ARTICLE IN PRESS Journal of Human Evolution

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Table 1
Comparative species measured by sex.
Family Genus Species \ _ U Total
Colobinae Colobus Colobus guereza 2 2 1 5
Colobus polykomos 1 0 6 7
Colobus sp. 0 0 1 1
Procolobus Procolobus badius 2 1 1 4
Paracolobus Paracolobus chemeroni 0 0 1 1
Cercopithecinae Cercopithecus ascanius 1 3 0 4
Cercopithecus diana 0 1 2 3
Cercopithecus erythrotis 0 0 1 1
Cercopithecus hamlyni 0 1 1 2
Cercopithecus mitis 3 7 1 11
Cercopithecus mona 0 2 0 2
Cercopithecus nictitans 0 0 1 1
Macaca Macaca arctoides 1 0 0 1
Macaca assamensis 0 2 1 3
Macaca fascicularis 18 19 0 37
Macaca fuscata 0 3 0 3
Macaca maura 0 1 0 1
Macaca mulatta 0 3 0 3
Macaca nemestrina 7 6 0 13
Macaca nigra 0 0 1 1
Macaca ochreata 1 0 0 1
Macaca sp. 0 1 0 1
Macaca sylvanus 2 0 2 4
Macaca thibetana 0 1 0 1
Macaca tonkeana 1 5 0 6
Mandrillus Mandrillus leucophaeus 0 1 0 1
Mandrillus sphinx 4 5 0 9
Papio Papio hamadryas a
2 16 0 18
Theropithecus Theropithecus gelada 1 2 0 3
Theropithecus oswaldi 0 0 2 2
with R 2 above 0.9 (Strasser, 1992). Estimated body mass of UB 330,
based on calcaneus body mass regression equations developed by
Strasser (1992), were compared to Plio-Pleistocene fossil primate
body mass estimates, as retrieved from the literature (Delson et al.,
2000).
All analyses were performed using SPSS (version 16.0) statistical
software. The jackknife DFA procedure was calculated using R.
Taxonomic comparisons and statistical results
Description
ARTICLE IN PRESS
Total 150
a Papio hamadryas includes individuals assigned to five subspecies: P. h. ursinus, P. h. hamadryas, P. h. anubis, P. h. cynocephalus, and P. h. papio.
A complete right calcaneus, catalogue number ‘Ubeidiya (UB)
330, was found in stratum III 12 (Fig. 2). The calcaneal tuber, lateral
process, and lateral edge are weathered. The calcaneal tuberosity is
in advanced epiphyseal fusion corresponding to stage C in the
development of appendicular bone in primates as defined by Galliari
(1988), and the articular surfaces are well defined and angular.
Age and bone fusion correlations for Old World monkeys (Papio and
Macaca) suggest that fusion of the calcaneus begins between the
ages of 3–4 years and is completed by the age of 7 (Bramblett, 1969;
Kimura and Hamada,1990). This would suggest an ‘‘adolescent’’ age
for UB 330 based on the age class of Kawai et al. (1983) and a nearly
mature adult maximum length (Scheuer and Black, 2004).
Within the Cercopithecidae, there are two subfamilial
morphological patterns that typify adaptations for terrestrial and
arboreal locomotion. The cercopithecine morphotype is adapted to
increased stress during plantarflexion and inteversion as well as
dorsiversion and eversion. In comparison, the colobine morphotype
is adapted to an increased ability in grasping and the supination
of the forefoot (Strasser, 1988). This results in a longer
proximal calcaneal region in cercopithecines (a longer lever arm
M. Belmaker / Journal of Human Evolution xxx (2009) 1–11
and a better leverage for foot plantar flexion), a larger insertion for
a bulkier m. triceps surae, and a wider and shorter posterior talar
facet (reducing the sliding function, proximal inversion and eversion,
and helicoid movement in the joint). In comparison, the
morphology of Colobinae exhibits a shorter proximal calcaneal
region, a longer and narrower m. triceps surae insertion, and
narrow and long posterior talar facets, which serve to increase the
precision and power in the mobility of the foot during movement
across difficult terrain, such as branches, during arboreal walking
and climbing (Langdon, 1986; Strasser, 1988, 1992).
Figure 3. Linear measurements taken on UB 330 and modern comparative calcanei. A:
Anterior view, B: Medial view, C: Superior view, D: Posterior view. These measurements
correspond to the descriptions of Cal 1–14 in Table S1.
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