Parallel evolution of hand anatomy in kangaroos and vombatiform marsupials: Functional and evolutionary implications

The anatomy of the mammalian hand is exposed to an intriguing interplay between phylogeny and function, and provides insights on phylogenetic affinities as well as locomotory habits of extinct species. Within the marsupial order Diprotodontia, terrestrial plantigrade quadrupedalism evolved twice, in the mostly extinct vombatiforms and in extant macropodoids. To assess the influence of functional and phylogenetic signal on the manus in these two clades, manual anatomy and digital proportions in specimens of eight extinct and three extant vombatiforms were investigated and compared with extant macropodoids and extant possums. The results reveal extensive parallelisms in the carpal region of vombatiforms and macropodoids, including flattened distal metacarpal facets, reduction of the palmar process of the hamatum, reduction of mid-wrist joint curve, extensive hamatum/scaphoid contact, and absence of a lunatum. These transformations appear to be related to stabilization of the wrist for plantigrade locomotion. Vombatiforms are apomorphic in scaphoid and triquetrum anatomy and their metacarpals are much more gracile than in other Diprotodontia. Manual diversity is greater in vombatiforms than in macropodoids, as probably was locomotor diversity. Digital proportions as well as wrist anatomy divide the extinct vombatiforms into species resembling arboreal diprotodontians, whereas others group with terrestrial quadrupedal kangaroos and wombats. The latter is suggested to be owing to plantigrade locomotion and/or large size. Carpal anatomy and digital proportions suggest that a range of earlier diverging vombatiforms may have been arboreal or scansorial. As such, we propose that the ancestor of extant vombatiforms (koalas and wombats) may have been arboreal, an option that deserves consideration in the reconstruction of vombatiform evolution.