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Partial Skeleton of Theropithecus Brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya

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Partial Skeleton of Theropithecus Brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya Journal of Human Evolution 61 (2011) 347e362 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol Partial skeleton of Theropithecus brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya Christopher C. Gilbert a,b,c,*, Emily D. Goble d, John D. Kingston e, Andrew Hill d a Department of Anthropology, Hunter College of the City University of New York, 695 Park Avenue, NY 10021, USA b Department of Anthropology, Graduate Center of the City University of New York, 365 Fifth Avenue, NY 10016, USA c New York Consortium in Evolutionary Primatology, New York, NY, USA d Department of Anthropology, Yale University, PO Box 208277, New Haven, CT 06520-8277, USA e Department of Anthropology, Emory University, 1557 Dickey Drive, Atlanta, GA 30322, USA article info abstract Article history: Here we describe a complete skull and partial skeleton of a large cercopithecoid monkey (KNM-TH Received 27 October 2010 46700) discovered in the Chemeron Formation of the Tugen Hills at BPRP Site #152 (2.63 Ma). Associated Accepted 21 April 2011 with the skeleton was a mandible of an infant cercopithecoid (KNM-TH 48364), also described here. KNM-TH 46700 represents an aged adult female of Theropithecus brumpti, a successful Pliocene papionin Keywords: taxon better known from the Omo Shungura Formation in Ethiopia and sites east and west of Lake Cercopithecoid Turkana, Kenya. While the morphology of male T. brumpti is well-documented, including a partial Papionin skeleton with both cranial and postcranial material, the female T. brumpti morphotype is not well- Crania fi Postcrania known. This skeleton represents some of the rst associated evidence of cranial and postcranial Baboons female T. brumpti remains. In addition to the complete skull, postcranial material includes elements of Plio-Pleistocene the axial skeleton and lower limb. While aspects of the skeleton conform to those of specimens previ- ously assigned to T. brumpti, other features on the femur and tibia appear to differ from those previously described for this species. It is unclear whether these differences represent general variation within the T. brumpti population, variation between the sexes in T. brumpti, or the incorrect assignment of previous isolated hindlimb specimens. In total, the observable morphological features of the hindlimb suggest that KNM-TH 46700 was a terrestrial quadruped similar to modern savannah baboons (Papio). From the available evidence, it is difficult to assess whether or not KNM-TH 46700 frequently engaged in the specialized squatting and shuffling behavior observed in extant geladas (Theropithecus gelada). Ó 2011 Elsevier Ltd. All rights reserved. Introduction extinct species’ paleobiology. In this case, accurate interpretations of T. brumpti’s locomotor behavior are also potentially important for Theropithecus brumpti is a well-known and distinctive fossil paleoenvironmental reconstructions. papionin commonly found at Turkana Basin paleontological sites Previous functional analyses of both associated and unassoci- from w3.5 Ma to 2.0 Ma (Leakey, 1993; Jablonski et al., 2008). The ated T. brumpti postcrania suggest a mix of arboreal and terrestrial species is easily recognized by its remarkable cranium, which is features (Ciochon, 1993; Krentz, 1993a,b; Jablonski et al., 2002, dominated by exceptionally large and flaring zygomatics. While 2008). In total, the available evidence suggests that T. brumpti hundreds of T. brumpti specimens have been recovered since its was a terrestrial quadruped that possessed features in the forelimb initial description (Arambourg, 1947), rarely are cranial and post- indicating it was also more arboreal than modern Theropithecus and cranial material found together in association. Even less common is Papio, a locomotor repertoire perhaps most similar to that of the the recovery of a partial or complete skeleton, a phenomenon that modern papionin genus Mandrillus (Ciochon,1993; Krentz, 1993a,b; is, unfortunately, rare throughout all of paleoanthropology. Well- Jablonski et al., 2002, 2008). The T. brumpti forelimb was further associated craniodental and skeletal remains are ultimately specialized for manual foraging, sharing derived characters such as crucial for comprehensive and accurate interpretations of an an elongated pollex and shortened second digit with modern geladas (Theropithecus gelada)(Jablonski, 1986; Krentz, 1993a,b; Jablonski et al., 2002). Also similar to modern geladas, the * Corresponding author. T. brumpti hindlimb is described as exhibiting features associated E-mail address: [email protected] (C.C. Gilbert). with a distinctive squatting and shuffling food-gathering behavior 0047-2484/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2011.04.007 348 C.C. Gilbert et al. / Journal of Human Evolution 61 (2011) 347e362 Figure 1. (a) Location of the Tugen Hills and Lake Baringo within the Kenyan Rift Valley. (b) Geologic map of the Chemeron Formation exposed in the Barsemoi, Ndau, and Kapthurin Rivers and associated drainages. Depicted are locations of vertebrate fossil localities including the location of BPRP #152 where the Theropithecus partial skeleton was found. (Krentz, 1993a). Such features of the hindlimb include a “reverse” to address some of the issues relating to the degree of arboreality or carrying angle of the femoral shaft and an angulated medial mal- terrestriality exhibited by T. brumpti, the number of derived features leolus of the tibia (Krentz, 1993a). shared with modern geladas, as well as any perceived differences Recently, a partial skeleton of a T. brumpti adult male was between male and female postcranial morphology and how this described from sediments dating to w3.3 Ma at the site of Lomekwi, affects locomotor reconstruction for this species. In addition, the West Turkana, Kenya, detailing many aspects of male T. brumpti new specimen allows us to comment on variation within the species morphology (Jablonski et al., 2002). Thus, from previous research on and morphological variation in females in particular. While the abundant craniodental and unassociated postcranial remains (e.g., current analysis is not intended to serve as a comprehensive Eck and Jablonski, 1987; Ciochon, 1993; Krentz, 1993a,b), as well as comparative study of papionin postcranial morphology and levels of the more recent description of the associated skeleton, the variation, the description of KNM-TH 46700 allows us to make some morphology of males is relatively well-known. In contrast, fewer preliminary observations with regards to these issues. complete cranial and postcranial remains represent the female morphotype. Because of their large size, T. brumpti males have been Geological context and associated fauna suggested by some to be unlikely arborealists (Jablonski et al., 2002, 2008), but by others to have been more arboreal than modern Fossiliferous sediments exposed west of Lake Baringo (Fig. 1a) in geladas or Theropithecus oswaldi (Krentz, 1993a). Given the large the Kenyan Rift Valley comprise the most chronologically extensive amount of sexual dimorphism and large differences in estimated Neogene clastic sequence known in East Africa, spanning the last body mass between T. brumpti male and female specimens (males 16 m.yr. (Fig. 2)(Hill, 2002). Within this succession, the Chemeron w36 kg, females w24 kg; see Delson et al., 2000), it is possible that Formation encompasses a series of discontinuous sediments and males and females exhibited slightly different frequencies of arbo- tuffs exposed over 40 km along the eastern foothills of the Tugen real locomotor activities. However, such differences between Hills. Chemeron deposits span over 3.7 m.yr. from about 5.3 Ma at T. brumpti males and females remain to be demonstrated. the base to less than 1.6 Ma at the top (Fig. 2)(Deino and Hill, 2002; Here, we describe a partial skeleton of an aged T. brumpti adult Deino et al., 2002). female recovered from BPRP site #152 in the Chemeron Formation In the southernmost exposures of the Chemeron Formation, of the Tugen Hills, Kenya. The associated partial skeleton allows us where BPRP #152 is situated (Fig. 1b), sediments are exposed as an C.C. Gilbert et al. / Journal of Human Evolution 61 (2011) 347e362 349 Figure 2. Generalized composite stratigraphic framework of the Tugen Hills Succession, indicating the stratigraphic level of BPRP #152 in the Chemeron Formation. eastward-dipping structural block (w20e30) and consist of the Tugen Hills in having the suid Metridiochoerus andrewsi as part terrigenous and lacustrine sediments, primarily mudstone, silt- of it. This is a relatively common pig in other east African regions stone, and sandstone with intercalations of tuff, diatomite, and over the time range represented by the Chemeron Formation, but conglomerate. Within this general succession, a distinctive litho- the four individuals at BPRP #152 are the only fossils of M. logic package characterized by a series of diatomite units and andrewsi so far known from the many sites in the entire Chemeron interbedded fluvial and alluvial fan detritus and tuffs can be traced succession (Bishop et al., 1999). The reason for the almost total >10 km NeS along strike. These sediments document significant, absence of M. andrewsi in the Chemeron, and for its so far exclu- intermittent lake systems within the axial portion of the rift that sive presence at this particular site, is not yet understood.
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