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Attempts to explain the origin and diversification of vertebrates have commonly invoked the evolution of feeding ecology, contrasting the passive suspension feeding of invertebrate chordates and larval lampreys with active predation in living jawed vertebrates. Of the extinct jawless vertebrates that phylogenetically intercalate these living groups, the feeding apparatus is well-preserved only in the early diverging stem-gnathostome heterostracans. However, its anatomy remains poorly understood. Here, we use X-ray microtomography to characterize the feeding apparatus of the pteraspid heterostracan Rhinopteraspis dunensis (Roemer, 1855). The apparatus is composed of 13 plates arranged approximately bilaterally, most of which articulate from the postoral plate. Our reconstruction shows that the oral plates were capable of rotating around the transverse axis, but likely with limited movement. It also suggests the nasohypophyseal organs opened internally, into the pharynx. The functional morphology of the apparatus in Rhinopteraspis precludes all proposed interpretations of feeding except for suspension/deposit feeding and we interpret the apparatus as having served primarily to moderate the oral gape. This is consistent with evidence that at least some early jawless gnathostomes were suspension feeders and runs contrary to macroecological scenarios that envisage early vertebrate evolution as characterized by a directional trend towards increasingly active food acquisition.

Skeletal simplification occurred in multiple vertebrate clades over the last 500 million years, including the evolution from premammalian cynodonts to mammals. This transition is characterised by the loss and reduction of cranial bones, the emergence of a novel jaw joint, and the rearrangement of the jaw musculature. These modifications have long been hypothesised to increase skull strength and efficiency during feeding. Here, we combine digital reconstruction and biomechanical modelling to show that there is no evidence for an increase in cranial strength and biomechanical performance. Our analyses demonstrate the selective functional reorganisation of the cranial skeleton, leading to reduced stresses in the braincase and the skull roof but increased stresses in the zygomatic region through this transition. This cranial functional reorganisation, reduction in mechanical advantage, and overall miniaturisation in body size are linked with a dietary specialisation to insectivory, permitting the subsequent morphological and ecological diversification of the mammalian lineage., (© 2023. The Author(s).)

The extent to which evolution is deterministic is a key question in biology, 1 , 2 , 3 , 4 , 5 , 6 , 7 , 8 , 9 with intensive debate on how adaptation 6 , 10 , 11 , 12 , 13 and constraints 14 , 15 , 16 might canalize solutions to ecological challenges. 4 , 5 , 6 Alternatively, unique adaptations 1 , 9 , 17 and phylogenetic contingency 1 , 3 , 18 may render evolution fundamentally unpredictable. 3 Information from the fossil record is critical to this debate, 1 , 2 , 11 but performance data for extinct taxa are limited. 7 This knowledge gap is significant, as general morphology may be a poor predictor of biomechanical performance. 17 , 19 , 20 High-fiber herbivory originated multiple times within ornithischian dinosaurs, 21 making them an ideal clade for investigating evolutionary responses to similar ecological pressures. 22 However, previous biomechanical modeling studies on ornithischian crania 17 , 23 , 24 , 25 have not compared early-diverging taxa spanning independent acquisitions of herbivory. Here, we perform finite-element analysis on the skull of five early-diverging members of the major ornithischian clades to characterize morphofunctional pathways to herbivory. Results reveal limited functional convergence among ornithischian clades, with each instead achieving comparable performance, in terms of reconstructed patterns and magnitudes of functionally induced stress, through different adaptations of the feeding apparatus. Thyreophorans compensated for plesiomorphic low performance through increased absolute size, heterodontosaurids expanded jaw adductor muscle volume, ornithopods increased jaw system efficiency, and ceratopsians combined these approaches. These distinct solutions to the challenges of herbivory within Ornithischia underpinned the success of this diverse clade. Furthermore, the resolution of multiple solutions to equivalent problems within a single clade through macroevolutionary time demonstrates that phenotypic evolution is not necessarily predictable, instead arising from the interplay of adaptation, innovation, contingency, and constraints. 1 , 2 , 3 , 7 , 8 , 9 , 18 ., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2022 The Author(s). Published by Elsevier Inc. All rights reserved.)

Proa valdearinnoensis is a relatively large-headed and stocky iguanodontian dinosaur from the latest Early Cretaceous of Spain. Its braincase is known from three specimens. Similar to that of other dinosaurs, it shows a mosaic ossification pattern in which most of the bones seem to have fused together indistinguishably while a few (frontoparietal, basioccipital) might have remained loosely attached. The endocasts of the three specimens are described based on CT data and digital reconstructions. They show unmistakable morphological similarities with the endocast of closely related taxa, such as Sirindhorna khoratensis (which is close in age but from Thailand). This supports a high conservatism of the endocranial cavity. The issue of volumetric correspondence between endocranial cavity and brain in dinosaurs is analyzed. Although a brain-to-endocranial cavity (BEC) index of 0.50 has been traditionally used, we employ instead 0.73. This is indeed the mid-value between the situation in adults of Alligator mississippiensis and Gallus gallus, which are members of the extant bracketing taxa of dinosaurs (Crocodilia and Aves). We thence gauge the level of encephalization of P. valdearinnoensis through the calculation of the encephalization quotient (EQ), which remains valuable as a metric for assessing the degree of cognitive function in extinct taxa, especially those with fully ossified braincases like dinosaurs and other archosaurs. The EQ obtained for P. valdearinnoensis (3.611) suggests that this species was significantly more encephalized than most if not all extant nonavian, nonmammalian amniotes. Our work adds to the growing body of data concerning theoretical cognitive capabilities in dinosaurs and supports the idea that an increasing encephalization was fostered not only in theropods but also in parallel in the shorter-lived lineage of ornithopods. P. valdearinnoensis was ill-equipped to respond to theropod dinosaurs and possibly lived in groups as a strategy to mitigate the risk of being predated upon. We hypothesize that group-living and protracted caring of juveniles in this and possibly many other iguanodontian ornithopods favored a degree of encephalization that was outstanding by reptile standards., (© 2021 Wiley Periodicals LLC.)

Pterosaurs, which lived during the Mesozoic, were the first known vertebrates to evolve powered flight. 1 , 2 Arboreal locomotion has been proposed for some taxa, 3 , 4 and even considered to have played a role in the origin of pterosaur flight. 5 , 6 Even so, there is still need for comprehensive quantitative ecomorphological analyses. 3 , 4 Furthermore, skeletal adaptations correlated to specialized lifestyles are often difficult to recognize and interpret in fossils. Here we report on a new darwinopteran pterosaur that inhabited a unique forest ecosystem from the Jurassic of China. The new species exhibits the oldest record of palmar (or true) opposition of the pollex, which is unprecedented for pterosaurs and represents a sophisticated adaptation related to arboreal locomotion. Principal-coordinate analyses suggest an arboreal lifestyle for the new species but not for other closely related species from the same locality, implying a possible case of ecological niche partitioning. The discovery adds to the known array of pterosaur adaptations and the history of arborealism in vertebrates. It also adds to the impressive early bloom of arboreal communities in the Jurassic of China, shedding light on the history of forest environments., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2021 Elsevier Inc. All rights reserved.)

The acquisition of elongated, sabre-like canines in multiple vertebrate clades during the last 265 Myr represents a remarkable example for convergent evolution. Due to striking superficial similarities in the cranial skeleton, the same or similar skull and jaw functions have been inferred for sabre-toothed species and interpreted as an adaptation to subdue large-bodied prey. However, although some sabre-tooth lineages have been classified into different ecomorphs (dirk-tooths and scimitar-tooths) the functional diversity within and between groups and the evolutionary paths leading to these specializations are unknown. Here, we use a suite of biomechanical simulations to analyse key functional parameters (mandibular gape angle, bending strength, bite force) to compare the functional performance of different groups and to quantify evolutionary rates across sabre-tooth vertebrates. Our results demonstrate a remarkably high functional diversity between sabre-tooth lineages and that different cranial function and prey killing strategies evolved within clades. Moreover, different biomechanical adaptations in coexisting sabre-tooth species further suggest that this functional diversity was at least partially driven by niche partitioning.

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 theropod Allosaurus fragilis and the living crocodylian Alligator 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., (© 2020 Anatomical Society.)

The origin of turtles is one of the most long-lasting debates in evolutionary research. During their evolution, a series of modifications changed their relatively kinetic and anapsid skull into an elongated akinetic structure with a unique pulley system redirecting jaw adductor musculature. These modifications were thought to be strongly correlated to functional adaptations, especially to bite performance. We conducted a series of Finite Element Analyses (FEAs) of several species, including that of the oldest fully shelled, Triassic stem-turtle Proganochelys, to evaluate the role of force distribution and to test existing hypotheses on the evolution of turtle skull architecture. We found no support for a relation between the akinetic nature of the skull or the trochlear mechanisms with increased bite forces. Yet, the FEAs show that those modifications changed the skull architecture into an optimized structure, more resistant to higher loads while allowing material reduction on specific regions. We propose that the skull of modern turtles is the result of a complex process of progressive correlation between their heads and highly flexible necks, initiated by the origin of the shell.

Ichthyosaurs are an extinct group of fully marine tetrapods that were well adapted to aquatic locomotion. During their approximately 160 Myr existence, they evolved from elongate and serpentine forms into stockier, fish-like animals, convergent with sharks and dolphins. Here, we use computational fluid dynamics (CFD) to quantify the impact of this transition on the energy demands of ichthyosaur swimming for the first time. We run computational simulations of water flow using three-dimensional digital models of nine ichthyosaurs and an extant functional analogue, a bottlenose dolphin, providing the first quantitative evaluation of ichthyosaur hydrodynamics across phylogeny. Our results show that morphology did not have a major effect on the drag coefficient or the energy cost of steady swimming through geological time. We show that even the early ichthyosaurs produced low levels of drag for a given volume, comparable to those of a modern dolphin, and that deep 'torpedo-shaped' bodies did not reduce the cost of locomotion. Our analysis also provides important insight into the choice of scaling parameters for CFD applied to swimming mechanics, and underlines the great influence of body size evolution on ichthyosaur locomotion. A combination of large bodies and efficient swimming modes lowered the cost of steady swimming as ichthyosaurs became increasingly adapted to a pelagic existence.

The evolution of the mammalian jaw is one of the most important innovations in vertebrate history, and underpins the exceptional radiation and diversification of mammals over the last 220 million years 1,2 . In particular, the transformation of the mandible into a single tooth-bearing bone and the emergence of a novel jaw joint-while incorporating some of the ancestral jaw bones into the mammalian middle ear-is often cited as a classic example of the repurposing of morphological structures 3,4 . Although it is remarkably well-documented in the fossil record, the evolution of the mammalian jaw still poses the paradox of how the bones of the ancestral jaw joint could function both as a joint hinge for powerful load-bearing mastication and as a mandibular middle ear that was delicate enough for hearing. Here we use digital reconstructions, computational modelling and biomechanical analyses to demonstrate that the miniaturization of the early mammalian jaw was the primary driver for the transformation of the jaw joint. We show that there is no evidence for a concurrent reduction in jaw-joint stress and increase in bite force in key non-mammaliaform taxa in the cynodont-mammaliaform transition, as previously thought 5-8 . Although a shift in the recruitment of the jaw musculature occurred during the evolution of modern mammals, the optimization of mandibular function to increase bite force while reducing joint loads did not occur until after the emergence of the neomorphic mammalian jaw joint. This suggests that miniaturization provided a selective regime for the evolution of the mammalian jaw joint, followed by the integration of the postdentary bones into the mammalian middle ear.

The evolution of the mammalian jaw during the transition from non-mammalian synapsids to crown mammals is a key event in vertebrate history and characterised by the gradual reduction of its individual bones into a single element and the concomitant transformation of the jaw joint and its incorporation into the middle ear complex. This osteological transformation is accompanied by a rearrangement and modification of the jaw adductor musculature, which is thought to have allowed the evolution of a more-efficient masticatory system in comparison to the plesiomorphic synapsid condition. While osteological characters relating to this transition are well documented in the fossil record, the exact arrangement and modifications of the individual adductor muscles during the cynodont-mammaliaform transition have been debated for nearly a century. We review the existing knowledge about the musculoskeletal evolution of the mammalian jaw adductor complex and evaluate previous hypotheses in the light of recently documented fossils that represent new specimens of existing species, which are of central importance to the mammalian origins debate. By employing computed tomography (CT) and digital reconstruction techniques to create three-dimensional models of the jaw adductor musculature in a number of representative non-mammalian cynodonts and mammaliaforms, we provide an updated perspective on mammalian jaw muscle evolution. As an emerging consensus, current evidence suggests that the mammal-like division of the jaw adductor musculature (into deep and superficial components of the m. masseter, the m. temporalis and the m. pterygoideus) was completed in Eucynodontia. The arrangement of the jaw adductor musculature in a mammalian fashion, with the m. pterygoideus group inserting on the dentary was completed in basal Mammaliaformes as suggested by the muscle reconstruction of Morganucodon oehleri. Consequently, transformation of the jaw adductor musculature from the ancestral ('reptilian') to the mammalian condition must have preceded the emergence of Mammalia and the full formation of the mammalian jaw joint. This suggests that the modification of the jaw adductor system played a pivotal role in the functional morphology and biomechanical stability of the jaw joint., (© 2016 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Temnospondyls were the morphologically and taxonomically most diverse group of early tetrapods with a near-global distribution during the Palaeozoic and Mesozoic. Members of this group occupied a range of different habitats (aquatic, amphibious, terrestrial), reflected by large morphological disparity of the cranium throughout their evolutionary history. A diagnostic feature of temnospondyls is the presence of an open palate with large interpterygoid vacuities, in contrast to the closed palate of most other early tetrapods and their fish-like relatives. Although the function of the interpterygoid vacuities has been discussed in the past, no quantitative studies have been performed to assess their biomechanical significance. Here, we applied finite element analysis, to test the possibility that the interpterygoid vacuities served for stress distribution during contraction of the jaw closing musculature. Different original and theoretical skull models, in which the vacuities differed in size or were completely absent, were compared for their mechanical performance. Our results demonstrate that palatal morphology played a considerable role in cranial biomechanics of temnospondyls. The presence of large cranial vacuities were found to offer the dual benefit of providing additional muscle attachment areas and allowing for more effective force transmission and thus an increase in bite force without compromising cranial stability.

Maniraptoriformes, the speciose group of derived theropod dinosaurs that ultimately gave rise to modern birds, display a diverse and remarkable suite of skeletal adaptations. Apart from the evolution of flight, a large-scale change in dietary behavior appears to have been one of the main triggers for specializations in the bauplan of these derived theropods. Among the different skeletal specializations, partial or even complete edentulism and the development of keratinous beaks form a recurring and persistent trend in from the evolution of derived nonavian dinosaurs. Therizinosauria is an enigmatic maniraptoriform clade, whose members display these and other osteological characters thought to be correlated with the shift from carnivory to herbivory. This makes therizinosaurians prime candidates to assess the functional significance of these morphological characters. Based on a highly detailed biomechanical model of Erlikosaurus andrewsi, a therizinosaurid from the Upper Cretaceous of Mongolia, different morphological configurations incorporating soft-tissue structures, such as a keratinous rhamphotheca, are evaluated for their biomechanical performance. Our results indicate that the development of beaks and the presence of a keratinous rhamphotheca would have helped to dissipate stress and strain, making the rostral part of the skull less susceptible to bending and displacement, and this benefit may extend to other vertebrate clades that possess rhamphothecae. Keratinous beaks, paralleled by edentulism, thus represent an evolutionary innovation developed early in derived theropods to enhance cranial stability, distinct to postulated mass-saving benefits associated with the origin of flight.

Understanding ontogenetic and developmental patterns is critical for reconstructing the life history of fossil vertebrates. In dinosaurs, ontogenetic studies have nearly exclusively focused on changes in the cranial and post-cranial skeleton, whereas ontogenetic changes in the endocranium have received little attention. Here, we present digital reconstructions of the brain and inner ear anatomy of two ontogenetic stages of the Jurassic ornithischian dinosaur Dysalotosaurus lettowvorbecki. Results show that the endocranial anatomy underwent considerable changes during growth, including a rostrocaudal elongation of the olfactory apparatus, a reduction in the cephalic and pontine flexure and an increase in cerebellum size. Functional elements, such as the cerebral hemispheres and the inner ear, were already well developed in early ontogenetic stages, indicating a large degree of precociality. The anisotropic pattern of size and shape changes in the endocranium further indicates that ontogenetic trajectories may be controlled by functional and environmental demands in the different growth stages in Dysalotosaurus lettowvorbecki. The occurrence of similar ontogenetic patterns in the endocranial anatomy of derived ornithopod dinosaurs suggests a more widespread distribution of this growth trajectory. [ABSTRACT FROM AUTHOR]