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The nasal passage performs multiple functions in amniotes, including olfaction and thermoregulation. These functions would have been present in extinct animals as well. However, fossils preserve only low‐resolution versions of the nasal passage due to loss of soft‐tissue structures after death. To test the effects of these lower resolution models on interpretations of nasal physiology, we performed a broadly comparative analysis of the nasal passages in extant diapsid representatives, e.g., alligator, turkey, ostrich, iguana, and a monitor lizard. Using computational fluid dynamics, we simulated airflow through 3D reconstructed models of the different nasal passages and compared these soft‐tissue‐bounded results to similar analyses of the same airways under the lower‐resolution limits imposed by fossilization. Airflow patterns in these bony‐bounded airways were more homogeneous and slower flowing than those of their soft‐tissue counterparts. These data indicate that bony‐bounded airway reconstructions of extinct animal nasal passages are far too conservative and place overly restrictive physiological limitations on extinct species. In spite of the diverse array of nasal passage shapes, distinct similarities in airflow were observed, including consistent areas of nasal passage constriction such as the junction of the olfactory region and main airway. These nasal constrictions can reasonably be inferred to have been present in extinct taxa such as dinosaurs.

 

Secondarily marine tetrapod lineages have independently evolved osmoregulatory adaptations for life in salt water but inferring physiological changes in extinct marine tetrapods is difficult. The Mesozoic crocodylomorph clade Thalattosuchia is unique in having both direct evidence from natural endocasts and several proposed osteological correlates for salt exocrine glands. Here, we investigate salt gland evolution in thalattosuchians by creating endocranial reconstructions from CT scans of eight taxa (one basal thalattosuchian, one teleosauroid, two basal metriorhynchoids and four metriorhynchids) and four outgroups (three extant crocodylians and the basal crocodyliform Protosuchus) to identify salt gland osteological correlates. All metriorhynchoids show dorsolateral nasal cavity expansions corresponding to the location of nasal salt glands in natural casts, but smaller expansions in teleosauroids correspond more with the cartilaginous nasal capsule. The different sizes of these expansions suggest the following evolutionary sequence: (1) plesiomorphically small glands present in semi-aquatic teleosauroids draining through the nasal vestibule; (2) moderately sized glands in the basalmost metriorhynchoid Pelagosaurus; and (3) hypertrophied glands in the clade comprising Eoneustes and metriorhynchids, with a pre-orbital fenestra providing a novel exit for salt drainage. The large gland size inferred from basal metriorhynchoids indicates advanced osmoregulation occurred while metriorhynchoids were semi-aquatic. This pattern does not precisely fit into current models of physiological evolution in marine tetrapods and suggests a unique sequence of changes as thalattosuchians transitioned from land to sea.

 

Abstract As computed tomography and related technologies have become mainstream tools across a broad range of scientific applications, each new generation of instrumentation produces larger volumes of more-complex 3D data. Lagging behind are step-wise improvements in computational methods to rapidly analyze these new large, complex datasets. Here we describe novel computational methods to capture and quantify volumetric information, and to efficiently characterize and compare shape volumes. It is based on innovative theoretical and computational reformulation of volumetric computing. It consists of two theoretical constructs and their numerical implementation: the spherical wave decomposition ( SWD ), that provides fast, accurate automated characterization of shapes embedded within complex 3D datasets; and symplectomorphic registration with phase space regularization by entropy spectrum pathways ( SYMREG ), that is a non-linear volumetric registration method that allows homologous structures to be correctly warped to each other or a common template for comparison. Together, these constitute the Shape Analysis for Phenomics from Imaging Data ( SAPID ) method. We demonstrate its ability to automatically provide rapid quantitative segmentation and characterization of single unique datasets, and both inter-and intra-specific comparative analyses. We go beyond pairwise comparisons and analyze collections of samples from 3D data repositories, highlighting the magnified potential our method has when applied to data collections. We discuss the potential of SAPID in the broader context of generating normative morphologies required for meaningfully quantifying and comparing variations in complex 3D anatomical structures and systems. 

Although the visual system of crocodylians has attracted interest regarding optical parameters and retinal anatomy, fundamental questions remain about the allometry of the eyeball and whether such scaling is the same across all crown groups of crocodylians. In addition, anatomy and identities of adnexal soft tissues that interact with the visual system are not well understood in many cases. We used contrast‐enhancing iodine stain and high‐resolution micro‐computed tomography to assess the anatomy of orbital soft tissues, including extraocular muscles and glands, in crocodylians. We also used regression analysis to estimate the allometric relationship between the bony orbit and eyeball acrossAlligator mississippiensisandCrocodylus niloticusfor the first time. Results revealed tight, negatively allometric relationships between the bony orbit and eyeball. Notably, the eyes ofC. niloticuswere larger for a given orbit size than the eyes ofA. mississippiensis, although the slope of the relationship was no different between these two crown crocodylian groups. Among the findings from our anatomical study, new details were uncovered about the homologies of muscles of the abducens complex. In particular, M. rectus lateralis of crocodylians is revealed to have a more complex form than previously appreciated, being adhered to the tendon of the nictitating membrane, which may be apomorphic for Crocodylia. Our calculation of the orbit‐eyeball allometric relationship and study of the adnexal soft tissues of the crocodylian visual system, in combination with previous work by other teams in other crown saurian clades, is a critical, formerly missing, piece in the Extant Phylogenetic Bracket for restoring the visual apparatus of extinct crocodyliforms and other archosauriform groups.

 

Body size has thermal repercussions that impact physiology. Large‐bodied dinosaurs potentially retained heat to the point of reaching dangerous levels, whereas small dinosaurs shed heat relatively easily. Elevated body temperatures are known to have an adverse influence on neurosensory tissues and require physiological mechanisms for selective brain and eye temperature regulation. Vascular osteological correlates in fossil dinosaur skulls from multiple clades representing different body‐size classes were identified and compared. Neurovascular canals were identified that differentiate thermoregulatory strategies involving three sites of evaporative cooling that are known in extant diapsids to function in selective brain temperature regulation. Small dinosaurs showed similarly sized canals, reflecting a plesiomorphic balanced pattern of blood supply and a distributed thermoregulatory strategy with little evidence of enhancement of any sites of thermal exchange. Large dinosaurs, however, showed a more unbalanced vascular pattern whereby certain sites of thermal exchange were emphasized for enhanced blood flow, reflecting a more focused thermal strategy. A quantitative, statistical analysis of canal cross‐sectional area was conducted to test these anatomical results, confirming that large‐bodied, and often large‐headed, species showed focused thermal strategies with enhanced collateral blood flow to certain sites of heat exchange. Large theropods showed evidence for a plesiomorphic balanced blood flow pattern, yet evidence for vascularization of the large antorbital paranasal air sinus indicates theropods may have had a fourth site of heat exchange as part of a novel focused thermoregulatory strategy. Evidence presented here for differing thermoregulatory strategies based on size and phylogeny helps refine our knowledge of dinosaur physiology. Anat Rec, 303:1075–1103, 2020. © 2019 American Association for Anatomy

 

Endocasts are increasingly relied upon to examine avian brain evolution because they can be used across extant and extinct species. The endocasts of birds appear to be relatively faithful representatives of the external morphology of their brains, but it is unclear how well the size of a surface feature visible on endocasts reflects the volume of the underlying brain region. The optic lobe and the Wulst are two endocast structures that are clearly visible on the external surface of avian endocasts. As they overlie two major visual regions of the brain, the optic tectum and hyperpallium, the surface areas of the optic lobe and Wulst, respectively, are often used to infer visual abilities. To determine whether the surface area of these features reflects the volume of the underlying brain regions, we compared the surface areas of the optic lobes and Wulsts from digital endocasts with the volumes of the optic tecta and hyperpallia from the literature or measured from histological series of brains of the same species. Regression analyses revealed strong, statistically significant correlations between the volumes of the brain regions and the surface areas of the overlying endocast structures. In other words, the size of the hyperpallium and optic tectum can be reliably inferred from the surface areas of the Wulst and optic lobe, respectively. This validation opens the possibility of estimating brain‐region volumes for extinct species in order to gain better insights in their visual ecology. It also emphasizes the importance of adopting a quantitative approach to the analysis of endocasts in the study of brain evolution.

 

Major evolutionary transitions, in which animals develop new body plans and adapt to dramatically new habitats and lifestyles, have punctuated the history of life. The origin of cetaceans from land-living mammals is among the most famous of these events. Much earlier, during the Mesozoic Era, many reptile groups also moved from land to water, but these transitions are more poorly understood. We use computed tomography to study changes in the inner ear vestibular system, involved in sensing balance and equilibrium, as one of these groups, extinct crocodile relatives called thalattosuchians, transitioned from terrestrial ancestors into pelagic (open ocean) swimmers. We find that the morphology of the vestibular system corresponds to habitat, with pelagic thalattosuchians exhibiting a more compact labyrinth with wider semicircular canal diameters and an enlarged vestibule, reminiscent of modified and miniaturized labyrinths of other marine reptiles and cetaceans. Pelagic thalattosuchians with modified inner ears were the culmination of an evolutionary trend with a long semiaquatic phase, and their pelagic vestibular systems appeared after the first changes to the postcranial skeleton that enhanced their ability to swim. This is strikingly different from cetaceans, which miniaturized their labyrinths soon after entering the water, without a prolonged semiaquatic stage. Thus, thalattosuchians and cetaceans became secondarily aquatic in different ways and at different paces, showing that there are different routes for the same type of transition.

 

Metriorhynchoid thalattosuchians were a marine clade of Mesozoic crocodylomorphs that evolved from semi‐aquatic, “gharial”‐like species into the obligately pelagic subclade Metriorhynchidae. To explore whether the sensory and physiological demands of underwater life necessitates a shift in rostral anatomy, both in neurology and vasculature, we investigate the trigeminal innervation and potential somatosensory abilities of metriorhynchoids by digitally segmenting the rostral neurovascular canals in CT scans of 10 extant and extinct crocodyliforms. The dataset includes the terrestrial, basal crocodyliformProtosuchus haughtoni, two semi‐aquatic basal metriorhynchoids, four pelagic metriorhynchids and three extant, semi‐aquatic crocodylians. In the crocodylian and basal metriorhynchoid taxa, we find three main neurovascular channels running parallel to one another posteroanteriorly down the length of the snout, whereas in metriorhynchids there are two, and inP. haughtonionly one. Crocodylians appear to be unique in their extensive trigeminal innervation, which is used to supply the integumentary sensory organs (ISOs) involved with their facial somatosensory abilities. Crocodylians have a far higher number of foramina on the maxillary bones than either metriorhynchoids orP. haughtoni, suggesting that the fossil taxa lacked the somatosensory abilities seen in extant species. We posit that the lack of ISO osteological correlates in metriorhynchoids is due to their basal position in Crocodyliformes, rather than a pelagic adaptation. This is reinforced by the hypothesis that extant crocodyliforms, and possibly some neosuchian clades, underwent a long “nocturnal bottleneck”—hinting that their complex network of ISOs evolved in Neosuchia, as a sensory trade‐off to compensate for poorer eyesight.

 

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 andin vivothermography 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.