Reconstruction of body cavity volume in terrestrial tetrapods.

Although it is generally assumed that herbivores have more voluminous body cavities due to larger digestive tracts required for the digestion of plant fiber, this concept has not been addressed quantitatively. We estimated the volume of the torso in 126 terrestrial tetrapods (synapsids including basal synapsids and mammals, and diapsids including birds, non-avian dinosaurs and reptiles) classified as either herbivore or carnivore in digital models of mounted skeletons, using the convex hull method. The difference in relative torso volume between diet types was significant in mammals, where relative torso volumes of herbivores were about twice as large as that of carnivores, supporting the general hypothesis. However, this effect was not evident in diapsids. This may either reflect the difficulty to reliably reconstruct mounted skeletons in non-avian dinosaurs, or a fundamental difference in the bauplan of different groups of tetrapods, for example due to differences in respiratory anatomy. Evidently, the condition in mammals should not be automatically assumed in other, including more basal, tetrapod lineages. In both synapsids and diapsids, large animals showed a high degree of divergence with respect to the proportion of their convex hull directly supported by bone, with animals like elephants or Triceratops having a low proportion, and animals such as rhinoceros having a high proportion of bony support. The relevance of this difference remains to be further investigated.

Toe function and dynamic pressure distribution in ostrich locomotion.

The ostrich is highly specialized in terrestrial locomotion and is the only extant bird that is both didactyl and exhibits a permanently elevated metatarsophalangeal joint. This extreme degree of digitigrady provides an excellent opportunity for the study of phalangeal adaptation towards fast, sustained bipedal locomotion. Data were gathered in a semi-natural setting with hand-raised, cooperative specimens. Dynamic pressure distribution, centre of pressure (CoP) trajectory and the positional inter-relationship of the toes during stance phase were investigated using pedobarography. Walking and running trials shared a J-shaped CoP trajectory with greater localization of CoP origin as speed increased. Slight variations of 4th toe position in walking affect CoP origin and modulation of 4th toe pressure on the substrate allows correction of balance, primarily at the beginning of stance phase at lower speeds. Load distribution patterns differed significantly between slow and fast trials. In walking, the 3rd and particularly the 4th toe exhibited notable variation in load distribution with minor claw participation only at push-off. Running trials yielded a distinctly triangular load distribution pattern defined by the 4th toe tip, the proximal part of the 3rd toe and the claw tip, with the sharp point of the claw providing an essential traction element at push-off. Consistency of CoP trajectory and load distribution at higher speeds arises from dynamic stability effects and may also reflect stringent limitations to degrees of freedom in hindlimb joint articulation that contribute to locomotor efficiency. This novel research could aid in the reconstruction of theropod locomotor modes and offers a systemic approach for future avian pedobarographic investigations.

The intertarsal joint of the ostrich (Struthio camelus): Anatomical examination and function of passive structures in locomotion.

The ostrich (Struthio camelus) is the largest extant biped. Being flightless, it exhibits advanced cursorial abilities primarily evident in its characteristic speed and endurance. In addition to the active musculoskeletal complex, its powerful pelvic limbs incorporate passive structures wherein ligaments interact with joint surfaces, cartilage and other connective tissue in their course of motion. This arrangement may enable energy conservation by providing joint stabilisation, optimised limb segment orientation and automated positioning of ground contact elements independently of direct muscle control. The intertarsal joint is of particular interest considering its position near the mid-point of the extended limb and its exposure to high load during stance with significant inertial forces during swing phase. Functional-anatomical analysis of the dissected isolated joint describes the interaction of ligaments with intertarsal joint contours through the full motion cycle. Manual manipulation identified a passive engage-disengage mechanism (EDM) that establishes joint extension, provides bi-directional resistance prior to a transition point located at 115 degrees and contributes to rapid intertarsal flexion at toe off and full extension prior to touch down. This effect was subsequently quantified by measurement of intertarsal joint moments in prepared anatomical specimens in a neutral horizontal position and axially-loaded vertical position. Correlation with kinematic analyses of walking and running ostriches confirms the contribution of the EDM in vivo. We hypothesise that the passive EDM operates in tandem with a stringently coupled multi-jointed muscle-tendon system to conserve the metabolic cost of locomotion in the ostrich, suggesting that a complete understanding of terrestrial locomotion across extinct and extant taxa must include functional consideration of the ligamentous system.