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Authors requiring further information regarding Elsevier’s archiving and manuscript policies are encouraged to visit: http://www.elsevier.com/copyright Author's personal copy Journal of Human Evolution 60 (2011) 731e745 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol Morphological systematics of the kipunji (Rungwecebus kipunji) and the ontogenetic development of phylogenetically informative characters in the Papionini Christopher C. Gilbert a,b,c,*, William T. Stanley d, Link E. Olson e, Tim R.B. Davenport f, Eric J. Sargis g,h a Department of Anthropology, Hunter College of the City University of New York, 695 Park Avenue, New York, NY 10021, USA b Department of Anthropology, Graduate Center of the City University of New York, 365 Fifth Avenue, New York, NY 10016, USA c New York Consortium in Evolutionary Primatology, New York, NY, USA d Department of Zoology, The Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605, USA e Department of Mammalogy, University of Alaska Museum, 907 Yukon Drive, Fairbanks, AK 99775, USA f Wildlife Conservation Society, Tanzania Program, P.O. Box 922, Zanzibar, Tanzania g Department of Anthropology, Yale University, P.O. Box 208277, New Haven, CT 06520-8277, USA h Divisions of Vertebrate Zoology and Vertebrate Paleontology, Peabody Museum of Natural History, Yale University, New Haven, CT 06520, USA article info abstract Article history: Since its discovery and description, the systematic position of the kipunji (Rungwecebus kipunji) has been Received 22 May 2010 a matter of debate. Although it was first placed in the mangabey genus Lophocebus, subsequent Accepted 8 January 2011 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 Keywords: a new genus, Rungwecebus. The recovery of an M2-stage subadult male kipunji voucher specimen, in Rungwecebus addition to the original M1-stage subadult male voucher specimen, has since allowed further study. Here, Papionin we describe the craniodental morphology of the newly acquired kipunji specimen and present Cercopithecine Crania a phylogenetic analysis of Rungwecebus craniodental morphology using quantitative and qualitative Juvenile characters. We examined the skulls of 76 M1- and M2-stage subadult males representing all extant Subadult papionin genera, taking note of character states that are static throughout ontogeny. To control for Phylogeny ontogenetic changes, only those characters expressing unchanged character states between subadult and Ontogeny adult specimens were coded for Rungwecebus and entered into a larger, recently published 151-character Allometry 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. Ó 2011 Elsevier Ltd. All rights reserved. Introduction Since its discovery and description, the kipunji’s systematic posi- tion has been a matter of some conjecture. On the basis of The kipunji (Rungwecebus kipunji) is an endemic Tanzanian morphological and behavioral observations in the field as well as papionin monkey known only from the Mt. Rungwe-Livingstone from photographs, Jones et al. (2005) first placed the kipunji in the forests in the Southern Highlands and the Ndundulu Forest in the mangabey genus Lophocebus as a new species (L. kipunji). Subse- Udzungwa Mountains (Jones et al., 2005; Davenport et al., 2006). quently, a subadult male voucher specimen with erupted M1s (FMNH 187122) was recovered from the type locality on Mt. Run- gwe in Tanzania. Morphological analysis of this specimen revealed * Corresponding author. further similarities with Lophocebus; however, molecular analyses E-mail address: [email protected] (C.C. Gilbert). on the basis of DNA obtained from this same specimen suggested 0047-2484/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2011.01.005 Author's personal copy 732 C.C. Gilbert et al. / Journal of Human Evolution 60 (2011) 731e745 that the kipunji is not a member of Lophocebus but is instead most might display (see Davenport et al., 2006), particularly because closely related to baboons (Papio)(Davenport et al., 2006; Olson papionin cranial shape is known to change throughout ontogeny et al., 2008). These findings necessitated the erection of a new (e.g., Freedman, 1962; Leigh et al., 2003; Leigh, 2006, 2007). genus, Rungwecebus, to accommodate the kipunji (R. kipunji), Singleton et al. (2010) used 3-D geometric morphometrics and which is morphologically distinct from Papio (Jones et al., 2005; different papionin models of ontogenetic cranial development to Davenport et al., 2006). A recent, detailed 3-D geometric morpho- predict the kipunji’s adult morphology, but it is impossible to know metric study of FMNH 187122 has also documented the phenetic how accurate these estimates are in the absence of an older kipunji distinctiveness of the kipunji, not only relative to Papio, but to the specimen for direct comparison. In addition, as Singleton et al. other papionin genera as well (Singleton, 2009). (2010) pointed out, it is difficult to estimate the adult expression of Recently, a series of studies has been published regarding the some phylogenetically informative qualitative morphological char- molecular systematics and origins of the kipunji. After acters, such as the orientation of the temporal and nuchal lines. the initial molecular work by Davenport et al. (2006) and Olson The phylogenetic position of Rungwecebus is not only important et al. (2008), Zinner et al. (2009) and Burrell et al. (2009) in and of itself, but also to the broader grouping of the Rungwece- demonstrated that the mtDNA of the kipunji from Mt. Rungwe is bus/Lophocebus/Papio/Theropithecus clade within the African not only nested within Papio, but is most similar to the mtDNA of papionins. Previous morphological and molecular phylogenetic the yellow baboon (P. cynocephalus) populations adjacent to the studies have failed to resolve the relationships within the Lopho- kipunji’s known distribution in Tanzania. Both studies suggested cebus/Papio/Theropithecus grouping (Disotell et al., 1992; Disotell, that the introgression of baboon mtDNA into the kipunji genome 1994, 2000; Harris and Disotell, 1998; Harris, 2000; Tosi et al., was the result of multiple hybridization events between kipunjis 2003; Gilbert and Rossie, 2007; Gilbert et al., 2009a), and the and adjacent yellow baboon populations. However, these studies addition of Rungwecebus adds new combinations of character states offered different hypotheses regarding the origins of the kipunji. and new information about character state polarities that could Burrell et al. (2009) hypothesized that the kipunji may be the result improve our understanding of the phylogenetic relationships of a relatively recent (w0.65 Ma) hybrid speciation event that within this clade. A greater understanding of phylogeny is neces- occurred between a population of Lophocebus mangabeys and sary to resolve lingering questions about morphological and yellow baboons, thereby explaining the presence of yellow baboon behavioral evolution within this clade. mtDNA in the kipunji genome. Alternatively, Zinner et al. (2009) In this paper, we describe the morphology of a recently hypothesized that the kipunji is the sister lineage to Papio and vouchered M2-stage subadult male kipunji specimen, Southern has since experienced an unknown degree of hybridization with Highlands Conservation Programme (SHCP) 2458, and compare it adjacent baboon populations. This scenario would explain why to the original M1-stage subadult voucher specimen (FMNH Zinner et al. (2009) found the kipunji nuclear genome to be distinct 187122). This additional, ontogenetically advanced specimen from Papio while observing that the mtDNA is most similar to allows for a more confident estimation of the kipunji’s adult male adjacent yellow baboon populations. morphology. We describe the specimen, and outline and summa- This