Contrast-enhanced XROMM reveals in vivo soft tissue interactions in the hip of Alligator mississippiensis.

Extant archosaurs exhibit highly divergent articular soft tissue anatomies between avian and crocodilian lineages. However, the general lack of understanding of the dynamic interactions among archosaur joint soft tissues has hampered further inferences about the function and evolution of these joints. Here we use contrast-enhanced computed tomography to generate 3D surface models of the pelvis, femora, and hip joint soft tissues in an extant archosaur, the American alligator. The hip joints were then animated using marker-based X-Ray Reconstruction of Moving Morphology (XROMM) to visualize soft tissue articulation during forward terrestrial locomotion. We found that the anatomical femoral head of the alligator travels beyond the cranial extent of the bony acetabulum and does not act as a central pivot, as has been suggested for some extinct archosaurs. Additionally, the fibrocartilaginous surfaces of the alligator's antitrochanter and femoral neck remain engaged during hip flexion and extension, similar to the articulation between homologous structures in birds. Moreover, the femoral insertion of the ligamentum capitis moves dorsoventrally against the membrane-bound portion of the medial acetabular wall, suggesting that the inner acetabular foramen constrains the excursion of this ligament as it undergoes cyclical stretching during the step cycle. Finally, the articular surface of the femoral cartilage model interpenetrates with those of the acetabular labrum and antitrochanter menisci; we interpret such interpenetration as evidence of compressive deformation of the labrum and of sliding movement of the menisci. Our data illustrate the utility of XROMM for studying in vivo articular soft tissue interactions. These results also allow us to propose functional hypotheses for crocodilian hip joint soft tissues, expanding our knowledge of vertebrate connective tissue biology and the role of joint soft tissues in locomotor behavior.