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The Presence of a Large Cercopithecine (Cf. Theropithecus Sp.) in the ‘Ubeidiya Formation (Early Pleistocene, Israel)

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The Presence of a Large Cercopithecine (Cf. Theropithecus Sp.) in the ‘Ubeidiya Formation (Early Pleistocene, Israel) ARTICLE IN PRESS Journal of Human Evolution xxx (2009) 1–11 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol The presence of a large cercopithecine (cf. Theropithecus sp.) in the ‘Ubeidiya formation (Early Pleistocene, Israel) Miriam Belmaker Department of Anthropology, Harvard University, 11 Divinity Ave, Cambridge MA 02138, USA article info abstract Article history: This study presents the discovery of a right cercopithecine calcaneus from the site of ‘Ubeidiya, Israel, Received 25 June 2008 dated to ca. 1.6 Ma. The fossil is described and statistically compared to bones of modern and fossil Accepted 20 August 2009 cercopithecids. The specimen can be attributed to a large-bodied cercopithecine and represents a new primate taxon previously unidentified in the Early Pleistocene of the Southern Levant. Among extant Keywords: genera, it is most clearly similar to calcanei of Theropithecus. However, it could also represent Para- Primate biogeography dolichopithecus, but this alternative is unlikely due to the morphological uniqueness of the latter taxon. Out of Africa I The finding of an African taxon in the Levant suggests a circum-Mediterranean dispersal route for the Cercopithecidae taxon out of Africa, and emphasizes the importance of the Levantine corridor as a biogeographic dispersal route between Africa and Eurasia during the Early Pleistocene. Evidence for the biogeography of large-bodied primates is essential for the understanding of the dispersal routes of ‘‘Out of Africa I’’ taxa and can help elucidate Homo dispersal patterns in the Early Pleistocene. Ó 2009 Elsevier Ltd. All rights reserved. Introduction Pleistocene contexts (Delson, 1980). Paradolichopithecus is known from the late Ruscinian to the middle Villafranchian in Europe and This study reports the discovery of a large cercopithecid calca- Asia (Delson, 1974; Szalay and Delson, 1979; Ardito and Mottura, neus from the Early Pleistocene site of ‘Ubeidiya, Israel. The spec- 1987; Delson et al., 2000; Rook and Martı´nez-Navarro, in press), imen, ‘Ubeidiya (UB) 330, was found in stratum III 12 during the and Theropithecus occurs sporadically in Eurasia in Early Pleistocene 1993 excavation season and is housed in the paleontological sediments, although it was widely distributed, ranging from the collection of the Hebrew University of Jerusalem, Israel (HUJI). The Iberian Peninsula in the west to the Indian sub-continent in the east only other recorded cercopithecid primate from the Levant during (Delson, 1993; Jablonski, 1993; Gibert et al., 1995; Delson et al., this period is the assemblage of Macaca sylavanus from ‘Ubeidiya 2000; Rook et al., 2004). In the Levant, the absence of large cerco- (Tchernov and Volokita, 1986). Readily observable size differences pithecids, often found sympatric with Macaca in other Eurasian between Macaca calcanei and UB 330 suggest that UB 330 repre- sites (Ardito and Mottura,1987), may be attributed to sampling bias. sents a different taxon, probably Theropithecus sp. (Belmaker, 2002). The goal of this study is to test the hypothesis that the specimen Taxonomic identification of primate foot bones remains chal- UB 330 cannot be attributed to Macaca sylvanus, and to evaluate the lenging. However, based on studies of modern taxa, primate calcanei alternative hypotheses that UB 330 represents one of the large have been shown to be morphologically distinct at the familial level cercopithecine genera present in Eurasia during the Early Pleisto- (Langdon, 1986; Strasser, 1988), and subfamilies and genera within cene, and, specifically, a species in the genus Theropithecus. The Old World monkeys can be distinguished based on linear mammalian fauna at the site of ‘Ubeidiya includes several African measurements and multivariate analyses of the calcaneus, which taxa such as Pelorovis oldwayensis and Kolpochoerus olduvaiensis probably relate to differences in locomotion and degree of terres- (Geraads, 1986). The presence of the African genus Theropithecus in triality vs. arboreality (Strasser, 1992; Yirga, 2002; Youlatos, 2003). ‘Ubeidiya would serve to further confirm an African-Asian dispersal At least three cercopithecine genera were present in Eurasia route along the Levantine corridor (Tchernov, 1981) and shed light during the Early Pleistocene: the Eurasian genera Macaca and Par- on possible hominin dispersal routes during this time period. adolichopithecus, and the African Theropithecus. Macaca sylvanus is the most common cercopithecine recovered from European Early Geological context The ‘Ubeidiya Formation lies about 3 km south of the Sea of E-mail address: [email protected] Galilee in Israel, on the flanks of the western escarpment of the 0047-2484/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2009.08.004 Please cite this article in press as: Belmaker, M., The presence of a large cercopithecine (cf. Theropithecus sp.) in..., J Hum Evol (2009), doi:10.1016/ j.jhevol.2009.08.004 ARTICLE IN PRESS 2 M. Belmaker / Journal of Human Evolution xxx (2009) 1–11 Jordan Rift (Fig. 1). The archaeological layers of the ‘Ubeidiya have been found in strata II 33 and II 23–24 in the Fi member and Formation have been systematically excavated since 1960 (Stekelis, have been assigned to the Cobb Mt. (1.215–1.190 Ma) and the Gilsa 1966a,b; Stekelis et al., 1969; Bar-Yosef and Goren-Inbar, 1993) (1.575–1.567 Ma), respectively (Sagi, 2005). The dating of these through the late 1990s (Stekelis,1966a; Stekelis et al.,1969; Bar-Yosef short polarity events is corroborated by local faunal turnovers. The and Goren-Inbar, 1993; Gue´rin et al., 1996,2003; Shea and Bar-Yosef, ‘Ubeidiya fauna can be assigned to a local mammalian fauna bio- 1998), and are known for rich faunal (Haas, 1966,1968; Tchernov, zone older than that in the sites of Bitzat Ruhama, Evron, and 1986; Belmaker, 2006) and lithic assemblages (Bar-Yosef and Goren- Latamne dated to ca. 1.0–1.2 Ma, suggesting that the ‘Ubeidiya Inbar, 1993; Shea and Bar-Yosef, 1998). The primate assemblage normal polarity events in strata II 23-24 and II 33 should both includes dental and postcranial material of Macaca sylvanus (Tcher- predate the Jaramillo (0.99–1.07 Ma). (For a detailed stratigraphic nov and Volokita,1986) as well as a small sample of Homo cf. ergaster/ correlation, see Supplementary Online Material Figure S1)(Bel- erectus dental material (Tobias, 1966a,b; Belmaker et al., 2002). maker, 2009). Furthermore, the large mammalian assemblage of Estimated dates for the fossil-bearing strata of the ‘Ubeidiya ‘Ubeidiya is similar to the Farneta faunal unit (the sites of Selvella Formation are between ca. 1.6–1.2 Ma. Paleomagnetic analysis of and Pietrafitta, Italy) (Belmaker, 2006; Martı´nez-Navarro et al., the ‘Ubeidiya Formation indicate that it overlies the ‘Erq el Ahmar 2009), which has been dated to ca. 1.6–1.2 Ma (Caloi and Palombo, Formation, which is dated at 1.96–1.78 Ma (Ron and Levi, 2001) and 1997, and references therein), and the lithic assemblage is similar to has a reversed polarity, suggesting that it predates the Brunhes– those from East African sites (Stekelis et al., 1969; Bar-Yosef and Matuyama reversal (Opdyke et al., 1983; Braun et al., 1991; Verosub Goren-Inbar, 1993) such as Olduvai Upper Bed II, dated to ca. 1.53– and Tchernov, 1991). Two short, normal paleomagnetic episodes 1.27 Ma (Gowlett, 1979; Cerling and Hay, 1986). Figure 1. Location of the site of ‘Ubeidiya in the Southern Levant. Please cite this article in press as: Belmaker, M., The presence of a large cercopithecine (cf. Theropithecus sp.) in..., J Hum Evol (2009), doi:10.1016/ j.jhevol.2009.08.004 ARTICLE IN PRESS M. Belmaker / Journal of Human Evolution xxx (2009) 1–11 3 Figure 2. UB 330, a right calcaneus. A: Medial view, B: Lateral view, C: Plantar (inferior) view, D: Dorsal (superior) view. The scale bar represents 5 cm. The total accumulation between the two normal episodes is ca. was divided by the geometric mean1, producing 14 size-adjusted 30 m. The micromorphological analysis of the paleolake ‘Ubeidiya ratio variables (designated by the variable name followed by the delta system included periods of hiatus, probably in the range of subscript ‘‘/GM,’’ e.g., Cal 1/GM). Second, 16 ratios were calculated several thousands of years during which pedogenic processes following Langdon (1986), Strasser (1992), and Yirga (2002). Thus, occurred, suggesting that the estimated duration of 400 k.yr. is not a total of 30 variables (14 Mosimann shape-adjusted and 16 ratios) inconsistent with the geomorphology of the site (Mallol, 2006). were used in this analysis (Tables S1 and S2). The specimen UB 330 described here was found in stratum III 12 Size-adjusted values of UB 330 were compared to modern cer- in the Li member. It is stratigraphically below the Fi member that copithecid generic means using the single observation means t-test contains the short, normal polarity events, indicating that it most (Sokal and Rohlf, 1995). Multiple comparisons, such as the one probably predates the Gilsa (ca. 1.575 Ma). Thus, the estimated date performed here, require adjusting the probability values for the for stratum III 12 and specimen UB 330 is ca.1.6–1.58 Ma (Figure S2). number of simultaneous tests to avoid Type I errors. To increase the power of the test, the sequential Bonferroni method was applied Materials and methods (Rice, 1989). A P value of 0.00033 was set as the test criterion of the single sample t-test. The fossil specimen UB 330 (Fig. 2) was compared to calcanei of Discriminant Function Analysis (DFA) uses correlation metrics to adult extant and fossil Cercopithecidae. Calcanei were measured address weight combinations of variables and emphasizes between from specimens of extant Cercopithecidae species from the Amer- group variation while minimizing within group variation.
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