Geobios 50 (2017) 197–209 Available online at ScienceDirect www.sciencedirect.com Original article A new Elasmotheriini (Perissodactyla, Rhinocerotidae) from the upper § Miocene of Samburu Hills and Nakali, northern Kenya a, b c d Naoto Handa *, Masato Nakatsukasa , Yutaka Kunimatsu , Hideo Nakaya a Museum of Osaka University, 1-20 Machikaneyama-cho, Toyonaka, Osaka 560-0043, Japan b Laboratory of Physical Anthropology, Graduate School of Science, Kyoto University, Kitashirakawa-oiwakecho, Sakyo-ku, Kyoto 606-8502, Japan c Faculty of Business Administration, Ryukoku University, 67 Tsukamoto-cho, Fukakusa, Fushimi-ku, Kyoto 612-8577, Japan d Graduate School of Science and Engineering, Kagoshima University, 1-21-35 Korimoto, Kagoshima 890-0065, Japan A R T I C L E I N F O A B S T R A C T Article history: Several rhinocerotid cheek teeth and mandibular fragments from the upper Miocene of the Samburu Received 15 August 2016 Hills and Nakali in northern Kenya are described. These specimens show characteristics that place them Accepted 20 April 2017 in the Tribe Elasmotheriini such as a constricted protocone, a developed antecrochet, and coronal Available online 8 May 2017 cement. The present specimens are compared with other Elasmotheriini species from Eurasia and sub- Saharan East Africa. They are found to be morphologically different from the previously known species of Keywords: Elasmotheriini. Morphologically, they are most similar to Victoriaceros kenyensis from the middle Elasmotheriini Miocene of Kenya, but differ from V. kenyensis in having the upper molars with the simple crochet, Rhinocerotidae lingual groove of the protocone and enamel ring in the medisinus. Therefore, the present specimens are Late Miocene assigned to a new genus and species of Elasmotheriini: Samburuceros ishidai. A cladistic analysis Samburu Hills Nakali tentatively places S. ishidai nov. gen., nov. sp. as a sister taxon of V. kenyensis. However, questions remain Kenya regarding a further detailed discussion of the phylogenetic relationship between the African Elasmotheriini and other Eurasian taxa because of the incompleteness of the specimens from Africa, as already noticed by several researchers. C 2017 Elsevier Masson SAS. All rights reserved. 1. Introduction and Gue´rin, 1974), and later in the late Miocene locality of the Samburu Hills in Kenya (Nakaya et al., 1987). Ougandatherium The Tribe Elasmotheriini (Heissig, 1973, 1989; = Elasmotheriina napakense was discovered in the early Miocene (20–19 Ma) locality in Antoine, 2002, 2003), an extinct group of Rhinocerotidae of Napak in Uganda (Gue´rin and Pickford, 2003). Geraads et al. (Mammalia, Perissodactyla), was widely distributed in Europe, (2012) described Victoriaceros kenyensis from the middle Miocene Asia and sub-Saharan East Africa, especially during the early to locality (ca. 15.5 Ma) of Maboko in Kenya. Recently, Geraads et al. middle Miocene of Europe (Cerden˜o and Nieto, 1995; Heissig, (2016) described a skull from the early Miocene (> 17.7 Ma) 1996, 1999), early to middle Miocene of Pakistan (Heissig, 1972; locality of Karungu in western Kenya, and tentatively assigned it as Antoine and Welcomme, 2000), early Miocene to late Pleistocene a second species of Victoriaceros, V. hooijeri. of China (Tong and Moigne, 2000; Deng and Downs, 2002; Antoine, The Kenya–Japan joint expedition carried out fieldwork in the 2003), and Pleistocene of Russia (Schvyreva, 2015). The first known Samburu Hills during the 1980s and 1990s (Ishida and Pickford, Elasmotheriini are from the early Miocene of Pakistan (Antoine and 1997) and has been working in Nakali since 2002 (Kunimatsu et al., Welcomme, 2000). This tribe diversified during the early to middle 2007). Abundant mammal fossils, including the large hominoid Miocene in Eurasia and became extinct during the Pleistocene in fossils Samburupithecus kiptalami and Nakalipithecus nakayamai, Europe and Asia (Heissig, 1989). have been discovered in these localities. Rhinocerotid fossils were In contrast, only a few species have been reported so far from also discovered through fieldwork in those localities (Nakaya et al., sub-Saharan East Africa (Fig. 1). Kenyatherium bishopi was first 1987; Kunimatsu et al., 2007; Fukuchi et al., 2008; Handa et al., discovered in the late Miocene locality of Nakali in Kenya (Aguirre 2015; Handa, 2016). Nakaya (1994) reported a taxon of rhinoce- rotid from the upper Miocene Namurungule Formation in the Samburu Hills as ‘‘Iranotheriinae sp. nov.’’. This taxon was first § Corresponding editor: Pierre-Olivier Antoine. reported as Rhinocerotidae gen. et sp. indet. (Nakaya et al., 1987). * Corresponding author. Tsujikawa (2005) cited the identification of Nakaya (1994) as the E-mail address: [email protected] (N. Handa). http://dx.doi.org/10.1016/j.geobios.2017.04.002 C 0016-6995/ 2017 Elsevier Masson SAS. All rights reserved. 198 N. Handa et al. / Geobios 50 (2017) 197–209 Fig. 1. Map showing the African Elasmotherhiini-bearing localities. Modified from Sawada et al., 1998. number of species of rhinocerotids from the Namurungule this area (Kunimatsu et al., 2007; Sakai et al., 2013). The Nakali Formation. However, a detailed taxonomic description of those Formation is unconformably overlaid by the Nasorut Formation, specimens has not been undertaken since Nakaya (1994). Further- which is composed of trachytic or basaltic lava and volcaniclastics. more, a few undescribed specimens that would belong to The thickness of the Nakali Formation is about 340 m. This ‘‘Iranotheriinae sp. nov.’’ were also found in the Nakali Formation formation is divided into three units: the Lower, Middle, and Upper in Nakali. Here, we describe those specimens as a new genus and Members in ascending order (Kunimatsu et al., 2007; Sakai et al., species of Elasmotheriini. 2013). The Lower Member is composed mainly of lacustrine and fluvio-lacustrine deposits. The Middle Member consists of a pyroclastic flow deposit, whose thickness is ca. 40 m. The Upper 2. Geological setting Member is characterized by fluvio-lacustrine and lacustrine deposits. Nakalipithecus nakayamai was collected from the Upper The Samburu Hills is located 50 km south of Lake Turkana Member (Kunimatsu et al., 2007). The studied rhinocerotid (Fig. 1). The Miocene succession (including the Nachola, Aka specimens from the Nakali Formation were found in the Upper 40 39 Aiteputh, Namurungule, and Kongia formations) and Pliocene and Lower Members. Ar/ Ar dating provided ages of 9.82 Æ 0.09 succession (corresponding to the Tirr Tirr Formation) are and 9.90 Æ 0.09 Ma for the uppermost part of the Lower Member of distributed in this area (Saneyoshi et al., 2006; Sakai et al., this formation (Kunimatsu et al., 2007). The paleomagnetic stratigra- 2010). Among them, the Namurungule Formation overlies the Aka phy of the uppermost level of the Lower Member and the lowermost Aitheputh Formation conformably and underlies the Kongia level of the Upper Member correlates with Chron C5n.1r (9.88 to Formation unconformably. Some parts of the boundary between 9.92 Ma; Kunimatsu et al., 2007). the Aka Aiteputh and Namurungule formations are local faults. The Namurungule Formation is about 200 m thick and is divided into the Upper and Lower Members, which consist of alluvial fan and 3. Review of the Rhinocerotidae from the Namurungule and fluvio-lacustrine, and fluvio-lacustrine delta and fluvial deposits, Nakali formations respectively (Saneyoshi et al., 2006; Sakai et al., 2010). Samburupi- thecus kiptalami was found from the Lower Member (Ishida and Nakaya et al. (1984, 1987) reported abundant specimens of the Pickford, 1997). There is a lahar deposit between the Lower and Rhinocerotidae from the Samburu Hills. After that, additional Upper Members. The studied rhinocerotid specimens from the mammalian fossils were recovered through the fieldwork sessions Namurungule Formation were discovered from the Lower Mem- in 1990s (Tsujikawa, 2005). Nakaya et al. (1984) preliminarily ber. The K/Ar age of the hominoid fossil-bearing horizon of the reported several rhinocerotid fossils from the Namurungule Lower Member is estimated to be 9.57 Æ 0.22 Ma and 9.47 Æ 0.22 Ma Formation, such as Brachypotherium sp. and Rhinocerotidae gen. (Sawada et al., 1998, 2006). In the paleomagnetic stratigraphy, the et sp. indet. Nakaya et al. (1987) reported additional specimens, Upper and Lower Members are correlated with Chron C4Ar.2n which were preliminarily identified as Chilotheridium sp., Para- (9.64 to 9.58 Ma) and Chrons C4Ar.2r to C4Ar.1n (9.58 to 9.31 Ma), diceros sp., K. bishopi, and Rhinocerotidae gen. et sp. indet. Later, respectively (Sawada et al., 1998, 2006). Nakaya (1994) discussed the faunal changes among the late Nakali is located 60 km south of the Samburu Hills (Fig. 1). The Miocene terrestrial mammals in sub-Saharan East Africa. In this upper Miocene Nakali and Nasorut formations are distributed in context, Rhinocerotidae gen. et sp. indet., reported by Nakaya et al. N. Handa et al. / Geobios 50 (2017) 197–209 199 (1987) was transferred to Iranotheriinae sp. nov. (Nakaya, 1994: Abbreviations: M: upper molar; m: lower molar; P: upper p. 17). Tsujikawa (2005) described additional rhinocerotid fossils premolar; p: lower premolar; MB: Maboko, Kenya; NA: Nakali, discovered from the Namurungule Formation in 1986, 1998 and Kenya; NC: Nyakach, Kenya; SH: Samburu Hills, Kenya; MN: 1999; he reported Chilotheridium pattersoni, Paradiceros mukirii, European Neogene Mammal Zones. and Rhinocerotidae gen. et sp. indet. Fukuchi et al. (2008) noted a cheek teeth row specimen of P.sp. from the Namurungule 5. Systematic paleontology Formation as Diceros sp. There are a few reports of rhinocerotid fossils from Nakali. Aguirre and Gue´rin (1974) first reported a new Class Mammalia Linnaeus, 1758 species, K. bishopi, based on material collected in the 1960s. Since Order Perissodactyla Owen, 1848 2000, a great amount of additional mammalian fossils was Family Rhinocerotidae Owen, 1845 recovered from new fieldwork, and the presence of Diceros sp. Subfamily Rhinocerotinae Dollo, 1885 was reported but without description and illustration (Kunimatsu Tribe Elasmotheriini Bonaparte, 1845 et al., 2007; Fukuchi et al., 2008).