Andrew R. Cuff

Feeding Mechanics in Spinosaurid Theropods and Extant Crocodilians

A number of extant and extinct archosaurs evolved an elongate, narrow rostrum. This longirostrine condition has been associated with a diet comprising a higher proportion of fish and smaller prey items compared to taxa with broader, more robust snouts. The evolution of longirostrine morphology and a bulbous anterior rosette of premaxillary teeth also occurs in the spinosaurid theropod dinosaurs, leading to suggestions that at least some members of this clade also had a diet comprising a notable proportion of fish or other small vertebrates. Here we compare the rostral biomechanics of the spinosaurs Baryonyx walkeri and Spinosaurus c.f. S. aegyptiacus to three extant crocodilians: two longistrine taxa, the African slender-snouted crocodile Mecistops cataphractus and the Indian gharial Gavialis gangeticus; and the American alligator Alligator mississippiensis. Using computed tomography (CT) data, the second moments of area and moments of inertia at successive transverse slices along the rostrum were calculated for each of the species. Size-independent results tested the biomechanical benefits of material distribution within the rostra. The two spinosaur rostra were both digitally reconstructed from CT data and compared against all three crocodilians. Results show that African slender-snouted crocodile skulls are more resistant to bending than an equivalent sized gharial. The alligator has the highest resistances to bending and torsion of the crocodiles for its size and greater than that of the spinosaurs. The spinosaur rostra possess similar resistance to bending and torsion despite their different morphologies. When size is accounted for, B. walkeri performs mechanically differently from the gharial, contradicting previous studies whereas Spinosaurus does not. Biomechanical data support known feeding ecology for both African slender-snouted crocodile and alligator, and suggest that the spinosaurs were not obligate piscivores with diet being determined by individual animal size.

Validation experiments on finite element models of an ostrich (Struthio camelus) cranium

The first finite element (FE) validation of a complete avian cranium was performed on an extant palaeognath, the ostrich (Struthio camelus). Ex-vivo strains were collected from the cranial bone and rhamphotheca. These experimental strains were then compared to convergence tested, specimen-specific finite element (FE) models. The FE models contained segmented cortical and trabecular bone, sutures and the keratinous rhamphotheca as identified from micro-CT scan data. Each of these individual materials was assigned isotropic material properties either from the literature or from nanoindentation, and the FE models compared to the ex-vivo results. The FE models generally replicate the location of peak strains and reflect the correct mode of deformation in the rostral region. The models are too stiff in regions of experimentally recorded high strain and too elastic in regions of low experimentally recorded low strain. The mode of deformation in the low strain neurocranial region is not replicated by the FE models, and although the models replicate strain orientations to within 10° in some regions, in most regions the correlation is not strong. Cranial sutures, as has previously been found in other taxa, are important for modifying both strain magnitude and strain patterns across the entire skull, but especially between opposing the sutural junctions. Experimentally, we find that the strains on the surface of the rhamphotheca are much lower than those found on nearby bone. The FE models produce much higher principal strains despite similar strain ratios across the entirety of the rhamphotheca. This study emphasises the importance of attempting to validate FE models, modelling sutures and rhamphothecae in birds, and shows that whilst location of peak strain and patterns of deformation can be modelled, replicating experimental data in digital models of avian crania remains problematic.

Retrodeformation and muscular reconstruction of ornithomimosaurian dinosaur crania

Ornithomimosaur dinosaurs evolved lightweight, edentulous skulls that possessed keratinous rhamphothecae. Understanding the anatomy of these taxa allows for a greater understanding of “ostrich-mimic” dinosaurs and character change during theropod dinosaur evolution. However, taphonomic processes during fossilisation often distort fossil remains. Retrodeformation offers a means by which to recover a hypothesis of the original anatomy of the specimen, and 3D scanning technologies present a way to constrain and document the retrodeformation process. Using computed tomography (CT) scan data, specimen specific retrodeformations were performed on three-dimensionally preserved but taphonomically distorted skulls of the deinocheirid Garudimimus brevipes Barsbold, 1981 and the ornithomimids Struthiomimus altus Lambe, 1902 and Ornithomimus edmontonicus Sternberg, 1933. This allowed for a reconstruction of the adductor musculature, which was then mapped onto the crania, from which muscle mechanical advantage and bite forces were calculated pre- and post-retrodeformation. The extent of the rhamphotheca was varied in each taxon to represent morphologies found within modern Aves. Well constrained retrodeformation allows for increased confidence in anatomical and functional analysis of fossil specimens and offers an opportunity to more fully understand the soft tissue anatomy of extinct taxa.

The scaling of postcranial muscles in cats (Felidae) I: forelimb, cervical, and thoracic muscles

The body masses of cats (Mammalia, Carnivora, Felidae) span a ~300‐fold range from the smallest to largest species. Despite this range, felid musculoskeletal anatomy remains remarkably conservative, including the maintenance of a crouched limb posture at unusually large sizes. The forelimbs in felids are important for body support and other aspects of locomotion, as well as climbing and prey capture, with the assistance of the vertebral (and hindlimb) muscles. Here, we examine the scaling of the anterior postcranial musculature across felids to assess scaling patterns between different species spanning the range of felid body sizes. The muscle architecture (lengths and masses of the muscle‐tendon unit components) for the forelimb, cervical and thoracic muscles was quantified to analyse how the muscles scale with body mass. Our results demonstrate that physiological cross‐sectional areas of the forelimb muscles scale positively with increasing body mass (i.e. becoming relatively larger). Many significantly allometric variables pertain to shoulder support, whereas the rest of the limb muscles become relatively weaker in larger felid species. However, when phylogenetic relationships were corrected for, most of these significant relationships disappeared, leaving no significantly allometric muscle metrics. The majority of cervical and thoracic muscle metrics are not significantly allometric, despite there being many allometric skeletal elements in these regions. When forelimb muscle data were considered in isolation or in combination with those of the vertebral muscles in principal components analyses and MANOVAs, there was no significant discrimination among species by either size or locomotory mode. Our results support the inference that larger felid species have relatively weaker anterior postcranial musculature compared with smaller species, due to an absence of significant positive allometry of forelimb or vertebral muscle architecture. This difference in strength is consistent with behavioural changes in larger felids, such as a reduction of maximal speed and other aspects of locomotor abilities.

Big cat, small cat: reconstructing body size evolution in living and extinct Felidae

The evolution of body mass is a fundamental topic in evolutionary biology, because it is closely linked to manifold life history and ecological traits and is readily estimable for many extinct taxa. In this study, we examine patterns of body mass evolution in Felidae (Placentalia, Carnivora) to assess the effects of phylogeny, mode of evolution, and the relationship between body mass and prey choice in this charismatic mammalian clade. Our data set includes 39 extant and 26 extinct taxa, with published body mass data supplemented by estimates based on condylobasal length. These data were run through ‘SURFACE’ and ‘bayou’ to test for patterns of body mass evolution and convergence between taxa. Body masses of felids are significantly different among prey choice groupings (small, mixed and large). We find that body mass evolution in cats is strongly influenced by phylogeny, but different patterns emerged depending on inclusion of extinct taxa and assumptions about branch lengths. A single Ornstein–Uhlenbeck optimum best explains the distribution of body masses when first‐occurrence data were used for the fossil taxa. However, when mean occurrence dates or last known occurrence dates were used, two selective optima for felid body mass were recovered in most analyses: a small optimum around 5 kg and a large one around 100 kg. Across living and extinct cats, we infer repeated evolutionary convergences towards both of these optima, but, likely due to biased extinction of large taxa, our results shift to supporting a Brownian motion model when only extant taxa are included in analyses.

Functional anatomy and feeding biomechanics of a giant Upper Jurassic pliosaur (Reptilia: Sauropterygia) from Weymouth Bay, Dorset, UK

Pliosaurs were among the largest predators in Mesozoic seas, and yet their functional anatomy and feeding biomechanics are poorly understood. A new, well‐preserved pliosaur from the Kimmeridgian of Weymouth Bay (UK) revealed cranial adaptations related to feeding. Digital modelling of computed tomography scans allowed reconstruction of missing, distorted regions of the skull and of the adductor musculature, which indicated high bite forces. Size‐corrected beam theory modelling showed that the snout was poorly optimised against bending and torsional stresses compared with other aquatic and terrestrial predators, suggesting that pliosaurs did not twist or shake their prey during feeding and that seizing was better performed with post‐symphyseal bites. Finite element analysis identified biting‐induced stress patterns in both the rostrum and lower jaws, highlighting weak areas in the rostral maxillary‐premaxillary contact and the caudal mandibular symphysis. A comparatively weak skull coupled with musculature that was able to produce high forces, is explained as a trade‐off between agility, hydrodynamics and strength. In the Kimmeridgian ecosystem, we conclude that Late Jurassic pliosaurs were generalist predators at the top of the food chain, able to prey on reptiles and fishes up to half their own length.

Regional differentiation of felid vertebral column evolution: a study of 3D shape trajectories

Recent advances in geometric morphometrics provide improved techniques for extraction of biological information from shape and have greatly contributed to the study of ecomorphology and morphological evolution. However, the vertebral column remains an under-studied structure due in part to a concentration on skull and limb research, but most importantly because of the difficulties in analysing the shape of a structure composed of multiple articulating discrete units (i.e. vertebrae). Here, we have applied a variety of geometric morphometric analyses to three-dimensional landmarks collected on 19 presacral vertebrae to investigate the influence of potential ecological and functional drivers, such as size, locomotion and prey size specialisation, on regional morphology of the vertebral column in the mammalian family Felidae. In particular, we have here provided a novel application of a method—phenotypic trajectory analysis (PTA)—that allows for shape analysis of a contiguous sequence of vertebrae as functionally linked osteological structures. Our results showed that ecological factors influence the shape of the vertebral column heterogeneously and that distinct vertebral sections may be under different selection pressures. While anterior presacral vertebrae may either have evolved under stronger phylogenetic constraints or are ecologically conservative, posterior presacral vertebrae, specifically in the post-T10 region, show significant differentiation among ecomorphs. Additionally, our PTA results demonstrated that functional vertebral regions differ among felid ecomorphs mainly in the relative covariation of vertebral shape variables (i.e. direction of trajectories, rather than in trajectory size) and, therefore, that ecological divergence among felid species is reflected by morphological changes in vertebral column shape.

Endocranial Morphology of the Extinct North American Lion (Panthera atrox)

The extinct North American lion (Panthera atrox) is one of the largest felids (Mammalia, Carnivora) to have ever lived, and it is known from a plethora of incredibly well-preserved remains. Despite this abundance of material, there has been little research into its endocranial anatomy. CT scans of a skull of P. atrox from the Pleistocene La Brea Tar pits were used to generate the first virtual endocranium for this species and to elucidate previously unknown details of its brain size and gross structure, cranial nerves, and inner-ear morphology. Results show that its gross brain anatomy is broadly similar to that of other pantherines, although P. atrox displays less cephalic flexure than either extant lions or tigers, instead showing a brain shape that is reminiscent of earlier felids. Despite this unusual reduction in flexure, the estimated absolute brain size for this specimen is one of the largest reported for any felid, living or extinct. Its encephalization quotient (brain size as a fraction of the expected brain mass for a given body mass) is also larger than that of extant lions but similar to that of the other pantherines. The advent of CT scans has allowed nondestructive sampling of anatomy that cannot otherwise be studied in these extinct lions, leading to a more accurate reconstruction of endocranial morphology and its evolution.