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Nacholapithecus Kerioi

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Nacholapithecus Kerioi Anthropological Science Advance Publication Material Report New forelimb long bone specimens of Nacholapithecus kerioi from the Middle Miocene of northern Kenya Tomo Takano1*, Masato Nakatsukasa2, Marta Pina3, Yutaka Kunimatsu4, Yoshihiko Nakano5, Naoki Morimoto2, Naomichi Ogihara6**, Hidemi Ishida7 1Japan Monkey Centre, Inuyama, Aichi 484-0081, Japan 2Laboratory of Physical Anthropology, Graduate School of Science, Kyoto University, Kyoto 606-8502, Japan 3Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain 4Faculty of Business Administration, Ryukoku University, Kyoto 612-8577, Japan 5Laboratory of Biological Anthropology, Department of Human Sciences, Osaka University, Osaka 565-0871, Japan 6Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Tokyo 113-0033, Japan 7Professor Emeritus, Kyoto University, Kyoto 606-8502, Japan Received 9 May 2019; accepted 16 January 2020 Abstract This article reports eight new humeral, ulnar, and radial fragments of Nacholapithecus keri- oi collected from Nachola, Kenya during the 1998/1999 field seasons. The study refines the description of its forelimb bones, which was mostly based on a single partial skeleton. The most distinctive feature of the distal humerus is a large, globular, medially tilted capitulum. The groove between the capitulum and the zona conoidea is quite deep. The medial part of the humeral trochlea is also diagnostic in show- ing a less salient medial border. The medial epicondyle is moderately long and more posteriorly reflected than was previously presumed. The coronoid process of the ulna is quite wide. Its medial portion is dis- tinctly concave. The ulnar shaft is anteroposteriorly deep in its proximal half, slender, straight in frontal view, and weakly anteriorly bowing. The elbow of Nacholapithecus exhibits a primitive functional pat- tern as a hominoid, including lack of universal stability of the humeroulnar joint through full extension and flexion, restriction of hyperextension of the elbow, and relatively anteroposteriorly oriented loading at the proximal ulna. On the other hand, it is derived in terms of enhanced rotational mobility and stabil- ity of the radius, incipiently increased stability at the humeroulnar joint, and more frequent maximum extension of the elbow compared to proconsulids. This mosaic morphology is different from both early Miocene proconsulids and later suspensory or orthograde European fossil apes. Although Na- cholapithecus was neither suspensory nor orthograde, its forelimbs may have played a greater role for body support or balance maintenance, more frequently reaching to and exploiting overhead supports than in early Miocene proconsulids. Key words: fossils, hominoid evolution, humerus, ulna, radius 1999). Since KNM-BG 35250 includes many skeletal ele- Introduction ments, comparative anatomical study has largely relied on it. Nacholapithecus kerioi is a Middle Miocene (16–15 Ma) However, unfortunately, most fossils from BG-K, including hominoid (kenyapithecine: Alba et al., 2012) discovered at KNM-BG 35250, suffer from plastic deformation and/or the Nachola, northern Kenya (Ishida et al., 1999; Nakatsukasa cracking due to compression during fossilization. Due to this and Kunimatsu, 2009; Kunimatsu et al., 2019). The majority taphonomic issue, proper characterization of Nacholapithecus of Nacholapithecus specimens, including the holotype anatomy is not easy (e.g. Takano et al., 2018). KNM-BG 35250 partial skeleton, have been collected at site After the discovery of KNM-BG 35250, excavation con- BG-K at Nachola (Nakatsukasa et al., 1998; Ishida et al., tinued at the fossiliferous site BG-K, and additional fossil specimens, including several forelimb bones, were un- * Correspondence to: Tomo Takano, Japan Monkey Centre, Kanrin earthed during the 1998/1999 field seasons. This article re- 26, Inuyama, Aichi 484-0081, Japan. ports on those specimens to amend the current description of E-mail: [email protected] the forelimb long bones of Nacholapithecus (Takano et al., ** Correspondence to: Naomichi Ogihara, Department of Biological Sciences, Graduate School of Science, The University of Tokyo, 2018). Notably, this collection includes a well-preserved Tokyo 113-0033, Japan. proximal portion of ulna with relatively minor deformation. E-mail: [email protected] A morphometric study of Nacholapithecus ulnae is being Published online 17 March 2020 prepared as a separate analytical paper. in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.200116 © 2020 The Anthropological Society of Nippon 1 2 T. TAKANO ET AL. Anthropological Science Table 1. New forelimb specimens of Nacholapithecus Accession no. Part side Collection information KNM-BG 38384 distal humerus R BG-K in situ (Trench 99-2) KNM-BG 37303 humeral trochlea fragment R BG-K surface collection (1998) KNM-BG 38610A humeral shaft R BG-K in situ (Trench 99-2) KNM-BG 38391B proximal ulna R BG-K in situ (Trench 99-2) KNM-BG 38610B proximal ulna without olecranon R BG-K in situ (Trench 99-2) KNM-BG 37352 proximal ulna without olecranon L BG-K surface collection (1998) KNM-BG 38387 ulnar shaft R BG-K in situ (1999) KNM-BG 40021 proximal radius R BG-K in situ (Trench 99-2) Materials and Methods Descriptions The material reported here consists of three humeral, four KNM-BG 38384 (distal portion of right humerus) ulnar, and one radial fragment (Table 1). All specimens are This specimen is a c. 8 cm long distal portion of a right stored at the Earth Science Department, the National Muse- humerus (Figure 1). The size of this specimen is comparable ums of Kenya (KNM), and are now fully accessible for to that of the previously described KNM-BG 35250M study. Nacholapithecus is highly dimorphic in body size (Table 2). (Kikuchi et al., 2018). Sexing of new specimens was done The shaft is smashed anteroposteriorly and displays an based on their size using KNM-BG 35250 as the male refer- exaggeratedly wide appearance. The olecranon fossa is dam- ence. All of these specimens comfortably fall in the male aged and perforated. Thus, the anterior surface where the category due to their large size. coronoid fossa would have been is broken. The distal joint is A digital sliding caliper was used to take linear measure- better preserved than that of KNM-BG 35250M although the ments to a tenth of millimeter. Measurements and indices are trochlea is cracked proximomedially to distolaterally on the the same as those published by Takano et al. (2018) (Table 2, anterior surface (arrows in Figure 1a) and the articular sur- Table 3, Table 4. Table 5). Because most of these specimens face proximal to the crack is displaced posteriorly (arrow in are subject to plastic deformation and/or cracking, damage Figure 1f). was evaluated from computed tomography (CT) images cap- Due to the severe damage, the original cross-sectional tured with a CT scanner (X-CT Research SA+, Stratec Mediz- shape of the shaft is unknowable. The lateral supracondylar intechnik GmbH, Germany) with a resolution of 0.1 mm. ridge is slightly eroded. It lacks a prominent lateral flare. The Comparative extant primate specimens used in this study are lateral epicondyle is massive and proximodistally long. It is the same as those used in Takano et al. (2018). For illustrative distorted and deflected slightly anteriorly (Figure 1c). The purposes (visual comparison and virtual restoration), some height of the lateral epicondyle from the distal surface of the original fossil specimens were surface-scanned with a capitulum is 23.5 mm. The radial fossa is shallow. NextEngine 3D scanner (Nextengine Inc., Santa Monica, CA, The capitulum is large, tall, and globular (Table 2, USA) using the Macro and HD (high point density) settings. Table 3). The proximodistal height is 16.2 mm. The capitu- Figure 1. KNM-BG 38384 right humerus: (a) anterior; (b) posterior; (c) lateral; (d) medial; (e) distal views. (f) CT section at the level of the dotted lines in (a). Despite artifacts due to the beam hardening effect, regularly aligned trabeculae are observed in the capitulum and the medial part of trochlea, providing evidence that the original structure is locally maintained. Arrows in (a) and (f) indicate cracking and displacement. NEW FORELIMB SPECIMENS OF NACHOLAPITHECUS 3 Table 2. Humeral measurements. Comparative data are from Takano et al. (2018). Measurement KNM-BG 38384 KNM-BG 37352 KNM-BG35250M Definition Bicondylar breadth (>43.9)1 — (<53.4)1 (c) in Rose et al. (1992) Capitular height 16.2 — (<16.9) (f) in Rose (1988) Articular width 31.3 — (<38.7) (a) in Rose (1988) Capitular + zona width (17.4) — 18.9 (d) in Rose (1988) Trochlear width (anterior) (>13.9) (<18.9) (e) in Rose (1988) Capitular width 12.7 — 14.1 (i) in Rose (1988) Medial trochlear rim height 16.1 15.02 — (c) in Rose (1988) Lateral trochlear rim height — 13.72 (13.9) (j) in Rose (1988) Trochlear notch height (>12.1) 10.52 (11.1) (k) in Rose (1988) Posterior breadth of distal articulation 19.1 — (15.8) (1) in Harrison (1982) 1 Values in parentheses represent estimates or values affected by deformation. 2 Due to damage, measured in the left counterpart. Table 3. Humeral ratios. Comparative data are from Takano et al. (2018). Medial trochlear rim Articular width/ Capitular + zona Capitular height/ Taxon Specimen height/lateral trochlear bicondylar width width/trochlear width capitular width rim height Nacholapithecus KNM-BG 38384 (0.71) (1.25>)
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