Ecomorphological patterns in trigeminal canal branching among sauropsids reveal sensory shift in suchians.

The vertebrate trigeminal nerve is the primary mediator of somatosensory information from nerve endings across the face, extending nerve branches through bony canals in the face and mandibles, terminating in sensory receptors. Reptiles evolved several extreme forms of cranial somatosensation in which enhanced trigeminal tissues are present in species engaging in unique mechanosensory behaviors. However, morphology varies by clade and ecology among reptiles. Few lineages approach the extreme degree of tactile somatosensation possessed by crocodylians, the only remaining members of a clade that underwent an ecological transition from the terrestrial to semiaquatic habitat, also evolving a specialized trigeminal system. It remains to be understood how trigeminal osteological correlates inform how adaptations for enhanced cranial sensation evolved in crocodylians. Here we identify an increase in sensory abilities in Early Jurassic crocodylomorphs, preceding the transitions to a semiaquatic habitat. Through quantification of trigeminal neurovascular canal branching patterns in an extant phylogenetic bracket we quantify and identify morphologies associated with sensory behaviors in representative fossil taxa, we find stepwise progression of increasing neurovascular canal density, complexity, and distribution from the primitive archosaurian to the derived crocodilian condition. Model-based inferences of sensory ecologies tested on quantified morphologies of extant taxa with known sensory behaviors indicate a parallel increase in sensory abilities among pseudosuchians. These findings establish patterns of reptile trigeminal ecomorphology, revealing evolutionary patterns of somatosensory ecology.

2D and 3D visualizations of archosaur jaw muscle mechanics, ontogeny and phylogeny using ternary diagrams and 3D modeling.

Comparing patterns of performance and kinematics across behavior, development and phylogeny is crucial to understand the evolution of complex musculoskeletal systems such as the feeding apparatus. However, conveying 3D spatial data of muscle orientation throughout a feeding cycle, ontogenetic pathway or phylogenetic lineage is essential to understanding the function and evolution of the skull in vertebrates. Here, we detail the use of ternary plots for displaying and comparing the 3D orientation of muscle data. First, we illustrate changes in 3D jaw muscle resultants during jaw closing taxa the American alligator (Alligator mississippiensis). Second, we show changes in 3D muscle resultants of jaw muscles across an ontogenetic series of alligators. Third, we compare 3D resultants of jaw muscles of avian-line dinosaurs, including extant (Struthio camelus, Gallus gallus, Psittacus erithacus) and extinct (Tyrannosaurus rex) species to outline the reorganization of jaw muscles that occurred along the line to modern birds. Finally, we compare 3D resultants of jaw muscles of the hard-biting species in our sample (A. mississippiensis, T. rex, P. erithacus) to illustrate how disparate jaw muscle resultants are employed in convergent behaviors in archosaurs. Our findings show that these visualizations of 3D components of jaw muscles are immensely helpful towards identifying patterns of cranial performance, growth and diversity. These tools will prove useful for testing other hypotheses in functional morphology, comparative biomechanics, ecomorphology and organismal evolution.

The roles of joint tissues and jaw muscles in palatal biomechanics of the savannah monitor (Varanus exanthematicus) and their significance for cranial kinesis.

Numerous vertebrates exhibit cranial kinesis, or movement between bones of the skull and mandible other than at the jaw joint. Many kinetic species possess a particular suite of features to accomplish this movement, including flexible cranial joints and protractor musculature. Whereas the musculoskeletal anatomy of these kinetic systems is well understood, how these joints are biomechanically loaded, how different soft tissues affect joint loading and kinetic capacity, and how the protractor musculature loads the skull remain poorly understood. Here, we present a finite element model of the savannah monitor, Varanus exanthematicus, a modestly kinetic lizard, to better elucidate the roles of soft tissue in mobile joints and protractor musculature in cranial loading. We describe the 3D resultants of jaw muscles and the histology of palatobasal, otic and jaw joints. We tested the effects of joint tissue type, bite point and muscle load to evaluate the biomechanical role of muscles on the palate and braincase. We found that the jaw muscles have significant mediolateral components that can impart stability across palatocranial joints. Articular tissues affect the magnitude of strains experienced around the palatobasal and otic joints. Without protractor muscle loading, the palate, quadrate and braincase experience higher strains, suggesting this muscle helps insulate the braincase and palatoquadrate from high loads. We found that the cross-sectional properties of the bones of V. exanthematicus are well suited for performing under torsional loads. These findings suggest that torsional loading regimes may have played a more important role in the evolution of cranial kinesis in lepidosaurs than previously appreciated.

Joint histology in Alligator mississippiensis challenges the identification of synovial joints in fossil archosaurs and inferences of cranial kinesis.

Archosaurs, like all vertebrates, have different types of joints that allow or restrict cranial kinesis, such as synovial joints and fibrous joints. In general, synovial joints are more kinetic than fibrous joints, because the former possess a fluid-filled cavity and articular cartilage that facilitate movement. Even though there is a considerable lack of data on the microstructure and the structure-function relationships in the joints of extant archosaurs, many functional inferences of cranial kinesis in fossil archosaurs have hinged on the assumption that elongated condylar joints are (i) synovial and/or (ii) kinetic. Cranial joint microstructure was investigated in an ontogenetic series of American alligators, Alligator mississippiensis All the presumably synovial, condylar joints found within the head of the American alligator (the jaw joint, otic joint and laterosphenoid-postorbital (LS-PO) joint) were studied by means of paraffin histology and undecalcified histology paired with micro-computed tomography data to better visualize three-dimensional morphology. Results show that among the three condylar joints of A. mississippiensis, the jaw joint was synovial as expected, but the otherwise immobile otic and LS-PO joints lacked a synovial cavity. Therefore, condylar morphology does not always imply the presence of a synovial articulation nor mobility. These findings reveal an undocumented diversity in the joint structure of alligators and show that crocodylians and birds build novel, kinetic cranial joints differently. This complicates accurate identification of synovial joints and functional inferences of cranial kinesis in fossil archosaurs and tetrapods in general.

Cranial joint histology in the mallard duck (Anas platyrhynchos): new insights on avian cranial kinesis.

The evolution of avian cranial kinesis is a phenomenon in part responsible for the remarkable diversity of avian feeding adaptations observable today. Although osteological, developmental and behavioral features of the feeding system are frequently studied, comparatively little is known about cranial joint skeletal tissue composition and morphology from a microscopic perspective. These data are key to understanding the developmental, biomechanical and evolutionary underpinnings of kinesis. Therefore, here we investigated joint microstructure in juvenile and adult mallard ducks (Anas platyrhynchos; Anseriformes). Ducks belong to a diverse clade of galloanseriform birds, have derived adaptations for herbivory and kinesis, and are model organisms in developmental biology. Thus, new insights into their cranial functional morphology will refine our understanding of avian cranial evolution. A total of five specimens (two ducklings and three adults) were histologically sampled, and two additional specimens (a duckling and an adult) were subjected to micro-computed tomographic scanning. Five intracranial joints were sampled: the jaw joint (quadrate-articular); otic joint (quadrate-squamosal); palatobasal joint (parasphenoid-pterygoid); the mandibular symphysis (dentary-dentary); and the craniofacial hinge (a complex flexion zone involving four different pairs of skeletal elements). In both the ducklings and adults, the jaw, otic and palatobasal joints are all synovial, with a synovial cavity and articular cartilage on each surface (i.e. bichondral joints) ensheathed in a fibrous capsule. The craniofacial hinge begins as an ensemble of patent sutures in the duckling, but in the adult it becomes more complex: laterally it is synovial; whereas medially, it is synostosed by a bridge of chondroid bone. We hypothesize that it is chondroid bone that provides some of the flexible properties of this joint. The heavily innervated mandibular symphysis is already fused in the ducklings and remains as such in the adult. The results of this study will serve as reference for documenting avian cranial kinesis from a microanatomical perspective. The formation of: (i) secondary articular cartilage on the membrane bones of extant birds; and (ii) their unique ability to form movable synovial joints within two or more membrane bones (i.e. within their dermatocranium) might have played a role in the origin and evolution of modern avian cranial kinesis during dinosaur evolution.

Ontogeny of bite force in a validated biomechanical model of the American alligator.

Three-dimensional computational modeling offers tools with which to investigate forces experienced by the skull during feeding and other behaviors. American alligators (Alligator mississippiensis) generate some of the highest measured bite forces among extant tetrapods. A concomitant increase in bite force accompanies ontogenetic increases in body mass, which has been linked with dietary changes as animals increase in size. Because the flattened skull of crocodylians has substantial mediolaterally oriented muscles, crocodylians are an excellent model taxon in which to explore the role of mediolateral force components experienced by the feeding apparatus. Many previous modeling studies of archosaur cranial function focused on planar analysis, ignoring the mediolateral aspects of cranial forces. Here, we used three-dimensionally accurate anatomical data to resolve 3D muscle forces. Using dissection, imaging and computational techniques, we developed lever and finite element models of an ontogenetic series of alligators to test the effects of size and shape on cranial loading and compared estimated bite forces with those previously measured in vivo in A. mississippiensis We found that modeled forces matched in vivo data well for intermediately sized individuals, and somewhat overestimated force in smaller specimens and underestimated force in larger specimens, suggesting that ontogenetically static muscular parameters and bony attachment sites alone cannot account for all the variation in bite force. Adding aponeurotic muscle attachments would likely improve force predictions, but such data are challenging to model and integrate into analyses of extant taxa and are generally unpreserved in fossils. We conclude that anatomically accurate modeling of muscles can be coupled with finite element and lever analyses to produce reliable, reasonably accurate estimate bite forces and thus both skeletal and joint loading, with known sources of error, which can be applied to extinct taxa.

Diffusible iodine-based contrast-enhanced computed tomography (diceCT): an emerging tool for rapid, high-resolution, 3-D imaging of metazoan soft tissues.

Morphologists have historically had to rely on destructive procedures to visualize the three-dimensional (3-D) anatomy of animals. More recently, however, non-destructive techniques have come to the forefront. These include X-ray computed tomography (CT), which has been used most commonly to examine the mineralized, hard-tissue anatomy of living and fossil metazoans. One relatively new and potentially transformative aspect of current CT-based research is the use of chemical agents to render visible, and differentiate between, soft-tissue structures in X-ray images. Specifically, iodine has emerged as one of the most widely used of these contrast agents among animal morphologists due to its ease of handling, cost effectiveness, and differential affinities for major types of soft tissues. The rapid adoption of iodine-based contrast agents has resulted in a proliferation of distinct specimen preparations and scanning parameter choices, as well as an increasing variety of imaging hardware and software preferences. Here we provide a critical review of the recent contributions to iodine-based, contrast-enhanced CT research to enable researchers just beginning to employ contrast enhancement to make sense of this complex new landscape of methodologies. We provide a detailed summary of recent case studies, assess factors that govern success at each step of the specimen storage, preparation, and imaging processes, and make recommendations for standardizing both techniques and reporting practices. Finally, we discuss potential cutting-edge applications of diffusible iodine-based contrast-enhanced computed tomography (diceCT) and the issues that must still be overcome to facilitate the broader adoption of diceCT going forward.

Developmental exposure to bisphenol A (BPA) alters sexual differentiation in painted turtles (Chrysemys picta).

Environmental chemicals can disrupt endocrine signaling and adversely impact sexual differentiation in wildlife. Bisphenol A (BPA) is an estrogenic chemical commonly found in a variety of habitats. In this study, we used painted turtles (Chrysemys picta), which have temperature-dependent sex determination (TSD), as an animal model for ontogenetic endocrine disruption by BPA. We hypothesized that BPA would override TSD and disrupt sexual development. We incubated farm-raised turtle eggs at the male-producing temperature (26°C), randomly assigned individuals to treatment groups: control, vehicle control, 17ß-estradiol (E2, 20ng/g-egg) or 0.01, 1.0, 100µgBPA/g-egg and harvested tissues at hatch. Typical female gonads were present in 89% of the E2-treated "males", but in none of the control males (n=35). Gonads of BPA-exposed turtles had varying amounts of ovarian-like cortical (OLC) tissue and disorganized testicular tubules in the medulla. Although the percentage of males with OLCs increased with BPA dose (BPA-low=30%, BPA-medium=33%, BPA-high=39%), this difference was not significant (p=0.85). In all three BPA treatments, SOX9 patterns revealed disorganized medullary testicular tubules and ß-catenin expression in a thickened cortex. Liver vitellogenin, a female-specific liver protein commonly used as an exposure biomarker, was not induced by any of the treatments. Notably, these results suggest that developmental exposure to BPA disrupts sexual differentiation in painted turtles. Further examination is necessary to determine the underlying mechanisms of sex reversal in reptiles and how these translate to EDC exposure in wild populations.

Grey parrot (Psittacus erithacus) beak papillae and nerves identified using novel 2-D and 3-D imaging modalities.

OBJECTIVE: The avian beak is a complex organ containing bone, neurovascular tissue, and keratinized covering (rhamphotheca). Nerve-rich papillae extend through bone into rhamphotheca providing sensory input from the beak tip. Beak trimming is a common procedure in avian species and is used for corrective, cosmetic, and behavioral modification purposes. Yet, practitioners are not well versed in complete beak anatomy, and therefore, beak trimming often disregards neurovasculature, injuring the patient and hampering recovery. Here, using comprehensive anatomical description, we aim to provide recommendations on how to safely perform beak trimming without damaging underlying sensory papillae. ANIMALS: Here, we evaluated beaks of 2 deceased grey parrots (Psittacus erithacus). PROCEDURES: In one, we used a novel stain and microcomputed tomography to visualize papillae in the upper and lower beaks. In a second, we hand isolated the upper and lower beak dermal papillae and used high-resolution photography plus traditional paraffin histology. RESULTS: Papillae and their nerves were easily identified in these 2- and 3-dimensional approaches. This allowed us to determine the approximate lengths of papillae within the upper and lower beak. CLINICAL RELEVANCE: Based on these findings, the authors recommend lateral radiographs of the bird's head and beak to identify the location of the underlying bone relative to the overlying rhamphotheca before performing beak trims. Specifically in grey parrots, the authors recommend the upper and lower beak should not be trimmed closer than 8 to 10 mm from the underlying bone. Further work is needed to support these recommendations and provide guidelines for other species.

Cranial biomechanics of Diplodocus (Dinosauria, Sauropoda): testing hypotheses of feeding behaviour in an extinct megaherbivore.

Sauropod dinosaurs were the largest terrestrial herbivores and pushed at the limits of vertebrate biomechanics and physiology. Sauropods exhibit high craniodental diversity in ecosystems where numerous species co-existed, leading to the hypothesis that this biodiversity is linked to niche subdivision driven by ecological specialisation. Here, we quantitatively investigate feeding behaviour hypotheses for the iconic sauropod Diplodocus. Biomechanical modelling, using finite element analysis, was used to examine the performance of the Diplodocus skull. Three feeding behaviours were modelled: muscle-driven static biting, branch stripping and bark stripping. The skull was found to be 'over engineered' for static biting, overall experiencing low stress with only the dentition enduring high stress. When branch stripping, the skull, similarly, is under low stress, with little appreciable difference between those models. When simulated for bark stripping, the skull experiences far greater stresses, especially in the teeth and at the jaw joint. Therefore, we refute the bark-stripping hypothesis, while the hypotheses of branch stripping and/or precision biting are both consistent with our findings, showing that branch stripping is a biomechanically plausible feeding behaviour for diplodocids. Interestingly, in all simulations, peak stress is observed in the premaxillary-maxillary 'lateral plates', supporting the hypothesis that these structures evolved to dissipate stress induced while feeding. These results lead us to conclude that the aberrant craniodental form of Diplodocus was adapted for food procurement rather than resisting high bite forces.

A 3D ontogenetic atlas of Alligator mississippiensis cranial nerves and their significance for comparative neurology of reptiles.

Cranial nerves are key features of the nervous system and vertebrate body plan. However, little is known about the anatomical relationships and ontogeny of cranial nerves in crocodylians and other reptiles, hampering understanding of adaptations, evolution, and development of special senses, somatosensation, and motor control of cranial organs. Here we share three dimensional (3D) models an of the cranial nerves and cranial nerve targets of embryonic, juvenile, and adult American Alligators (Alligator mississippiensis) derived from iodine-contrast CT imaging, for the first time, exploring anatomical patterns of cranial nerves across ontogeny. These data reveal the tradeoffs of using contrast-enhanced CT data as well as patterns in growth and development of the alligator cranial nervous system. Though contrast-enhanced CT scanning allows for reconstruction of numerous tissue types in a nondestructive manner, it is still limited by size and resolution. The position of alligator cranial nerves varies little with respect to other cranial structures yet grow at different rates as the skull elongates. These data constrain timing of trigeminal and sympathetic ganglion fusion and reveal morphometric differences in nerve size and path during growth. As demonstrated by these data, alligator cranial nerve morphology is useful in understanding patterns of neurological diversity and distribution, evolution of sensory and muscular innervation, and developmental homology of cranial regions, which in turn, lead to inferences of physiology and behavior.

Dispatches from the age of crocodiles: New discoveries from ancient lineages.

Crocodilians inspire researchers and the public alike with their explosive hunting methodologies, distinct craniofacial and dental morphology, and resplendent fossil record. This special issue highlights recent advances in the biology and paleontology of this fascinating lineage of vertebrates. The authors in this volume bring crocodylians and their extinct ancestors to life using a variety of approaches including fieldwork, imaging, 3D modeling, developmental biology, physiological monitoring, dissection, and a host of other comparative methods. Our journey begins with early crocodylomorphs from the Triassic, carries us through the radiation of crocodyliforms during the rest of the Mesozoic Era, and finally celebrates the diversification development and biology of extant crocodylians. Crocodyliform science has grown appreciably the past few decades. New fossil species and genetic evidence continue to keep phylogenies and our understanding of relationships wavering in key places of the tree such as the relationships of the extinct marine thalattosuchians as well as still living species like gharials. The application of imaging approaches and 3D modeling to both preserved tissues as well as living specimens is now revealing patterns in brain and lung evolution and function, growth strategies, and feeding and locomotor behaviors across the lineage. Comparative anatomical studies are offering new data on genitals, cephalic venous drainage and thoracoabdominal pressures. The new discoveries found here only reveal there is far more work to be done to understand the biology and behavior responsible for the great radiation extinct suchians and their crocodylian descendants experienced during their conquest of Mesozoic and Tertiary ecosystems.

Ontogeny of the trigeminal system and associated structures in Alligator mississippiensis.

From the appearance of the vertebrate head, the trigeminal system has played a role in behavioral and ecological adaptation. The trigeminal nerve is the primary cranial somatosensory nerve, also innervating the jaw muscles. In crocodylians, the trigeminal nerve plays a role in modulating the high bite force and unique integumentary sensation. In association with these behaviors, crocodylians are known for large trigeminal nerves, a high volume of trigeminal-innervated musculature, and densely packed, specialized sensory receptors. These innovations also occurred in concert with a restructuring of the lateral braincase wall. These morphologies have previously been investigated in phylogenetic and evolutionary contexts, but an ontogenetic, whole-system investigation of trigeminal tissue and associated musculature, cartilage, and bone is lacking, as is an understanding of developmental timing of morphologies significant to hypotheses of homology. Here, we use contrast-enhanced computed tomography imaging to provide description and analysis of the trigeminal system in an ontogenetic series of Alligator mississippiensis from embryonic to adult form. We explore growth rates and allometric relationships of structures and discuss the significance to hypotheses of homology. We find a high growth rate and allometric trajectory of the trigeminal nerve in comparison to other cranial nerves, likely associated with the large volume of trigeminal musculature and high densities of sensory receptors. We identify a similar trend in the pterygoideus dorsalis muscle, the highest contributor to bite force. We narrow ontogenetic timing of features related to the trigeminal topological paradigm and the undeveloped epipterygoid. Overall, we provide a basis for understanding trigeminal development in crocodylians, which upon comparison across reptiles will reveal ontogenetic origins of morphological variation.

Biomechanical performance of the cranio-mandibular complex of the small notosuchian Araripesuchus gomesii (Notosuchia, Uruguaysuchidae).

Notosuchia is a clade of crocodyliforms that was highly successful and diverse in the Cretaceous of Gondwana. Araripesuchus gomesii is a small notosuchian from the Early Cretaceous of Brazil that belongs to Uruguaysuchidae, one of the subgroups of notosuchians that first radiated, during the Aptian-Albian. Here we present a finite element analysis of A. gomesii based on a model reconstructed from CT scans and performed using published bone properties for crocodiles. The adductor musculature and their respective attachment areas were reconstructed based on Extant Phylogenetic Bracket. Different functional scenarios were tested applying an estimated 158 N bite force: unilateral bite, bilateral bite, pullback, head-shake, and head-twist. The results obtained were compared with those of Alligator mississippiensis, one of its closest living relatives. In the different simulations, the skull and lower jaws of Araripesuchus suffers more stress in the head-shake movement, followed by the unilateral and pullback bites with stress focalized in the premaxillary region. In contrast, the head-twist is the one with smaller stress values. Araripesuchus possess an oreinirostral skull that may provide greater overall resistance in the different scenarios on average, unlike Alligator that has a platyrostral skull with less resistance to dorsoventral mechanical loads. Previous hypotheses that considered A. gomesii as omnivorous coupled with our results, its small size, and likely limited bite force, suggest this taxon probably fed on small prey and other trophic items that could catch and handle entirely with its mouth, such as insects and small vertebrates.

The effects of skull flattening on suchian jaw muscle evolution.

Jaw muscles are key features of the vertebrate feeding apparatus. The jaw musculature is housed in the skull whose morphology reflects a compromise between multiple functions, including feeding, housing sensory structures, and defense, and the skull constrains jaw muscle geometry. Thus, jaw muscle anatomy may be suboptimally oriented for the production of bite force. Crocodylians are a group of vertebrates that generate the highest bite forces ever measured with a flat skull suited to their aquatic ambush predatory style. However, basal members of the crocodylian line (e.g., Prestosuchus) were terrestrial predators with plesiomorphically tall skulls, and thus the origin of modern crocodylians involved a substantial reorganization of the feeding apparatus and its jaw muscles. Here, we reconstruct jaw muscles across a phylogenetic range of crocodylians and fossil suchians to investigate the impact of skull flattening on muscle anatomy. We used imaging data to create 3D models of extant and fossil suchians that demonstrate the evolution of the crocodylian skull, using osteological correlates to reconstruct muscle attachment sites. We found that jaw muscle anatomy in early fossil suchians reflected the ancestral archosaur condition but experienced progressive shifts in the lineage leading to Metasuchia. In early fossil suchians, musculus adductor mandibulae posterior and musculus pterygoideus (mPT) were of comparable size, but by Metasuchia, the jaw musculature is dominated by mPT. As predicted, we found that taxa with flatter skulls have less efficient muscle orientations for the production of high bite force. This study highlights the diversity and evolution of jaw muscles in one of the great transformations in vertebrate evolution.

New frontiers in imaging, anatomy, and mechanics of crocodylian jaw muscles.

New imaging and biomechanical approaches have heralded a renaissance in our understanding of crocodylian anatomy. Here, we review a series of approaches in the preparation, imaging, and functional analysis of the jaw muscles of crocodylians. Iodine-contrast microCT approaches are enabling new insights into the anatomy of muscles, nerves, and other soft tissues of embryonic as well as adult specimens of alligators. These imaging data and other muscle modeling methods offer increased accuracy of muscle sizes and attachments without destructive methods like dissection. 3D modeling approaches and imaging data together now enable us to see and reconstruct 3D muscle architecture which then allows us to estimate 3D muscle resultants, but also measurements of pennation in ways not seen before. These methods have already revealed new information on the ontogeny, diversity, and function of jaw muscles and the heads of alligators and other crocodylians. Such approaches will lead to enhanced and accurate analyses of form, function, and evolution of crocodylians, their fossil ancestors and vertebrates in general.

The impact of bone and suture material properties on mandibular function in Alligator mississippiensis: testing theoretical phenotypes with finite element analysis.

The functional effects of bone and suture stiffness were considered here using finite element models representing three different theoretical phenotypes of an Alligator mississippiensis mandible. The models were loaded using force estimates derived from muscle architecture in dissected specimens, constrained at the 18th and 19th teeth in the upper jaw and 19th tooth of the lower jaw, as well as at the quadrate-articular joint. Stiffness was varied systematically in each theoretical phenotype. The three theoretical phenotypes included: (i) linear elastic isotropic bone of varying stiffness and no sutures; (ii) linear elastic orthotropic bone of varying stiffness with no sutures; and (iii) linear elastic isotropic bone of a constant stiffness with varying suture stiffness. Variation in the isotropic material properties of bone primarily resulted in changes in the magnitude of principal strain. By comparison, variation in the orthotropic material properties of bone and isotropic material properties of sutures resulted in: a greater number of bricks becoming either more compressive or more tensile, changing between being either dominantly compressive or tensile, and having larger changes in the orientation of maximum principal strain. These data indicate that variation in these model properties resulted in changes to the strain regime of the model, highlighting the importance of using biologically verified material properties when modeling vertebrate bones. When bones were compared within each set, the response of each to changing material properties varied. In two of the 12 bones in the mandible, varied material properties within sutures resulted in a decrease in the magnitude of principal strain in bricks adjacent to the bone/suture interface and decreases in stored elastic energy. The varied response of the mandibular bones to changes in suture stiffness highlights the importance of defining the appropriate functional unit when addressing relationships of performance and morphology.

More than one way to be a giant: Convergence and disparity in the hip joints of saurischian dinosaurs.

Saurischian dinosaurs evolved seven orders of magnitude in body mass, as well as a wide diversity of hip joint morphology and locomotor postures. The very largest saurischians possess incongruent bony hip joints, suggesting that large volumes of soft tissues mediated hip articulation. To understand the evolutionary trends and functional relationships between body size and hip anatomy of saurischians, we tested the relationships among discrete and continuous morphological characters using phylogenetically corrected regression. Giant theropods and sauropods convergently evolved highly cartilaginous hip joints by reducing supraacetabular ossifications, a condition unlike that in early dinosauromorphs. However, transitions in femoral and acetabular soft tissues indicate that large sauropods and theropods built their hip joints in fundamentally different ways. In sauropods, the femoral head possesses irregularly rugose subchondral surfaces for thick hyaline cartilage. Hip articulation was achieved primarily using the highly cartilaginous femoral head and the supraacetabular labrum on the acetabular ceiling. In contrast, theropods covered their femoral head and neck with thinner hyaline cartilage and maintained extensive articulation between the fibrocartilaginous femoral neck and the antitrochanter. These findings suggest that the hip joints of giant sauropods were built to sustain large compressive loads, whereas those of giant theropods experienced compression and shear forces.

The Frontoparietal Fossa and Dorsotemporal Fenestra of Archosaurs and Their Significance for Interpretations of Vascular and Muscular Anatomy in Dinosaurs.

The attachments of jaw muscles are typically implicated in the evolution and shape of the dorsotemporal fenestra on the skull roof of amniotes. However, the dorsotemporal fenestrae of many archosaurian reptiles possess smooth excavations rostral and dorsal to the dorsotemporal fossa which closely neighbors the dorsotemporal fenestra and jaw muscle attachments. Previous research has typically identified this region, here termed the frontoparietal fossa, to also have attachment surfaces for jaw-closing muscles. However, numerous observations of extant and extinct archosaurs described here suggest that other tissues are instead responsible for the size and shape of the frontoparietal fossa. This study reviewed the anatomical evidence that support soft-tissue hypotheses of the frontoparietal fossa and its phylogenetic distribution among sauropsids. Soft-tissue hypotheses (i.e., muscle, pneumatic sinus, vascular tissues) were analyzed using anatomical, imaging and in vivo thermography techniques within a phylogenetic framework using extant and extinct taxa to determine the inferential power underlying the reconstruction of the soft tissues in the skull roofs of dinosaurs, pseudosuchians, and other reptiles. Relevant anatomical features argue for rejection of the default hypothesis-that the fossa was muscular-due to a complete lack of osteological correlates reflective of muscle attachment. The most-supported inference of soft tissues is that the frontoparietal fossa contained a large vascular structure and adipose tissue. Despite the large sizes and diverse morphologies of these fossae found among dinosaur taxa, these data suggest that non-avian dinosaurs had the anatomical foundation to support physiologically significant vascular devices and/or vascular integumentary structures on their skull roofs. Anat Rec, 303:1060-1074, 2020. © 2019 Wiley Periodicals, Inc.