationships between the relative volumes of three sensory structures (olfactory bulbs, orbits, and cochleae) and diet. We found a positive relationship between frugivory and both olfactory and orbit size. However, we also found a negative relationship between nectarivory and cochlea size. Ancestral state estimates suggest that larger orbits and olfactory bulbs were present in the common ancestor of family Phyllostomidae, but not in other noctilionoid. This constellation of traits indicates a shift toward omnivory at the base of Phyllostomidae, predating their radiation into an exceptionally broad range of dietary niches. This is consistent with a scenario in which changes in sensory systems associated with foraging and feeding set the stage for subsequent morphological modification and diversification.

Anatomy, variation, and asymmetry of the bronchial tree in the African grey parrot (Psittacus erithacus).

The avian bronchial tree has a unique and elaborate architecture for the maintenance of unidirectional airflow. Gross descriptions of this bronchial arrangement have traditionally relied upon dissection and casts of the negative (air-filled) spaces. In this study, the bronchial trees of five deceased African grey parrots (Psittacus erithacus) were segmented from micro-computed tomography (µCT) scans into three-dimensional (3D) surface models, and then compared. Select metrics of the primary bronchi and major secondary branches in the µCT scans of 11 specimens were taken to assess left-right asymmetry and quantify gross lung structure. Analysis of the 3D surface models demonstrates variation in the number and distribution of secondary bronchi with consistent direct connections to specific respiratory air sacs. A single model of the parabronchi further reveals indirect connections to all but two of the nine total air sacs. Statistical analysis of the metrics show significant left-right asymmetry between the primary bronchi and the origins of the first four secondary bronchi (the ventrobronchi), consistently greater mean values for all right primary bronchus length metrics, and relatively high coefficients of variation for cross-sectional area metrics of the primary bronchi and secondary bronchi ostia. These findings suggest that the lengths of the primary bronchi distal to the ventrobronchi do not preserve lung symmetry, and that aerodynamic valving can functionally accommodate a wide range of bronchial proportions.

Anatomy, ontogeny, and evolution of the archosaurian respiratory system: A case study on Alligator mississippiensis and Struthio camelus.

The avian lung is highly specialized and is both functionally and morphologically distinct from that of their closest extant relatives, the crocodilians. It is highly partitioned, with a unidirectionally ventilated and immobilized gas-exchanging lung, and functionally decoupled, compliant, poorly vascularized ventilatory air-sacs. To understand the evolutionary history of the archosaurian respiratory system, it is essential to determine which anatomical characteristics are shared between birds and crocodilians and the role these shared traits play in their respective respiratory biology. To begin to address this larger question, we examined the anatomy of the lung and bronchial tree of 10 American alligators (Alligator mississippiensis) and 11 ostriches (Struthio camelus) across an ontogenetic series using traditional and micro-computed tomography (µCT), three-dimensional (3D) digital models, and morphometry. Intraspecific variation and left to right asymmetry were present in certain aspects of the bronchial tree of both taxa but was particularly evident in the cardiac (medial) region of the lungs of alligators and the caudal aspect of the bronchial tree in both species. The cross-sectional area of the primary bronchus at the level of the major secondary airways and cross-sectional area of ostia scaled either isometrically or negatively allometrically in alligators and isometrically or positively allometrically in ostriches with respect to body mass. Of 15 lung metrics, five were significantly different between the alligator and ostrich, suggesting that these aspects of the lung are more interspecifically plastic in archosaurs. One metric, the distances between the carina and each of the major secondary airways, had minimal intraspecific or ontogenetic variation in both alligators and ostriches, and thus may be a conserved trait in both taxa. In contrast to previous descriptions, the 3D digital models and CT scan data demonstrate that the pulmonary diverticula pneumatize the axial skeleton of the ostrich directly from the gas-exchanging pulmonary tissues instead of the air sacs. Global and specific comparisons between the bronchial topography of the alligator and ostrich reveal multiple possible homologies, suggesting that certain structural aspects of the bronchial tree are likely conserved across Archosauria, and may have been present in the ancestral archosaurian lung.

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.

Asymmetric and Spiraled Genitalia Coevolve with Unique Lateralized Mating Behavior.

Asymmetric genitalia and lateralized mating behaviors occur in several taxa, yet whether asymmetric morphology in one sex correlates or coevolves with lateralized mating behavior in the other sex remains largely unexplored. While lateralized mating behaviors are taxonomically widespread, among mammals they are only known in the harbor porpoise (Phocoena phocoena). Males attempt copulation by approaching a female exclusively on her left side. To understand if this unusual lateralized behavior may have coevolved with genital morphology, we quantified the shape of female and male harbor porpoise reproductive tracts using 2D geometric morphometrics and 3D models of the vaginal lumen and inflated distal penis. We found that the vaginas varied individually in shape and that the vaginas demonstrated both significant directional and fluctuating asymmetry. This asymmetry resulted from complex 3D spirals and vaginal folds with deep recesses, which may curtail the depth or direction of penile penetration and/or semen movement. The asymmetric shapes of the vaginal lumen and penis tip were both left-canted with similar angular bends that mirrored one another and correspond with the left lateral mating approach. We suggest that the reproductive anatomy of both sexes and their lateral mating behavior coevolved.

The evolutionary diversity of locomotor innovation in rodents is not linked to proximal limb morphology.

Rodents are the most species-rich order within Mammalia and have evolved disparate morphologies to accommodate numerous locomotor niches, providing an excellent opportunity to understand how locomotor innovation can drive speciation. To evaluate the connection between the evolutionary success of rodents and the diversity of rodent locomotor ecologies, we used a large dataset of proximal limb CT scans from across Myomorpha and Geomyoidea to examine internal and external limb shape. Only fossorial rodents displayed a major reworking of their proximal limbs in either internal or external morphology, with other locomotor modes plotting within a generalist morphospace. Fossorial rodents were also the only locomotor mode to consistently show increased rates of humerus/femur morphological evolution. We propose that these rodent clades were successful at spreading into ecological niches due to high behavioral plasticity and small body sizes, allowing them to modify their locomotor mode without requiring major changes to their proximal limb morphology.

The postcranial skeleton of the erythrosuchid archosauriform Garjainia prima from the Early Triassic of European Russia.

Erythrosuchidae were large-bodied, quadrupedal, predatory archosauriforms that dominated the hypercarnivorous niche in the aftermath of the Permo-Triassic mass extinction. Garjainia, one of the oldest members of the clade, is known from the late Olenekian of European Russia. The holotype of Garjainia prima comprises a well-preserved skull, but highly incomplete postcranium. Recent taxonomic reappraisal demonstrates that material from a bone bed found close to the type locality, previously referred to as 'Vjushkovia triplicostata', is referable to G. prima. At least, seven individuals comprising cranial remains and virtually the entire postcranium are represented, and we describe this material in detail for the first time. An updated phylogenetic analysis confirms previous results that a monophyletic Garjainia is the sister taxon to a clade containing Erythrosuchus, Shansisuchus and Chalishevia. Muscle scars on many limb elements are clear, allowing reconstruction of the proximal locomotor musculature. We calculate the body mass of G. prima to have been 147-248 kg, similar to that of an adult male lion. Large body size in erythrosuchids may have been attained as part of a trend of increasing body size after the Permo-Triassic mass extinction and allowed erythrosuchids to become the dominant carnivores of the Early and Middle Triassic.

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.