Morphological evolution and functional consequences of giantism in tyrannosauroid dinosaurs.

Tyrannosauroids are a clade of theropod dinosaur taxa that varied greatly in their body size distribution. We investigated the feeding performance of six tyrannosaur genera of variable body size and skull morphology. We used 3D finite element analysis to test whether skull shape becomes more or less resistant to feeding-induced forces. Cranial and mandibular models were scaled by adult Tyrannosaurus's surface area to analyze the influence of shape on skull function. It was found that Tyrannosaurus experienced higher absolute stresses compared to small-bodied relatives. When surface area values were equalized across genera to account for the effect of size and test efficiency of skull shape, smaller individuals experience notably greater stresses than larger relatives due to the robust cranial osteology characterized in the allometry of tyrannosaurids. These results may indicate that the wide crania of tyrannosaurids convey a functional advantage that basal tyrannosauroids, juvenile tyrannosauroids, and alioramins lacked.

Brazilian fossils reveal homoplasy in the oldest mammalian jaw joint.

The acquisition of the load-bearing dentary-squamosal jaw joint was a key step in mammalian evolution1-5. Although this innovation has received decades of study, questions remain over when and how frequently a mammalian-like skull-jaw contact evolved, hindered by a paucity of three-dimensional data spanning the non-mammaliaform cynodont-mammaliaform transition. New discoveries of derived non-mammaliaform probainognathian cynodonts from South America have much to offer to this discussion. Here, to address this issue, we used micro-computed-tomography scanning to reconstruct the jaw joint anatomy of three key probainognathian cynodonts: Brasilodon quadrangularis, the sister taxon to Mammaliaformes6-8, the tritheledontid-related Riograndia guaibensis9 and the tritylodontid Oligokyphus major. We find homoplastic evolution in the jaw joint in the approach to mammaliaforms, with ictidosaurs (Riograndia plus tritheledontids) independently evolving a dentary-squamosal contact approximately 17 million years before this character first appears in mammaliaforms of the Late Triassic period10-12. Brasilodon, contrary to previous descriptions6-8, lacks an incipient dentary condyle and squamosal glenoid and the jaws articulate solely using a plesiomorphic quadrate-articular joint. We postulate that the jaw joint underwent marked evolutionary changes in probainognathian cynodonts. Some probainognathian clades independently acquired 'double' craniomandibular contacts, with mammaliaforms attaining a fully independent dentary-squamosal articulation with a conspicuous dentary condyle and squamosal glenoid in the Late Triassic. The dentary-squamosal contact, which is traditionally considered to be a typical mammalian feature, therefore evolved more than once and is more evolutionary labile than previously considered.

Endocranial development in non-avian dinosaurs reveals an ontogenetic brain trajectory distinct from extant archosaurs.

Modern birds possess highly encephalized brains that evolved from non-avian dinosaurs. Evolutionary shifts in developmental timing, namely juvenilization of adult phenotypes, have been proposed as a driver of head evolution along the dinosaur-bird transition, including brain morphology. Testing this hypothesis requires a sufficient developmental sampling of brain morphology in non-avian dinosaurs. In this study, we harness brain endocasts of a postnatal growth series of the ornithischian dinosaur Psittacosaurus and several other immature and mature non-avian dinosaurs to investigate how evolutionary changes to brain development are implicated in the origin of the avian brain. Using three-dimensional characterization of neuroanatomical shape across archosaurian reptiles, we demonstrate that (i) the brain of non-avian dinosaurs underwent a distinct developmental trajectory compared to alligators and crown birds; (ii) ornithischian and non-avialan theropod dinosaurs shared a similar developmental trajectory, suggesting that their derived trajectory evolved in their common ancestor; and (iii) the evolutionary shift in developmental trajectories is partly consistent with paedomorphosis underlying overall brain shape evolution along the dinosaur-bird transition; however, the heterochronic signal is not uniform across time and neuroanatomical region suggesting a highly mosaic acquisition of the avian brain form.

Modelling take-off moment arms in an ornithocheiraean pterosaur.

Take-off is a vital part of powered flight which likely constrains the size of birds, yet extinct pterosaurs are known to have reached far larger sizes. Three different hypothesised take-off motions (bipedal burst launching, bipedal countermotion launching, and quadrupedal launching) have been proposed as explanations for how pterosaurs became airborne and circumvented this proposed morphological limit. We have constructed a computational musculoskeletal model of a 5 m wingspan ornithocheiraean pterosaur, reconstructing thirty-four key muscles to estimate the muscle moment arms throughout the three hypothesised take-off motions. Range of motion constrained hypothetical kinematic sequences for bipedal and quadrupedal take-off motions were modelled after extant flying vertebrates. Across our simulations we did not find higher hindlimb moment arms for bipedal take-off motions or noticeably higher forelimb moment arms in the forelimb for quadrupedal take-off motions. Despite this, in all our models we found the muscles utilised in the quadrupedal take-off have the largest total launch applicable moment arms throughout the entire take-off sequences and for the take-off pose. This indicates the potential availability of higher leverage for a quadrupedal take-off than hypothesised bipedal motions in pterosaurs pending further examination of muscle forces.

Widespread convergence towards functional optimization in the lower jaws of crocodile-line archosaurs.

Extant crocodilian jaws are subject to functional demands induced by feeding and hydrodynamics. However, the morphological and ecological diversity of extinct crocodile-line archosaurs is far greater than that of living crocodilians, featuring repeated convergence towards disparate ecologies including armoured herbivores, terrestrial macropredators and fully marine forms. Crocodile-line archosaurs, therefore, present a fascinating case study for morphological and functional divergence and convergence within a clade across a wide range of ecological scenarios. Here, we build performance landscapes of two-dimensional theoretical jaw shapes to investigate the influence of strength, speed and hydrodynamics in the morphological evolution of crocodile-line archosaur jaws, and test whether ecologically convergent lineages evolved similarly optimal jaw function. Most of the 243 sampled jaw morphologies occupy optimized regions of theoretical morphospace for either rotational efficiency, resistance to Von Mises stress, hydrodynamic efficiency or a trade-off between multiple functions, though some seemingly viable shapes remain unrealized. Jaw speed is optimized only in a narrow region of morphospace whereas many shapes possess optimal jaw strength, which may act as a minimum boundary rather than a strong driver for most taxa. This study highlights the usefulness of theoretical morphology in assessing functional optimality, and for investigating form-function relationships in diverse clades.

Anatomy of the maxillary canal of Riograndia guaibensis (Cynodontia, Probainognathia)-A prozostrodont from the Late Triassic of southern Brazil.

Investigating the evolutionary trajectory of synapsid sensory and cephalic systems is pivotal for understanding the emergence and diversification of mammals. Recent studies using CT-scanning to analyze the rostral foramina and maxillary canals morphology in fossilized specimens of probainognathian cynodonts have contributed to clarifying the homology and paleobiological interpretations of these structures. In the present work, µCT-scannings of three specimens of Riograndia guaibensis, an early Norian cynodont from southern Brazil, were analyzed and revealed an incomplete separation between the lacrimal and maxillary canals, with points of contact via non-ossified areas. While the maxillary canal exhibits a consistent morphological pattern with other Prozostrodontia, featuring three main branches along the lateral region of the snout, the rostral alveolar canal in Riograndia displays variability in the number of extra branches terminating in foramina on the lateral surface of the maxilla, showing differences among individuals and within the same skull. Additionally, pneumatization is observed in the anterior region of the skull, resembling similar structures found in reptiles and mammals. Through this pneumatization, certain branches originating from the maxillary canal extend to the canine alveolus. Further investigation is warranted to elucidate the functionality of this structure and its occurrence in other cynodont groups.

New evidence from high-resolution computed microtomography of Triassic stem-mammal skulls from South America enhances discussions on turbinates before the origin of Mammaliaformes.

The nasal cavity of living mammals is a unique structural complex among tetrapods, acquired along a series of major morphological transformations that occurred mainly during the Mesozoic Era, within the Synapsida clade. Particularly, non-mammaliaform cynodonts document several morphological changes in the skull, during the Triassic Period, that represent the first steps of the mammalian bauplan. We here explore the nasal cavity of five cynodont taxa, namely Thrinaxodon, Chiniquodon, Prozostrodon, Riograndia, and Brasilodon, in order to discuss the main changes within this skull region. We did not identify ossified turbinals in the nasal cavity of these taxa and if present, as non-ossified structures, they would not necessarily be associated with temperature control or the development of endothermy. We do, however, notice a complexification of the cartilage anchoring structures that divide the nasal cavity and separate it from the brain region in these forerunners of mammals.

Finite element analysis of kangaroo astragali: A new angle on the ankle.

Using finite element analysis on the astragali of five macropodine kangaroos (extant and extinct hoppers) and three sthenurine kangaroos (extinct proposed bipedal striders) we investigate how the stresses experienced by the ankle in similarly sized kangaroos of different hypothesized/known locomotor strategy compare under different simulation scenarios, intended to represent the moment of midstance at different gaits. These tests showed a clear difference between the performance of sthenurines and macropodines with the former group experiencing lower stress in simulated bipedal strides in all species compared with hopping simulations, supporting the hypothesis that sthenurines may have utilized this gait. The Pleistocene macropodine Protemnodon also performed differently from all other species studied, showing high stresses in all simulations except for bounding. This may support the hypothesis of Protemnodon being a quadrupedal bounder.

Predatory synapsid ecomorphology signals growing dynamism of late Palaeozoic terrestrial ecosystems.

Terrestrial ecosystems evolved substantially through the Palaeozoic, especially the Permian, gaining much new complexity, especially among predators. Key among these predators were non-mammalian synapsids. Predator ecomorphology reflect interactions with prey and competitors, which are key controls on carnivore diversity and ecology. Therefore, carnivorous synapsids may offer insight on wider ecological evolution as the first complex, tetrapod-dominated, terrestrial ecosystems formed through the late Palaeozoic. Using morphometric and phylogenetic comparative methods, we chart carnivorous synapsid trophic morphology from the latest Carboniferous to the earliest Triassic (307-251.2 Ma). We find a major morphofunctional shift in synapsid carnivory between the early and middle Permian, via the addition of new feeding modes increasingly specialised for greater biting power or speed that captures the growing antagonism and dynamism of terrestrial tetrapod predator-prey interactions. The further evolution of new hypo- and hypercarnivorous synapsids highlight the nascent intrinsic pressures and complexification of terrestrial ecosystems across the mid-late Permian.

Environmental impacts on the structural integrity of British rhodoliths.

Coralline algae form complex habitats which are biodiversity hotspots. Experimental studies suggest that climate change will decrease coralline algal structural integrity. These experiments, however, lack information on local morphological variability and how much structural change would be needed to threaten habitat formation. Here, using finite element modelling, we assess variability in cellular structure and chemical composition of the carbonate skeleton of four coralline algal species from Britain in contemporary and historical specimens collected over the last 130 years. Cellular and mineral properties are highly variable within species, between sites and through time, with structurally weaker cells in the southern species and contemporary material compared to northern taxa and historical material. Yet, temporal differences in strength were smaller than spatial differences. Our work supports long term experiments which show the adaptation potential of this group. Our results suggest that future anthropogenic climate change may lead to loss of habitat complexity in the south and expansion of structurally weaker southern species into northern sites.

Extracellular matrix assembly stress initiates Drosophila central nervous system morphogenesis.

Forces controlling tissue morphogenesis are attributed to cellular-driven activities, and any role for extracellular matrix (ECM) is assumed to be passive. However, all polymer networks, including ECM, can develop autonomous stresses during their assembly. Here, we examine the morphogenetic function of an ECM before reaching homeostatic equilibrium by analyzing de novo ECM assembly during Drosophila ventral nerve cord (VNC) condensation. Asymmetric VNC shortening and a rapid decrease in surface area correlate with the exponential assembly of collagen IV (Col4) surrounding the tissue. Concomitantly, a transient developmentally induced Col4 gradient leads to coherent long-range flow of ECM, which equilibrates the Col4 network. Finite element analysis and perturbation of Col4 network formation through the generation of dominant Col4 mutations that affect assembly reveal that VNC morphodynamics is partially driven by a sudden increase in ECM-driven surface tension. These data suggest that ECM assembly stress and associated network instabilities can actively participate in tissue morphogenesis.

Functional reorganisation of the cranial skeleton during the cynodont-mammaliaform transition.

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.

Functional space analyses reveal the function and evolution of the most bizarre theropod manual unguals.

Maniraptoran dinosaurs include the ancestors of birds, and most used their hands for grasping and in flight, but early-branching maniraptorans had extraordinary claws of mysterious function. Alvarezsauroids had short, strong arms and hands with a stout, rock-pick-like, single functional finger. Therizinosaurians had elongate fingers with slender and sickle-like unguals, sometimes over one metre long. Here we develop a comprehensive methodological framework to investigate what the functions of these most bizarre bony claws are and how they formed. Our analysis includes finite element analysis and a newly established functional-space analysis and also involves shape and size effects in an assessment of function and evolution. We find a distinct functional divergence among manual unguals of early-branching maniraptorans, and we identify a complex relationship between their structural strength, morphological specialisations, and size changes. Our analysis reveals that efficient digging capabilities only emerged in late-branching alvarezsauroid forelimbs, rejecting the hypothesis of functional vestigial structures like T. rex. Our results also support the statement that most therizinosaurians were herbivores. However, the bizarre, huge Therizinosaurus had sickle-like unguals of such length that no mechanical function has been identified; we suggest they were decorative and lengthened by peramorphic growth linked to increased body size.

Dental form and function in the early feeding diversification of dinosaurs.

Dinosaurs evolved a remarkable diversity of dietary adaptations throughout the Mesozoic, but the origins of different feeding modes are uncertain, especially the multiple origins of herbivory. Feeding habits of early dinosaurs have mostly been inferred from qualitative comparisons of dental morphology with extant analogs. Here, we use biomechanical and morphometric methods to investigate the dental morphofunctional diversity of early dinosaurs in comparison with extant squamates and crocodylians and predict their diets using machine learning classification models. Early saurischians/theropods are consistently classified as carnivores. Sauropodomorphs underwent a dietary shift from faunivory to herbivory, experimenting with diverse diets during the Triassic and Early Jurassic, and early ornithischians were likely omnivores. Obligate herbivory was a late evolutionary innovation in both clades. Carnivory is the most plausible ancestral diet of dinosaurs, but omnivory is equally likely under certain phylogenetic scenarios. This early dietary diversity was fundamental in the rise of dinosaurs to ecological dominance.

Early tetrapod cranial evolution is characterized by increased complexity, constraint, and an offset from fin-limb evolution.

The developmental underpinnings and functional consequences of modifications to the limbs during the origin of the tetrapod body plan are increasingly well characterized, but less is understood about the evolution of the tetrapod skull. Decrease in skull bone number has been hypothesized to promote morphological and functional diversification in vertebrate clades, but its impact during the initial rise of tetrapods is unknown. Here, we test this by quantifying topological changes to cranial anatomy in fossil and living taxa bracketing the fin-to-limb transition using anatomical network analysis. We find that bone loss across the origin of tetrapods is associated not only with increased complexity of bone-to-bone contacts but also with decreasing topological diversity throughout the late Paleozoic, which may be related to developmental and/or mechanical constraints. We also uncover a 10-Ma offset between fin-limb and cranial morphological evolution, suggesting that different evolutionary drivers affected these features during the origin of tetrapods.

The efficacy of computed tomography scanning versus surface scanning in 3D finite element analysis.

Finite element analysis (FEA) is a commonly used application in biomechanical studies of both extant and fossil taxa to assess stress and strain in solid structures such as bone. FEA can be performed on 3D structures that are generated using various methods, including computed tomography (CT) scans and surface scans. While previous palaeobiological studies have used both CT scanned models and surface scanned models, little research has evaluated to what degree FE results may vary when CT scans and surface scans of the same object are compared. Surface scans do not preserve the internal geometries of 3D structures, which are typically preserved in CT scans. Here, we created 3D models from CT scans and surface scans of the same specimens (crania and mandibles of a Nile crocodile, a green sea turtle, and a monitor lizard) and performed FEA under identical loading parameters. It was found that once surface scanned models are solidified, they output stress and strain distributions and model deformations comparable to their CT scanned counterparts, though differing by notable stress and strain magnitudes in some cases, depending on morphology of the specimen and the degree of reconstruction applied. Despite similarities in overall mechanical behaviour, surface scanned models can differ in exterior shape compared to CT scanned models due to inaccuracies that can occur during scanning and reconstruction, resulting in local differences in stress distribution. Solid-fill surface scanned models generally output lower stresses compared to CT scanned models due to their compact interiors, which must be accounted for in studies that use both types of scans.

Increasing morphological disparity and decreasing optimality for jaw speed and strength during the radiation of jawed vertebrates.

The Siluro-Devonian adaptive radiation of jawed vertebrates, which underpins almost all living vertebrate biodiversity, is characterized by the evolutionary innovation of the lower jaw. Multiple lines of evidence have suggested that the jaw evolved from a rostral gill arch, but when the jaw took on a feeding function remains unclear. We quantified the variety of form in the earliest jaws in the fossil record from which we generated a theoretical morphospace that we then tested for functional optimality. By drawing comparisons with the real jaw data and reconstructed jaw morphologies from phylogenetically inferred ancestors, our results show that the earliest jaw shapes were optimized for fast closure and stress resistance, inferring a predatory feeding function. Jaw shapes became less optimal for these functions during the later radiation of jawed vertebrates. Thus, the evolution of jaw morphology has continually explored previously unoccupied morphospace and accumulated disparity through time, laying the foundation for diverse feeding strategies and the success of jawed vertebrates.

Walking with early dinosaurs: appendicular myology of the Late Triassic sauropodomorph Thecodontosaurus antiquus.

Dinosaur evolution is marked by numerous independent shifts from bipedality to quadrupedality. Sauropodomorpha is one of the lineages that transitioned from small bipedal forms to graviportal quadrupeds, with an array of intermediate postural strategies evolving in non-sauropodan sauropodomorphs. This locomotor shift is reflected by multiple modifications of the appendicular skeleton, coupled with a drastic rearrangement of the limb musculature. Here, we describe the osteological correlates of appendicular muscle attachment of the Late Triassic sauropodomorph Thecodontosaurus antiquus from multiple well-preserved specimens and provide the first complete forelimb and hindlimb musculature reconstruction of an early-branching sauropodomorph. Comparisons with other sauropodomorphs and early dinosaurs reveal a unique combination of both plesiomorphic and derived musculoskeletal features. The diversity of appendicular osteological correlates among early dinosaurs and their relevance in muscle reconstruction are discussed. In line with previous evidence, aspects of the limb muscle arrangement, such as conspicuous correlates of lower limb extensors and flexors and low moment arms of hip extensors and flexors, suggest Thecodontosaurus was an agile biped. This reconstruction helps to elucidate the timing of important modifications of the appendicular musculature in the evolution of sauropodomorphs which facilitated the transition to quadrupedalism and contributed to their evolutionary success.

Divergent locomotor evolution in "giant" kangaroos: Evidence from foot bone bending resistances and microanatomy.

The extinct sthenurine (giant, short-faced) kangaroos have been proposed to have a different type of locomotor behavior to extant (macropodine) kangaroos, based both on physical limitations (the size of many exceeds the proposed limit for hopping) and anatomical features (features of the hind limb anatomy suggestive of weight-bearing on one leg at a time). Here, we use micro computerised tomography (micro-CT) scans of the pedal bones of six kangaroos, three sthenurine, and three macropodine, ranging from ~50 to 150 kg, to investigate possible differences in bone resistances to bending and cortical bone distribution that might relate to differences in locomotion. Using second moment of area analysis, we show differences in resistance to bending between the two subfamilies. Distribution of cortical bone shows that sthenurines had less resistant calcaneal tubers, implying a different foot posture during locomotion, and the long foot bones were more resistant to the medial bending stresses. These differences were the most pronounced between Pleistocene monodactyl sthenurines (Sthenurus stirlingi and Procoptodon browneorum) and the two species of Macropus (the extant M. giganteus and the extinct M. cf. M. titan) and support the hypothesis that these derived sthenurines employed bipedal striding. The Miocene sthenurine Hadronomas retains some more macropodine-like features of bone resistance to bending, perhaps reflecting its retention of the fifth pedal digit. The Pleistocene macropodine Protemnodon has a number of unique features, possibly indicative of a type of locomotion unlike the other kangaroos.

Cranial functional morphology of the pseudosuchian Effigia and implications for its ecological role in the Triassic.

Pseudosuchians, archosaurian reptiles more closely related to crocodylians than to birds, exhibited high morphological diversity during the Triassic with numerous examples of morphological convergence described between Triassic pseudosuchians and post-Triassic dinosaurs. One example is the shuvosaurid Effigia okeeffeae which exhibits an "ostrich-like" bauplan comprising a gracile skeleton with edentulous jaws and large orbits, similar to ornithomimid dinosaurs and extant palaeognaths. This bauplan is regarded as an adaptation for herbivory, but this hypothesis assumes morphological convergence confers functional convergence, and has received little explicit testing. Here, we restore the skull morphology of Effigia, perform myological reconstructions, and apply finite element analysis to quantitatively investigate skull function. We also perform finite element analysis on the crania of the ornithomimid dinosaur Ornithomimus edmontonicus, the extant palaeognath Struthio camelus and the extant pseudosuchian Alligator mississippiensis to assess the degree of functional convergence with a taxon that exhibit "ostrich-like" bauplans and its closest extant relatives. We find that Effigia possesses a mosaic of mechanically strong and weak features, including a weak mandible that likely restricted feeding to the anterior portion of the jaws. We find limited functional convergence with Ornithomimus and Struthio and limited evidence of phylogenetic constraints with extant pseudosuchians. We infer that Effigia was a specialist herbivore that likely fed on softer plant material, a niche unique among the study taxa and potentially among contemporaneous Triassic herbivores. This study increases the known functional diversity of pseudosuchians and highlights that superficial morphological similarity between unrelated taxa does not always imply functional and ecological convergence.