The thumb of Miocene apes: new insights from Castell de Barberà (Catalonia, Spain).

Primate hands display a major selective compromise between locomotion and manipulation. The thumb may or may not participate in locomotion, but it plays a central role in most manipulative activities. Understanding whether or not the last common ancestor of humans and Pan displayed extant-ape-like hand proportions (i.e., relatively long fingers and a short thumb) can be clarified by the analysis of Miocene ape hand remains. Here we describe new pollical remains-a complete proximal phalanx and a partial distal phalanx-from the middle/late Miocene site of Castell de Barberà (ca., 11.2-10.5 Ma, Vallès-Penedès Basin), and provide morphometric and qualitative comparisons with other available Miocene specimens as well as extant catarrhines (including humans). Our results show that all available Miocene taxa (Proconsul, Nacholapithecus, Afropithecus, Sivapithecus, Hispanopithecus, Oreopithecus, and the hominoid from Castell de Barberà) share a similar phalangeal thumb morphology: the phalanges are relatively long, and the proximal phalanges have a high degree of curvature, marked insertions for the flexor muscles, a palmarly bent trochlea and a low basal height. All these features suggest that these Miocene apes used their thumb with an emphasis on flexion, most of them to powerfully assist the fingers during above-branch, grasping arboreal locomotion. Moreover, in terms of relative proximal phalangeal length, the thumb of Miocene taxa is intermediate between the long-thumbed humans and the short-thumbed extant apes. Together with previous evidence, this suggests that a moderate-length hand with relatively long thumb-involved in locomotion-is the original hand morphotype for the Hominidae.