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  1. Abstract

    Eopneumatosuchus colbertiCrompton and Smith, 1980, known from a single partial skull, is an enigmatic crocodylomorph from the Lower Jurassic Kayenta Formation. In spite of its unique morphology, an exceptionally pneumatic braincase, and presence during a critical time period of crocodylomorph evolution, relatively little is known about this taxon. Here, we redescribe the external cranial morphology ofE.colberti, present novel information on its endocranial anatomy, evaluate its phylogenetic position among early crocodylomorphs, and seek to better characterize its ecology. Our examination clarifies key aspects of cranial suture paths and braincase anatomy. Comparisons with related taxa (e.g.,Protosuchus haughtoni) demonstrate that extreme pneumaticity of the braincase may be more widespread in protosuchids than previously appreciated. Computed tomography scans reveal an endocranial morphology that resembles that of other early crocodylomorphs, in particular the noncrocodyliform crocodylomorphAlmadasuchus figarii. There are, however, key differences in olfactory bulb and cerebral hemisphere morphology, which demonstrate the endocranium of crocodylomorphs is not as conserved as previously hypothesized. Our phylogenetic analysis recoversE.colbertias a close relative ofProtosuchus richardsoniandEdentosuchus tienshanensis, contrasting with previous hypotheses of a sister group relationship with Thalattosuchia. Previous work suggested the inner ear has some similarities to semi‐aquatic crocodyliforms, but the phylogenetic placement ofE.colbertiamong protosuchids with a terrestrial postcranial skeletal morphology complicates paleoecological interpretation.

     
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  2. Abstract

    Thalattosuchians represent one of the several independent transitions into the marine realm among crocodylomorphs. The extent of their aquatic adaptations ranges from the semiaquatic teleosauroids, superficially resembling extant gharials, to the almost cetacean‐like pelagic metriorhynchids. Understanding the suite of osteological, physiological, and sensory changes that accompanied this major transition has received increased attention, but is somewhat hindered by a dearth of complete three‐dimensionally preserved crania. Here, we describe the cranial and endocranial anatomy of a well‐preserved three‐dimensional specimen ofMacrospondylus bollensisfrom the Toarcian of Yorkshire, UK. The trigeminal fossa contains two similar‐sized openings separated by a thin lamina of prootic, a configuration that appears unique to a subset of teleosauroids.Macrospondylus bollensisresembles other thalattosuchians in having pyramidal semicircular canals with elongate cochlear ducts, enlarged carotid canals leading to an enlarged pituitary fossa, enlarged orbital arteries, enlarged endocranial venous sinuses, reduced pharyngotympanic sinuses, and a relatively straight brain with a hemispherical cerebral expansion. We describe for the first time the olfactory region and paranasal sinuses of a teleosauroid. A relatively large olfactory region suggests greater capacity for airborne olfaction in teleosauroids than in the more aquatically adapted metriorhynchoids. Additionally, slight swellings in the olfactory region suggest the presence of small salt glands of lower secretory capacity than those of metriorhynchoids. The presence of osteological correlates for salt glands in a teleosauroid corroborates previous hypotheses that these glands originated in the common ancestor of Thalattosuchia, facilitating their rapid radiation into the marine realm.

     
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  3. Abstract

    Caimaninae is one of the few crocodylian lineages that still has living representatives. Today, most of its six extant species are restricted to South and Central America. However, recent discoveries have revealed a more complex evolutionary history, with a fossil record richer than previously thought and a possible North American origin. Among the oldest caimanines isEocaiman cavernensis, from the Eocene of Patagonia, Argentina. It was described by George G. Simpson in the 1930s, representing the first caimanine reported for the Palaeogene. Since then,E. cavernensishas been ubiquitous in phylogenetic studies on the group, but a more detailed morphological description and revision of the taxon were lacking. Here, we present a reassessment ofE. cavernensis, based on first‐hand examination and micro‐computed tomography of the holotype, and reinterpret different aspects of its morphology. We explore the phylogenetic affinities ofE. cavernensisand other caimanines using parsimony and Bayesian inference approaches. Our results provide evidence for a monophyleticEocaimangenus within Caimaninae, even though some highly incomplete taxa (including the congenericEocaiman itaboraiensis) represent significant sources of phylogenetic instability. We also foundCulebrasuchus mesoamericanusas sister to all other caimanines and the North American globidontans (i.e.Brachychampsaand closer relatives) outside Caimaninae. A time‐calibrated tree, obtained using a fossilized birth–death model, shows a possible Campanian origin for the group (76.97 ± 6.7 Ma), which is older than the age estimated using molecular data, and suggests that the earliest cladogenetic events of caimanines took place rapidly and across the K–Pg boundary.

     
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  4. Abstract

    Theropod dinosaurs were relatively scarce in the Late Cretaceous ecosystems of southeast Brazil. Instead, hypercarnivorous crocodyliforms known as baurusuchids were abundant and probably occupied the ecological role of apex predators. Baurusuchids exhibited a series of morphological adaptations hypothesized to be associated with this ecological role, but quantitative biomechanical analyses of their morphology have so far been lacking. Here, we employ a biomechanical modelling approach, applying finite element analysis (FEA) to models of the skull and mandibles of a baurusuchid specimen. This allows us to characterize the craniomandibular apparatus of baurusuchids, as well as to compare the functional morphology of the group with that of other archosaurian carnivores, such as theropods and crocodylians. Our results support the ecological role of baurusuchids as specialized apex predators in the continental Late Cretaceous ecosystems of South America. With a relatively weak bite force (~600 N), the predation strategies of baurusuchids likely relied on other morphological specializations, such as ziphodont dentition and strong cervical musculature. Comparative assessments of the stress distribution and magnitude of scaled models of other predators (the theropodAllosaurus fragilisand the living crocodylianAlligator mississippiensis) consistently show different responses to loadings under the same functional scenarios, suggesting distinct predatory behaviors for these animals. The unique selective pressures in the arid to semi‐arid Late Cretaceous ecosystems of southeast Brazil, which were dominated by crocodyliforms, possibly drove the emergence and evolution of the biomechanical features seen in baurusuchids, which are distinct from those previously reported for other predatory taxa.

     
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  5. Abstract

    Extant crocodylomorphs are semiaquatic ambush predators largely restricted to freshwater or estuarine environments, but the group is ancestrally terrestrial and inhabited a variety of ecosystems in the past. Despite its rich ecological history, little effort has focused on elucidating the historical pattern of ecological transitions in the group. Traditional views suggested a single shift from terrestrial to aquatic in the Early Jurassic. However, new fossil discoveries and phylogenetic analyses tend to imply a multiple-shift model. Here we estimate ancestral habitats across a comprehensive phylogeny and show at least three independent shifts from terrestrial to aquatic and numerous other habitat transitions. Neosuchians first invade freshwater habitats in the Jurassic, with up to four subsequent shifts into the marine realm. Thalattosuchians first appear in marine habitats in the Early Jurassic. Freshwater semiaquatic mahajangasuchids are derived from otherwise terrestrial notosuchians. Within nearly all marine groups, some species return to freshwater environments. Only twice have crocodylomorphs reverted from aquatic to terrestrial habitats, both within the crown group. All living non-alligatorid crocodylians have a keratinised tongue with salt-excreting glands, but the lack of osteological correlates for these adaptations complicates pinpointing their evolutionary origin or loss. Based on the pattern of transitions to the marine realm, our analysis suggests at least four independent origins of saltwater tolerance in Crocodylomorpha.

     
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  6. Abstract

    Cranial endocasts, or the internal molds of the braincase, are a crucial correlate for investigating the neuroanatomy of extinct vertebrates and tracking brain evolution through deep time. Nevertheless, the validity of such studies pivots on the reliability of endocasts as a proxy for brain morphology. Here, we employ micro‐computed tomography imaging, including diffusible iodine‐based contrast‐enhancedCT, and a three‐dimensional geometric morphometric framework to examine both size and shape differences between brains and endocasts of two exemplar archosaur taxa – the American alligator (Alligator mississippiensis) and the domestic chicken (Gallus gallus). With ontogenetic sampling, we quantitatively evaluate how endocasts differ from brains and whether this deviation changes during development. We find strong size and shape correlations between brains and endocasts, divergent ontogenetic trends in the brain‐to‐endocast correspondence between alligators and chickens, and a comparable magnitude between brain–endocast shape differences and intraspecific neuroanatomical variation. The results have important implications for paleoneurological studies in archosaurs. Notably, we demonstrate that the pattern of endocranial shape variation closely reflects brain shape variation. Therefore, analyses of endocranial morphology are unlikely to generate spurious conclusions about large‐scale trends in brain size and shape. To mitigate any artifacts, however, paleoneurological studies should consider the lower brain–endocast correspondence in the hindbrain relative to the forebrain; higher size and shape correspondences in chickens than alligators throughout postnatal ontogeny; artificially ‘pedomorphic’ shape of endocasts relative to their corresponding brains; and potential biases in both size and shape data due to the lack of control for ontogenetic stages in endocranial sampling.

     
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  7. Effective interpretation of historical selective regimes requires comprehensive in vivo performance evaluations and well-constrained ecomorphological prox- ies. The feeding apparatus is a frequent target of such evolutionary studies due to a direct relationship between feeding and survivorship, and the durability of craniodental elements in the fossil record. Among vertebrates, behaviors such as bite force have been central to evaluation of clade dynamics; yet, in the absence of detailed performance studies, such evaluations can misidentify potential selective factors and their roles. Here, we combine the results of a total-clade performance study with fossil-inclusive, phylogenetically informed methods to assess bite-force proxies throughout mesoeucrocodylian evolution. Although bite-force shifts were previously thought to respond to changing rostrodental selective regimes, we find body-size dependent conservation of performance proxies throughout the history of the clade, indicating stabilizing selection for bite-force potential. Such stasis reveals that mesoeucrocodylians with dietary ecologies as disparate as herbivory and hypercarnivory maintain similar bite-force-to-body-size relationships, a pattern which contrasts the pre- cept that vertebrate bite forces should vary most strongly by diet. Furthermore, it may signal that bite-force conservation supported mesoeucrocodylian craniodental disparity by providing a stable performance foundation for the exploration of novel ecomorphospace. 
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  8. null (Ed.)
    Relative brain size has long been considered a reflection of cognitive capacities and has played a fundamental role in developing core theories in the life sciences. Yet, the notion that relative brain size validly represents selection on brain size relies on the untested assumptions that brain-body allometry is restrained to a stable scaling relationship across species and that any deviation from this slope is due to selection on brain size. Using the largest fossil and extant dataset yet assembled, we find that shifts in allometric slope underpin major transitions in mammalian evolution and are often primarily characterized by marked changes in body size. Our results reveal that the largest-brained mammals achieved large relative brain sizes by highly divergent paths. These findings prompt a reevaluation of the traditional paradigm of relative brain size and open new opportunities to improve our understanding of the genetic and developmental mechanisms that influence brain size. 
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  9. null (Ed.)
    Crocodylians are currently facing evolutionary decline. This is evinced by the rich fossil record of their extinct relatives, crocodylomorphs, which show not only significantly higher levels of biodiversity in the past but also remarkable morphological disparity and higher ecological diversity. In terms of body size, crocodylians are mostly large animals (>2m), especially when com-pared to other extant reptiles. In contrast, extinct crocodylomorphs exhibited a 10-fold range in body sizes, with early terrestrial forms often quite small.Recent research has shed new light on the tempo and mode of crocodylomorph body size evolution,demonstrating a close relationship with ecology, in which physiological constraints contribute to the larger sizes of marine species. Abiotic environmental factors can also play an important role with in individual subgroups. Crocodylians, for instance,have been experiencing an average size increase during Cenozoic, which seems to be related to along-term process of global cooling. 
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  10. 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.

     
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