Extant and fossil crocodilians have long been divided into taxonomic and/or ecological groups based on broad patterns of skull shape, particularly the relative length and width of the snout. However, these patterns have not been quantitatively analyzed in detail, and their biomechanical and functional implications are similarly understudied. Here, we use geometric morphometrics and finite element analysis to explore the patterns of variation in crocodilian skull morphology and the functional implications of those patterns. Our results indicate that skull shape variation in extant crocodiles is much more complex than previously recognized. Differences in snout length and width are the main components of shape variation, but these differences are correlated with changes in other regions of the skull. Additionally, there is considerable disparity within general classes such as longirostrine and brevirostrine forms. For example, Gavialis and Tomistoma occupy different parts of morphospace implying a significant difference in skull shape, despite the fact that both are traditionally considered longirostrine. Skull length and width also strongly influence the mechanical performance of the skull; long and narrow morphotypes (e.g., Tomistoma) experience the highest amount of stress during biting, whereas short and broad morphotypes (e.g., Caiman latirostris) experience the least amount of stress. Biomechanical stress and the hydrodynamic properties of the skull show a strong relationship with the distribution of crocodilians in skull morphospace, whereas phylogeny and biogeography show weak or no correlation. Therefore, ecological specializations related to feeding and foraging likely have the greatest influence on crocodilian skull shape.

The origin of tetrapods and the transition from swimming to walking was a pivotal step in the evolution and diversification of terrestrial vertebrates. During this time, modifications of the limbs—particularly the specialization of joints and the structures that guide their motions—fundamentally changed the ways in which early tetrapods could move. Nonetheless, little is known about the functional consequences of limb anatomy in early tetrapods and how that anatomy influenced locomotion capabilities at this very critical stage in vertebrate evolution. Here we present a three-dimensional reconstruction of the iconic Devonian tetrapod Ichthyostega and a quantitative and comparative analysis of limb mobility in this early tetrapod. We show that Ichthyostega could not have employed typical tetrapod locomotory behaviours, such as lateral sequence walking. In particular, it lacked the necessary rotary motions in its limbs to push the body off the ground and move the limbs in an alternating sequence. Given that long-axis rotation was present in the fins of tetrapodomorph fishes, it seems that either early tetrapods evolved through an initial stage of restricted shoulder and hip joint mobility or that Ichthyostega was unique in this respect. We conclude that early tetrapods with the skeletal morphology and limb mobility of Ichthyostega were unlikely to have made some of the recently described Middle Devonian trackways.

Thalattosuchians were highly specialised aquatic archosaurs of the Jurassic and Early Cretaceous, and represent a peak of aquatic adaptation among crocodylomorphs. Relatively little is known of their endocranial anatomy or its relevance for the evolution of sensory systems, physiology, and other aspects of biology. Nevertheless, such data have significance for two reasons: (1) thalattosuchians represent an important data point regarding adaptation to marine life in tetrapods; and (2) as early-diverging members of the crocodylian stem-lineage, thalattosuchians provide information on the evolutionary assembly of the brain and other endocranial structures in crocodylomorphs. Here we use µCT data to virtually reconstruct the endocranial anatomy of Pelagosaurus typus, an early thalattosuchian with plesiomorphic traits of relevance to the split between the two major subgroups: Teleosauroidea and Metriorhynchoidea. Interpretation of these data in a broad comparative context indicate that several key endocranial features may be unique to thalattosuchians, including: a pyramidal morphology of the semicircular canals, the presence of an elongate endosseous cochlear duct that may indicate enhanced hearing ability, the presence of large, paired canals extending anteriorly from an enlarged pituitary fossa, a relatively straight brain (possibly due to the presence of large, laterally placed orbits), and an enlarged venous sinus projecting dorsally from the endocast that is confluent with the paratympanic sinus system. Notably, we document a large expansion of the nasal cavity anterior to the orbits in Pelagosaurus as an osteological correlate of an enlarged salt gland previously only documented in Late Jurassic metriorhynchoids. This is the first anatomical evidence of this structure in early thalattosuchians. Pelagosaurus also shares the presence of paired olfactory bulbs with metriorhynchoids, and shows an enlarged cerebrum, which may also be present in teleosauroids. Taken together, our findings indicate that physiological and sensory adaptations to marine life occurred early in thalattosuchian evolution, predating the origins of flippers, tail flukes, and hydrodynamic body forms seen later in metriorhynchoids.

ABSTRACT-Geometric morphometrics (GM) and finite element analysis (FEA) are increasingly common techniques for the study of form and function. We show how principles of quantitative evolution in continuous phenotypic traits can link the two techniques, allowing hypotheses about the relative importance of different functions to be tested in a phylogenetic and evolutionary framework. Finite element analysis is used to derive quantitative surfaces that describe the comparative performance of different morphologies in a morphospace derived from GM. The combination of two or more performance surfaces describes a quantitative adaptive landscape that can be used to predict the direction morphological evolution would take if a combination of functions was selected for. Predicted paths of evolution also can be derived for hypotheses about the relative importance of multiple functions, which can be tested against evolutionary pathways that are documented by phylogenies or fossil sequences. Magnitudes of evolutionary trade-offs between functions can be estimated using maximum likelihood. We apply these methods to an earlier study of carapace strength and hydrodynamic efficiency in emydid turtles. We find that strength and hydrodynamic efficiency explain about 45% of the variance in shell shape; drift and other unidentified functional factors are necessary to explain the remaining variance. Measurement of the proportional trade-off between shell strength and hydrodynamic efficiency shows that throughout the Cenozoic aquatic turtles generally sacrificed strength for streamlining and terrestrial species favored stronger shells; this suggests that the selective regime operating on small to mid-sized emydids has remained relatively static.SUPPLEMENTAL DATA-Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Polly, P. D., C. T. Stayton, E. R. Dumont, S. E. Pierce, E. J. Rayfield, and K. D. Angielczyk. 2016. Combining geometric morphometrics and finite element analysis with evolutionary modeling: towards a synthesis. Journal of Vertebrate Paleontology.