Mechanisms of hind foot reversal in climbing mammals

Many climbing mammals are able to reverse normal hind foot posture to effect the grip necessary to descend headfirst or to hang upside down. Such hind foot reversal is known in sciurids, procyonids, felids, viverrids, tupaiids, prosimians, and marsupials. The joint movements involved, however, have never been documented unequivocally although various interpretations (some contradictory) have been made. We report here radiographic data from species of the genera Didelphis, Felis, Nasua, Nycticebus, Potos, Sciurus, and Tupaia. In the six eutherians studied, three joints are involved, and there is a common pattern in the mechanism: crurotalar plantarflexion, subtalar inversion, and transverse tarsal supination. Hind foot reversal represents the development of an unusual degree of excursion at these joints, rather than the appearance of any new type of movement. In Didelphis the mechanism is quite different: a bicondylar, spiral tibiotalar joint is the principal site of inversion/abduction movements. This specialization is characteristic of didelphids and phalangerids, and occurs in the extinct multituberculates as well; it is not found in macropodids (which are like eutherians in crurotalar joint structure) or other marsupial families. This diversity in pedal structure and function is evidently the result of parallel evolution from the type of tibiotalar joint of cynodonts and early mammals. In Morganucodon the bulbous, hemispheroidal proximal surface of the talus bears two tibial facets. These facets are represented in didelphids and multituberculates as sulci, whereas in macropodids and eutherians they developed as the proximal and medial surfaces of the talar trochlea. Among living mammals, the primitive hemispheroidal joint is retained among monotremes as a ball and socket joint.