ARTICLE IN PRESS Journal of Human Evolution

More documents

Recommendations

Info

8 Table 4 Body mass (kg) (fossil estimates) for modern and fossil Cercopithecinae. Species Sex Body mass (kg) (fossil estimates) Known age range Modern genera Cercopithecus _ 1.8–8.0 Extant \ 1.8–5 Macaca _ 4.9–17.5 Extant \ 3.05–14.1 Mandrillus _ 27–45 Extant \ 10–17 Papio _ 15–37.2 Extant \ 8.8–20.5 Theropithecus _ 16.5–20.25 Extant \ 9–13.8 Fossil species Paradolichopithecus avernensis _ 25–41 2.5–1.6 Ma \ 12–23 Theropithecus oswaldi oswaldi a _ 20–86 2.5–1.2 Ma \ 13–36 Macaca sylvanus (fossil) _ 10–17 Late Miocene – Early Pleistocene \ 6.5–12.5 Macaca sylvanus (from ‘Ubeidiya) \ 6.5–9.5 1.6–1.2 Ma ones. Misclassifications of the DFA were very low between these two groups. Therefore, the null hypothesis was rejected with a high degree of probability, and UB 330 was assigned to a large Cercopithecinae previously unidentified in ‘Ubeidiya. All previous Pleistocene cercopithecid material in the Levant has been attributed to the small-bodied Macaca sylvanus, and UB 330 represents the finding of a new taxon in the Early Pleistocene of the Southern Levant. Assignment to genera within the large-bodied Cercopithecinae, Papio, Paradolichopithecus, orTheropithecus, is more difficult based on the current data set but some taxa are more probable, based on morphology, body size, and biogeography. Do the data support an assignment of UB 330 to the genus Papio? ARTICLE IN PRESS The question of the identification of UB 330 is very interesting from a biogeographic point of view. Papio hamadryas hamadryas is the only Papio to disperse beyond the African continent. It is currently found in the south of the Arabian Peninsula in the republic of Yemen and Saudi Arabia (Harrison and Bates,1991). Two alternative (although not mutually exclusive) routes have been suggested for the dispersal of Papio from Africa to Arabia: 1) a longer route, which includes a dispersal northward though the Nile valley and Sinai Peninsula into the Levant and then southward to the Arabian Peninsula, and 2) crossing the Bab el Mandeb strait during periods of low sea levels (Kummer, 1995). The finding of Papio in the Early Pleistocene Levant would provide support for the former northern route. Evidence presented in this study does not provide strong support for the assignment of UB 330 to the genus Papio. While UB 330 falls within the body size range of both modern and fossil Papio species (Delson et al., 2000), DFA could assign UB 330 to Papio with a median probability of
the Levant. Of more interest would be the implication for the role of African vs. Eurasian taxa in Early Pleistocene dispersal events. Rook et al. (2004) proposed that the Early Pleistocene dispersal of Homo from Africa to Eurasia occurred in parallel with a suite of four other African taxa that included Megantereon whitei, Kolpochoerus olduvaiensis, Hippopotamus antiquus, and Theropithecus oswaldi. The presence of an ‘‘African assemblage’’ in Eurasia has been suggested as a faunal marker for the presence of Homo and has been ascribed to global climatic change (Martínez-Navarro and Palmqvist, 1995; Martínez-Navarro, 2004; Rook et al., 2004). This was based, among other things, on the presence of Theropithecus in Cueva Victoria (Gibert et al., 1995), Pirro Nord (Rook et al., 2004; Rook and Martínez Navaro, in press), Mirzapur (Gupta and Sahni, 1981), and ‘Ubeidiya (Belmaker, 2002). However, if future analysis of specimens assigned to Theropithecus were to support a reassignment to the Eurasian Paradolichopithecus, as has been suggested for Pirro Nord (Patel et al., 2007), the dispersal hypothesis would require reevaluation. Do the data support an assignment of UB 330 to the genus Theropithecus? ARTICLE IN PRESS Of the genera studied here, UB 330 is most similar to the modern genus Theropithecus. This is the only cercopithecine taxon that does not differ from UB 330 in any size-adjusted measurement. DFA classification results indicate that the probability that UB 330 could be classified as Theropithecus is 98.5%. The body size estimates are consistent with those for T. oswaldi females dating between 2.5–1.2 Ma, and it has a high positive score along the first discriminant function, which is consistent with a high value of pedal power arm for large-bodied and terrestrial cercopithecines, as well as a high value for the ratio of anterior calcaneus height to length, which is consistent with ankle inversion and flexibility. Within the genus Theropithecus, UB 330 cannot be identified to species level. However, T. oswaldi is the only Theropithecus species to have been found in Eurasia during the time frame of ‘Ubeidiya (1.6–1.2 Ma) and is therefore the most probable species identification of UB 330. If this is correct, UB 330 represents the only occurrence of Theropithecus sp. in the eastern Mediterranean Levant and one of the oldest members of its genus out of Africa. The presence of Theropithecus in Eurasia during the Early Pleistocene was sporadic although widely distributed, ranging from the Iberian Peninsula in the west to the Indian sub-continent in the east (Jablonski, 1993). T. oswaldi has been found at the site of Cueva Victoria, Spain (Gibert et al., 1995; Martínez-Navarro et al., 2005), dated to ca. 1.2 Ma; at the site of Mirzapur, India, dated to ca. 1.0–0.1 Ma (Gupta and Sahni, 1981; Delson, 1993; Pickford,1993); and at the sites of Ternifine and Thomas Quarry, Algeria (Delson, 1993), ranging in date from 1.0– 0.4 Ma (Alemseged and Geraads, 1998; Raynal et al., 2001). The species has also been identified at the site of Pirro Nord, Italy, (Rook et al., 2004; Rook and Martínez-Navarro, in press) dated to ca. 1.6– 1.3 Ma, but this identification has recently been questioned (Patel et al., 2007). While Theropithecus fossils are rare in Eurasia, their finds document the dispersal routes of large-bodied primates from Africa into Eurasia during the Early Pleistocene and mirror the possible dispersal routes used by early Homo. To date, the presence of Theropithecus in Cueva Victoria in the west and in India in the east have suggested two parallel, although not mutually exclusive, dispersal routes: a westerly dispersal route via the Gibraltar straits, and a northern dispersal route along the Nile valley and Levantine corridor (Tchernov, 1988; Petraglia, 2003; Turner and O’Regan, 2007; O’Regan, 2008). An additional route across the Bab el Mandeb strait is not probable as there is no evidence for a land bridge at M. Belmaker / Journal of Human Evolution xxx (2009) 1–11 9 the Bab el Mandeb straits during this time period (Derricourt, 2006; Fernandes et al., 2006; Turner and O’Regan, 2007). The presence of Theropithecus in ‘Ubeidiya suggests a possible circum Mediterranean dispersal route via ‘Ubeidiya (ca. 1.6 Ma), Pirro Nord (ca. 1.6–1.3 Ma), and Cueva Victoria (ca. 1.2 Ma), which may have allowed for the dispersal of Theropithecus from Africa without requiring Theropithecus (and other taxa including Homo)to cross open bodies of water such as the Gibraltar straits. This route has been shown to be the most probable based on computer simulation of vegetation and climate models (Holmes, 2007), as well as biogeographic models (O’Regan, 2008). Computer simulation models for the dispersal of Theropithecus’ such as ‘‘Stepping Out,’’’ did not include the presence of this taxon in the Levantine corridor (Hughes et al., 2008); it would be of interest to rerun additional computer simulation programs with the inclusion of an ‘Ubeidiya Theropithecus to test the probability of such routes. The high proportion of African taxa in the mammalian faunal assemblage of ‘Ubeidiya, such as Pelorovis oldwayensis, Oryx sp., and Kolpochoerus olduvaiensis has suggested the presence of a well established dispersal route between East Africa and the Central Jordan Valley (Haas, 1966; Tchernov, 1986; Martínez-Navarro, 2004). This route would have supported the dispersal of early hominin taxa as well. The presence of an additional African taxon (Theropithecus sp.) in the Central Jordan Valley provides additional support for this dispersal route during the Early Pleistocene. Conclusions Cercopithecid calcaneal morphology can be used to distinguish genera based on body size and degree of terrestrial vs. arboreal locomotion. A new specimen, UB 330, a right calcaneus from stratum III 12, ‘Ubeidiya, Israel, which has been dated to ca. 1.6 Ma, can be attributed to a large-bodied cercopithecine and represents a new primate taxon previously unidentified in the Early Pleistocene of the Southern Levant. At the genus level, it can be attributed to Theropithecus sp. with the highest probability, and represents the only member of its genus in the Southern Levant and perhaps the earliest in Eurasia. It is assigned to Theropithecus based on the high value of its pedal power arm (indicative of terrestrial locomotion), large body size, and high value of the ratio of anterior height to length consistent with the inversion and ankle flexibility observed in modern Theropithecus. While UB 330 could potentially also be attributed to Paradolichopithecus, this alternative is less probable. The presence of an African taxon in the Central Jordan Valley at this date suggests a circum Mediterranean dispersal route for Theropithecus during the earlier part of the Pleistocene and supports the presence of the Levantine corridor as a biogeographic route between Africa and Eurasia. The finding of Theropithecus sp. in ‘Ubeidiya expands our knowledge of primate dispersals during the Early Pleistocene. The understanding and interpretation of the biogeography of large-bodied primates and the dispersal route of other ‘‘Out of Africa I’’ taxa is important in elucidating hominin dispersal patterns. Acknowledgments The research was made possible by generous grants from the Irene Levy Sala CARE Foundation, the L.S.B. Leakey Foundation, the Richard Carley Hunt Wenner-Gren postdoctoral Fellowship, and the American School of Prehistoric Research postdoctoral research fellowship at Harvard University. The photographs of UB 330 were taken by M. Barazani. I am indebted to O. Bar-Yosef, David Pilbeam, and the late E. Tchernov for their support and help throughout this research, as well as to Navot Morag and Alon Barash for their technical assistance. I would like to thank Rivka Rabinovich, Judy 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
Page 1 and 2: The presence of a large cercopithec
Page 3 and 4: The total accumulation between the
Page 5 and 6: Table 2 Pooled within-group correla
Page 7: ARTICLE IN PRESS the more terrestri
Page 11: ARTICLE IN PRESS Sagi, A., 2005. Ma

ARTICLE IN PRESS Journal of Human Evolution

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?