Morphological systematics of the kipunji (Rungwecebus kipunji) and the ontogenetic development of phylogenetically informative characters in the Papionini

Since its discovery and description, the systematic position of the kipunji (Rungwecebus kipunji) has been a matter of debate. Although it was first placed in the mangabey genus Lophocebus, subsequent molecular studies indicated that the kipunji is most closely related to baboons (Papio). However, the kipunji does not appear to possess cranial features typical of Papio, thus necessitating the erection of a new genus, Rungwecebus. The recovery of an M2-stage subadult male kipunji voucher specimen, in addition to the original M1-stage subadult male voucher specimen, has since allowed further study. Here, we describe the craniodental morphology of the newly acquired kipunji specimen and present a phylogenetic analysis of Rungwecebus craniodental morphology using quantitative and qualitative characters. We examined the skulls of 76 M1- and M2-stage subadult males representing all extant papionin genera, taking note of character states that are static throughout ontogeny. To control for ontogenetic changes, only those characters expressing unchanged character states between subadult and adult specimens were coded for Rungwecebus and entered into a larger, recently published 151-character matrix of adult male morphology. To account for allometry, the narrow allometric coding method and the general allometric coding method were applied. The resulting most parsimonious trees suggest that Rungwecebus is phylogenetically closest to Lophocebus, a result consistent with initial morphological descriptions. However, due to the large amount of missing data for Rungwecebus, there are low bootstrap support values associated with any relationships within the larger Theropithecus/Papio/Lophocebus/Rungwecebus grouping. Taken in combination with previous molecular, phenetic, and ecological studies, the results of this study suggest that Rungwecebus is best regarded as a distinct genus closely related to Papio, Lophocebus, and Theropithecus. Adult morphological specimens are necessary to fully understand the adult kipunji morphotype, and its phylogenetic position will only be more precisely resolved with additional morphological and molecular data.