Digital dissection and three-dimensional interactive models of limb musculature in the Australian estuarine crocodile (Crocodylus porosus).

Digital dissection is a relatively new technique that has enabled scientists to gain a better understanding of vertebrate anatomy. It can be used to rapidly disseminate detailed, three-dimensional information in an easily accessible manner that reduces the need for destructive, traditional dissections. Here we present the results of a digital dissection on the appendicular musculature of the Australian estuarine crocodile (Crocodylus porosus). A better understanding of this until now poorly known system in C. porosus is important, not only because it will expand research into crocodilian locomotion, but because of its potential to inform muscle reconstructions in dinosaur taxa. Muscles of the forelimb and hindlimb are described and three-dimensional interactive models are included based on CT and MRI scans as well as fresh-tissue dissections. Differences in the arrangement of musculature between C. porosus and other groups within the Crocodylia were found. In the forelimb, differences are restricted to a single tendon of origin for triceps longus medialis. For the hindlimb, a reduction in the number of heads of ambiens was noted as well as changes to the location of origin and insertion for iliofibularis and gastrocnemius externus.

What lies beneath: sub-articular long bone shape scaling in eutherian mammals and saurischian dinosaurs suggests different locomotor adaptations for gigantism.

Eutherian mammals and saurischian dinosaurs both evolved lineages of huge terrestrial herbivores. Although significantly more saurischian dinosaurs were giants than eutherians, the long bones of both taxa scale similarly and suggest that locomotion was dynamically similar. However, articular cartilage is thin in eutherian mammals but thick in saurischian dinosaurs, differences that could have contributed to, or limited, how frequently gigantism evolved. Therefore, we tested the hypothesis that sub-articular bone, which supports the articular cartilage, changes shape in different ways between terrestrial mammals and dinosaurs with increasing size. Our sample consisted of giant mammal and reptile taxa (i.e., elephants, rhinos, sauropods) plus erect and non-erect outgroups with thin and thick articular cartilage. Our results show that eutherian mammal sub-articular shape becomes narrow with well-defined surface features as size increases. In contrast, this region in saurischian dinosaurs expands and remains gently convex with increasing size. Similar trends were observed in non-erect outgroup taxa (monotremes, alligators), showing that the trends we report are posture-independent. These differences support our hypothesis that sub-articular shape scales differently between eutherian mammals and saurischian dinosaurs. Our results show that articular cartilage thickness and sub-articular shape are correlated. In mammals, joints become ever more congruent and thinner with increasing size, whereas archosaur joints remained both congruent and thick, especially in sauropods. We suggest that gigantism occurs less frequently in mammals, in part, because joints composed of thin articular cartilage can only become so congruent before stress cannot be effectively alleviated. In contrast, frequent gigantism in saurischian dinosaurs may be explained, in part, by joints with thick articular cartilage that can deform across large areas with increasing load.

Calcified cartilage shape in archosaur long bones reflects overlying joint shape in stress-bearing elements: Implications for nonavian dinosaur locomotion.

In nonavian dinosaur long bones, the once-living chondroepiphysis (joint surface) overlay a now-fossilized calcified cartilage zone. Although the shape of this zone is used to infer nonavian dinosaur locomotion, it remains unclear how much it reflects chondroepiphysis shape. We tested the hypothesis that calcified cartilage shape reflects the overlying chondroepiphysis in extant archosaurs. Long bones with intact epiphyses from American alligators (Alligator mississippiensis), helmeted guinea fowl (Numida meleagris), and juvenile ostriches (Struthio camelus) were measured and digitized for geometric morphometric (GM) analyses before and after chondroepiphysis removal. Removal of the chondroepiphysis resulted in significant element truncation in all examined taxa, but the amount of truncation decreased with increasing size. GM analyses revealed that Alligator show significant differences between chondroepiphysis shape and the calcified cartilage zone in the humerus, but display nonsignificant differences in femora of large individuals. In Numida, GM analysis shows significant shape differences in juvenile humeri, but humeri of adults and the femora of all guinea fowl show no significant shape difference. The juvenile Struthio sample showed significant differences in both long bones, which diminish with increasing size, a pattern confirmed with magnetic resonance imaging scans in an adult. Our data suggest that differences in extant archosaur long bone shape are greater in elements not utilized in locomotion and related stress-inducing activities. Based on our data, we propose tentative ranges of error for nonavian dinosaur long bone dimensional measurements. We also predict that calcified cartilage shape in adult, stress-bearing nonavian dinosaur long bones grossly reflects chondroepiphysis shape.

Differential limb scaling in the american alligator (Alligator mississippiensis) and its implications for archosaur locomotor evolution.

Bipedalism evolved multiple times within archosaurs, and relatively shorter forelimbs characterize both crocodyliforms and nonavian dinosaurs. Analysis of a comprehensive ontogenetic sequence of specimens (embryo to adult) of the sauropodomorph Massospondylus has shown that bipedal limb proportions result from negative forelimb allometry. We ask, is negative forelimb allometry a pattern basal to archosaurs, amplified in certain taxa to produce bipedalism? Given the phylogenetic position of extant crocodylians and their relatively shorter forelimb, we tested the hypothesis that prevalent negative forelimb allometry is present in Alligator mississippiensis from a sample of wild specimens from embryonic to adult sizes. Long bone lengths (humerus, radius, ulna, femur, tibia, fibula, third metapodials) were measured with their epiphyseal cartilage intact at all sizes. Our results show an overall isometric pattern for most elements regressed on femur length, humerus length, or total limb length. However, negative allometry was prevalent for the ulna, and the third metapodials scale with positive allometry embryonically. These data suggest that the general forelimb proportions in relation to the hindlimb do not change significantly with increasing size in A. mississippiensis. The negative allometry of the ulna and embryonicaly positive allometry of the third metapodials appears to be related to maintaining the functional integrity of the limbs. We show that this pattern is different from that of the sauropodomorph Massospondylus, and we suggest that if bipedalism in archosaurs is tied, in part, to negative forearm allometry, it was either secondarily lost through isometric scaling, or never developed in the ancestor of A. mississippiensis.