ARTICLE IN PRESS Journal of Human Evolution

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 

Article history: 

Received 25 June 2008 

Accepted 20 August 2009 

Keywords: 

Primate biogeography 

Out of Africa I 

Cercopithecidae 

Introduction 

abstract 

This study reports the discovery of a large cercopithecid calcaneus 

from the Early Pleistocene site of ‘Ubeidiya, Israel. The specimen, 

‘Ubeidiya (UB) 330, was found in stratum III 12 during the 

1993 excavation season and is housed in the paleontological 

collection of the Hebrew University of Jerusalem, Israel (HUJI). The 

only other recorded cercopithecid primate from the Levant during 

this period is the assemblage of Macaca sylavanus from ‘Ubeidiya 

(Tchernov and Volokita, 1986). Readily observable size differences 

between Macaca calcanei and UB 330 suggest that UB 330 represents 

a different taxon, probably Theropithecus sp. (Belmaker, 2002). 

Taxonomic identification of primate foot bones remains challenging. 

However, based on studies of modern taxa, primate calcanei 

have been shown to be morphologically distinct at the familial level 

(Langdon, 1986; Strasser, 1988), and subfamilies and genera within 

Old World monkeys can be distinguished based on linear 

measurements and multivariate analyses of the calcaneus, which 

probably relate to differences in locomotion and degree of terrestriality 

vs. arboreality (Strasser, 1992; Yirga, 2002; Youlatos, 2003). 

At least three cercopithecine genera were present in Eurasia 

during the Early Pleistocene: the Eurasian genera Macaca and Paradolichopithecus, 

and the African Theropithecus. Macaca sylvanus is 

the most common cercopithecine recovered from European Early 

E-mail address: belmaker@fas.harvard.edu 

0047-2484/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved. 

doi:10.1016/j.jhevol.2009.08.004 

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 

This study presents the discovery of a right cercopithecine calcaneus from the site of ‘Ubeidiya, Israel, 

dated to ca. 1.6 Ma. The fossil is described and statistically compared to bones of modern and fossil 

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 

genera, it is most clearly similar to calcanei of Theropithecus. However, it could also represent Paradolichopithecus, 

but this alternative is unlikely due to the morphological uniqueness of the latter taxon. 

The finding of an African taxon in the Levant suggests a circum-Mediterranean dispersal route for the 

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. 

Pleistocene contexts (Delson, 1980). Paradolichopithecus is known 

from the late Ruscinian to the middle Villafranchian in Europe and 

Asia (Delson, 1974; Szalay and Delson, 1979; Ardito and Mottura, 

1987; Delson et al., 2000; Rook and Martínez-Navarro, in press), 

and Theropithecus occurs sporadically in Eurasia in Early Pleistocene 

sediments, although it was widely distributed, ranging from the 

Iberian Peninsula in the west to the Indian sub-continent in the east 

(Delson, 1993; Jablonski, 1993; Gibert et al., 1995; Delson et al., 

2000; Rook et al., 2004). In the Levant, the absence of large cercopithecids, 

often found sympatric with Macaca in other Eurasian 

sites (Ardito and Mottura,1987), may be attributed to sampling bias. 

The goal of this study is to test the hypothesis that the specimen 

UB 330 cannot be attributed to Macaca sylvanus, and to evaluate the 

alternative hypotheses that UB 330 represents one of the large 

cercopithecine genera present in Eurasia during the Early Pleistocene, 

and, specifically, a species in the genus Theropithecus. The 

mammalian fauna at the site of ‘Ubeidiya includes several African 

taxa such as Pelorovis oldwayensis and Kolpochoerus olduvaiensis 

(Geraads, 1986). The presence of the African genus Theropithecus in 

‘Ubeidiya would serve to further confirm an African-Asian dispersal 

route along the Levantine corridor (Tchernov, 1981) and shed light 

on possible hominin dispersal routes during this time period. 

Geological context 

The ‘Ubeidiya Formation lies about 3 km south of the Sea of 

Galilee in Israel, on the flanks of the western escarpment of the 

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

2 


Jordan Rift (Fig. 1). The archaeological layers of the ‘Ubeidiya 

Formation have been systematically excavated since 1960 (Stekelis, 

1966a,b; Stekelis et al., 1969; Bar-Yosef and Goren-Inbar, 1993) 

through the late 1990s (Stekelis,1966a; Stekelis et al.,1969; Bar-Yosef 

and Goren-Inbar, 1993; Guérin et al., 1996,2003; Shea and Bar-Yosef, 

1998), and are known for rich faunal (Haas, 1966,1968; Tchernov, 

1986; Belmaker, 2006) and lithic assemblages (Bar-Yosef and Goren- 

Inbar, 1993; Shea and Bar-Yosef, 1998). The primate assemblage 

includes dental and postcranial material of Macaca sylvanus (Tchernov 

and Volokita,1986) as well as a small sample of Homo cf. ergaster/ 

erectus dental material (Tobias, 1966a,b; Belmaker et al., 2002). 

Estimated dates for the fossil-bearing strata of the ‘Ubeidiya 

Formation are between ca. 1.6–1.2 Ma. Paleomagnetic analysis of 

the ‘Ubeidiya Formation indicate that it overlies the ‘Erq el Ahmar 

Formation, which is dated at 1.96–1.78 Ma (Ron and Levi, 2001) and 

has a reversed polarity, suggesting that it predates the Brunhes– 

Matuyama reversal (Opdyke et al., 1983; Braun et al., 1991; Verosub 

and Tchernov, 1991). Two short, normal paleomagnetic episodes 

M. Belmaker / Journal of Human Evolution xxx (2009) 1–11 

Figure 1. Location of the site of ‘Ubeidiya in the Southern Levant. 

have been found in strata II 33 and II 23–24 in the Fi member and 

have been assigned to the Cobb Mt. (1.215–1.190 Ma) and the Gilsa 

(1.575–1.567 Ma), respectively (Sagi, 2005). The dating of these 

short polarity events is corroborated by local faunal turnovers. The 

‘Ubeidiya fauna can be assigned to a local mammalian fauna biozone 

older than that in the sites of Bitzat Ruhama, Evron, and 

Latamne dated to ca. 1.0–1.2 Ma, suggesting that the ‘Ubeidiya 

normal polarity events in strata II 23-24 and II 33 should both 

predate the Jaramillo (0.99–1.07 Ma). (For a detailed stratigraphic 

correlation, see Supplementary Online Material Figure S1) (Belmaker, 

2009). Furthermore, the large mammalian assemblage of 

‘Ubeidiya is similar to the Farneta faunal unit (the sites of Selvella 

and Pietrafitta, Italy) (Belmaker, 2006; Martínez-Navarro et al., 

2009), which has been dated to ca. 1.6–1.2 Ma (Caloi and Palombo, 

1997, and references therein), and the lithic assemblage is similar to 

those from East African sites (Stekelis et al., 1969; Bar-Yosef and 

Goren-Inbar, 1993) such as Olduvai Upper Bed II, dated to ca. 1.53– 

1.27 Ma (Gowlett, 1979; Cerling and Hay, 1986). 


j.jhevol.2009.08.004

The total accumulation between the two normal episodes is ca. 

30 m. The micromorphological analysis of the paleolake ‘Ubeidiya 

delta system included periods of hiatus, probably in the range of 

several thousands of years during which pedogenic processes 

occurred, suggesting that the estimated duration of 400 k.yr. is not 

inconsistent with the geomorphology of the site (Mallol, 2006). 

The specimen UB 330 described here was found in stratum III 12 

in the Li member. It is stratigraphically below the Fi member that 

contains the short, normal polarity events, indicating that it most 

probably predates the Gilsa (ca. 1.575 Ma). Thus, the estimated date 

for stratum III 12 and specimen UB 330 is ca.1.6–1.58 Ma (Figure S2). 

Materials and methods 


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 fossil specimen UB 330 (Fig. 2) was compared to calcanei of 

adult extant and fossil Cercopithecidae. Calcanei were measured 

from specimens of extant Cercopithecidae species from the American 

Museum of Natural History, the Museum of Comparative 

Zoology, as well as one specimen from the personal collection of 

Philip Rightmire. Comparative fossil material included an unpublished 

Macaca sylvanus from ‘Ubeidiya (UB 101), as well as two casts 

of Theropithecus oswaldi (from Kanjera, Kenya), and a cast of Paracolobus 

chemeroni (Chemeron Fm., Kenya) generously provided by 

E. Delson. All extant comparative specimens (n ¼ 146) were adult, 

based on tooth eruption and fusion of limb elements (Table 1). 

Measurements were taken with digital calipers on both modern 

and fossil specimens to an accuracy of two decimal places (Fig. 3). 

Measurements followed Langdon (1986) and Yirga (2002). Analysis 

was performed on size-adjusted data. Two methods of size 

adjustments were used. First, raw variables were transformed to 

size-adjusted Mosimann shape data (Mosimann, 1970; Falsetti 

et al., 1993; Jungers et al., 1995). Each of the 14 raw measurements 

1 

The geometric mean is calculated as the nth root of the product of n 

measurements. 

M. Belmaker / Journal of Human Evolution xxx (2009) 1–11 3 

was divided by the geometric mean 1 , producing 14 size-adjusted 

ratio variables (designated by the variable name followed by the 

subscript ‘‘/GM,’’ e.g., Cal 1/GM). Second, 16 ratios were calculated 

following Langdon (1986), Strasser (1992), and Yirga (2002). Thus, 

a total of 30 variables (14 Mosimann shape-adjusted and 16 ratios) 

were used in this analysis (Tables S1 and S2). 

Size-adjusted values of UB 330 were compared to modern cercopithecid 

generic means using the single observation means t-test 

(Sokal and Rohlf, 1995). Multiple comparisons, such as the one 

performed here, require adjusting the probability values for the 

number of simultaneous tests to avoid Type I errors. To increase the 

power of the test, the sequential Bonferroni method was applied 

(Rice, 1989). A P value of 0.00033 was set as the test criterion of the 

single sample t-test. 

Discriminant Function Analysis (DFA) uses correlation metrics to 

address weight combinations of variables and emphasizes between 

group variation while minimizing within group variation. In this 

study, a two-tier stepwise Linear Discriminant Function was 

applied to the size-adjusted variables using stepwise insertion of 

variables (maximizing the smallest F ratio) with UB 330 treated as 

a separate group (Sokal and Rohlf, 1995). First, a DFA was run at the 

subfamilial level to test the hypothesis that UB 330 could be 

identified as a member of Colobinae or Cercopithecinae. Second, 

a DFA was run at the generic level confined to cercopithecines 

(Cercopithecus, Macaca, Mandrillus, Papio, and Theropithecus). 

(Methodological considerations of the DFA are presented in the 

Supplementary Online Material.) Leave-one-out cross validation 

was used to assess the overall error rate for the DFA. Furthermore, 

bias and standard error around the predicted posterior probabilities 

for UB 330 were estimated using the jackknife procedure. This 

was performed by running the DFA while randomly removing 

a single observation at a time and iterated for the total number of 

specimens (n ¼ 150). 

Linear regressions of cercopithecid indices of calcaneus pedal 

power arm (Cal 20) and calcaneal load arm (Cal 21) with body mass, 

have shown significant correlations at the 0.001 significance level 


j.jhevol.2009.08.004

4 

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 


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 


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. 


j.jhevol.2009.08.004

Table 2 

Pooled within-group correlations between discriminating variables and standardized 

canonical discriminant functions*. 

A. Subfamilial DFA 

Function 1 

Cal 23 0.565 

Cal 25 0.565 

Cal 4/GM 

0.340 

Eigenvalues 0.506 

% of variance 100 

Canonical Correlation 0.50 

P value

6 

Table 3 

DFA classification results. 


A. Actual rows by predicted columns for subfamilial DFA classification results: counts and (%), DFA correctly classified 89.5% of original grouped cases. 

Cercopithecinae Colobinae Total 

Cercopithecinae 120 (89.6) 14 (10.4) 134 

Colobinae 2 (11.1) 16 (88.9) 18 

B. Actual rows by predicted columns for leave-one-out cross validation for subfamilial DFA classification results: counts and (%), DFA correctly classified 

ca. 89% of cross-validated grouped cases. 

Cercopithecinae Colobinae Total 

Cercopithecinae 119 (88.8) 15 (11.2) 134 

Colobinae 2 (11.1) 16 (88.9) 18 

C. Actual rows by predicted columns for cercopithecine genera DFA classification results: counts and (%), DFA correctly classified 84.1% of original grouped cases. 

Cercopithecus Macaca Mandrillus Papio Theropithecus Total 

Cercopithecus 21 (87.5) 3 (12.5) 0 0 0 24 

Macaca 6 (12) 63 (84) 2 (2.7) 0 1 (1.3) 75 

Mandrillus 0 1 (10) 7 (70) 2 (20) 0 10 

Papio 0 0 0 16 (88.9) 2 (11.1) 18 

Theropithecus 0 1 (20) 0 0 4 (80) 5 

D. Actual rows by predicted columns for leave-one-out cross validation for cercopithecine genera DFA classification results: counts and (%), DFA correctly 

classified ca. 75% of original grouped cases. 

Cercopithecus Macaca Mandrillus Papio Theropithecus Total 

Cercopithecus 19 (79.2) 5 (20.8) 0 0 0 24 

Macaca 11 (14.7) 59 (78.7) 2 (2.7) 0 3 (4.0) 75 

Mandrillus 1 (20) 1 (10) 5(50) 2 (20) 0 10 

Papio 0 0 1 (5.6) 14 (77.8) 3 (16.8) 18 

Theropithecus 0 1 (20) 0 1 (20) 3 (60) 5 

efficiency may be an additional contributing factor in long limb 

skeletal differences between the arboreal M. fascicularis and the 

terrestrial M. nemestrina (Rodman, 1979). While he did not consider 

the osteological morphology of the foot in his original study, it is 

intriguing to speculate if this hypothesis can be applied to the 

similarities observed here between Theropithecus and Macaca. 

Theropithecus forage over much shorter distances while feeding 

than the sympatric anubis baboons (Iwamoto, 1993), and perhaps 

provide an analogy to the M. fascicularis–M. nemestrina study by 

Rodman (1979). If this is correct, the misclassification between 

Theropithecus and Macaca fascicularis may be due to similarities in 

foraging efficiencies, which may confound the distinction in 

calcaneal morphology observed among arboreal and terrestrial 

cercopithecines. 

Posterior probabilities of classification to each taxon were 

obtained for each quartile. The results indicate that UB 330 should 

be assigned to the genus Theropithecus with a median probability of 

98.5% (with an inter-quartile range of 98.1–99.2%). Results indicated 

that UB 330 can be assigned to Cercopithecus with a median 

probability of 1.1% (with an inter-quartile range of 0.06–1.67%), and 

could be assigned to Papio with a median probability of 0.22% (with 

an inter-quartile range of 0.01–0.35%). Probability of assignment to 

Macaca and Mandrillus was less than 0.001 percent. The high 

posterior probability in assignment of UB 330 to Theropithecus with 

very narrow inter-quartile ranges provides strong support for the 

identification of UB 330 as Theropithecus. 

In order to understand the morphological differences that may 

be driving the distinction between the genera, a scatter plot of the 

two first functions can be observed (Fig. 4) and analyzed in relation 

to the results of the stepwise linear DFA (Table 2B). 

The DFA plots indicate that the major separation along the first 

function, which explains 68.4% of the variance, is between the 

larger and terrestrial cercopithecine genera (Papio, Mandrillus, and 

Theropithecus), which score positive values on the first function, 

and the smaller and more arboreal genera (Macaca and 


Cercopithecus), which score negative values on the first function. 

While UB 330 scores clearly in the large terrestrial cercopithecine 

group, it occupies a unique position on the plot, specifically in 

relation to Theropithecus, and will be discussed later. 

The first function is affected primarily by Cal 20 (pedal power 

arm). Arboreal taxa such as Macaca have a shorter pedal power arm 

compared to more terrestrial taxa, since the ‘‘high gear’’ ratio 

contributes to increasing take off velocity required for leaping 

locomotion in comparison to a ‘‘lower gear’’ ratio, which is found in 

Figure 4. Bivariate plot of the first two axes of the Discriminant Function Analysis 

(DFA) separating the five cercopithecine genera and UB 330. The unique position of UB 

330 is discussed in the text. Discriminant function 1 explains 68.4% of the total 

variance, and discriminant function 2 explains 21.1% of the total variance. Theropithecus 

gelada and T. oswaldi were grouped together in the analysis. 


j.jhevol.2009.08.004


the more terrestrial locomotion of baboons (Strasser, 1992). 

However, pedal power arm is also positively correlated with body 

mass (Langdon, 1986; Strasser, 1992). Thus, we cannot exclude 

a body mass component in the distinction between groups of cercopithecine 

genera along the first function. The average body 

weight calculated for Macaca and Cercopithecus is ca. 5.5 kg 

compared to the average body weight of 21.3 kg for Papio, Mandrillus, 

and Theropithecus. 

The distinction along the second dimension, which accounts for 

only 21.1% of the variance, is more difficult to explain. In the 

smaller-bodied cercopithecine group, there is a gradient from 

negative to positive scores, from Macaca (a more terrestrial genus) 

to Cercopithecus (a more arboreal genus on average). In the larger 

taxa, a reverse trend is observed from negative to positive scores, 

from Mandrillus (which is the most arboreal), to Papio, and then 

Theropithecus, which is the most terrestrial. This similarity in 

positive scores between Cercopithecus and Theropithecus is 

surprising since both have different behavioral positions. Most of 

the Cercopithecus species sampled in this study are arboreal, and 

the two that score the highest on the second function of the DFA are 

C. mitis and C. ascanius, while Theropithecus is an extremely 

terrestrial species with a ‘‘shuffling forward bipedally’’ type of 

locomotion (Wrangham, 1980). 

The variable that most affected the second dimension is Cal 22, 

anterior height to anterior length of the calcaneus. An increase in 

height relative to length provides additional strength and robusticity 

to the calcaneus and suggests an adaptation to stresses in the 

sagittal plane produced by m. triceps surae (Langdon, 1986; Youlatos, 

2003). A well-developed m. triceps surae provides the necessary 

force required for terrestrial locomotion, as opposed to the more 

slender m. triceps surae observed in more arboreal taxa (Strasser, 

1988; Youlatos, 2003). Thus, it is not surprising that we observe the 

trend in the smaller-bodied cercopithecines. The more terrestrial 

Macaca score negatively (i.e., have higher anterior calcanei relative 

to length) while the more arboreal Cercopithecus have more positive 

scores (i.e., have shorter anterior calcanei relative to length). 

However, the results of the larger-bodied cercopithecines are more 

difficult to explain as it may have been expected that Theropithecus 

would score negatively on the second function. However, unlike 

other terrestrial cercopithecines, Theropithecus invert their feet 

much of the time while feeding, so a more agile and flexible ankle is 

advantageous, and perhaps convergent in some of its morphology 

to more arboreal forms (Krentz, 1993). Theropithecus shares with 

the arboreal Colobus an angulated medial malleolus and 

a pronounced notch for m. tibialis posterior, which aids in inverting 

the foot and increased flexibility of the ankle (Krentz, 1993). 

It is interesting to note that the score for UB 330 on the second 

function falls above any of the observed values for the comparative 

samples. This is supported by the fact that the two Theropithecus 

oswaldi from Kanjera score the highest on the second dimension 

within the Theropithecus sample (T. oswaldi mean ¼ 2.53, 

S.D. ¼ 1.279; T. gelada mean ¼ 0.84, S.D. ¼ 0.645). However, the two 

species do not differ significantly along the second dimension (twotailed 

student t-test P value >0.1), probably due to the low sample 

size of the comparative Theropithecus sample, making it difficult to 

evaluate the significance of the extreme value of UB 330. While 

probabilistically UB 330 is more similar to Theropithecus than other 

genera, the unique position of UB 330 along the second function 

may indicate either an undocumented high variability in 

Theropithecus calcaneal morphology reflective of variable locomotion 

patterns (Elton, 2002), or an otherwise unknown calcaneus of 

a fossil taxon, such as Paradolichopithecus. However, current 

assessments of the positional behavior of the latter taxon point to 

an increased terrestrial locomotion (Sondaar et al., 2006; E. Delson 

and W. Harcourt-Smith, pers. comm.), whereas T. oswaldi has been 


shown to have occupied a more arboreal habitat than modern 

T. gelada (Elton, 2002), and ‘‘shuffling forward bipedally’’ is an early 

locomotor adaptation in the Theropithecus lineage (Krentz, 1993), 

both of which support the former hypothesis. 

Body mass estimates 

The body mass for UB 330 was 25.56 kg based on the pedal 

power arm regression equation and 22.83 kg based on the calcaneal 

load arm regression equation (Strasser, 1992). Since DFA results 

indicated that UB 330 could be classified as a cercopithecine, the 

body size of UB 330 was compared to fossil and extant cercopithecines 

only. The body size estimates for UB 330 are higher than 

modern Cercopithecus and Macaca (both sexes). It is also higher 

than modern Mandrillus, Papio, and Theropithecus females but 

within the range of the males of these genera. It is only slightly 

above the range of modern Theropithecus males (Delson et al., 

2000). This value is higher than Plio-Pleistocene fossil populations 

of Macaca in general (10–16 kg for males and 6.5–12.5 kg for 

females) and the ‘Ubeidiya Macaca sylvanus in particular 

(6.5–9.5 kg for females) (Delson et al., 2000). The body mass estimate 

is within the estimated range for T. oswaldi oswaldi, dated 

between 2.5–1.2 Ma, and obtained from both dental and postcranial 

measurements (13–36 kg for females and 20–86 kg for males) 

(Delson et al., 2000), and is consistent with the identification of UB 

330 as Theropithecus (Table 4). However, it is also just within the 

estimated body size range for Paradolichopithecus avernensis 

females (12–23 kg) and males (25–41 kg) (Delson et al., 2000). 

Discussion 

The identification of primate fossil material to genus based on 

postcranial material alone is a difficult task. This study indicates 

that cercopithecid calcanei can be used to distinguish between 

subfamily and genera using both univariate and multivariate 

methods. The measurements of specimen UB 330 may have been 

a slight underestimation of the full adult size due to its adolescent 

age. However, the estimated adult length was probably only an 

additional 1–2 mm due to incomplete ossification of the epiphysis. 

Primate calcanei attain their adult shape by adolescence. Since 

Macaca sylvanus are smaller than other terrestrial Cercopithecinae, 

the comparisons with adult Macaca provide a conservative size 

comparison with UB 330. 

Based on the results in this study, UB 330 can be identified as 

a cercopithecine with a very high level of confidence. DFA analysis 

identified UB 330 as a cercopithecine with a posterior probability 

of nearly 90%. Of the cercopithecine genera analyzed, UB 330 

differed significantly from Cercopithecus, Mandrillus, Papio, and 

Macaca. One sample t-test differed in one variable or more from 

each of these genera, and stepwise linear DFA classification results 

indicated that UB 330 could be classified as Cercopithecus, 

Mandrillus, or Macaca with a probability of less than 0.0001. 

Moreover, UB 330 fell above the estimated size range for 

Cercopithecus and Macaca as well as Mandrillus females. The 

distinction from Cercopithecus and Mandrillus is not surprising as 

their current and fossil biogeographic distribution is confined to 

sub-Sahara Africa (Pickford, 1987). 

The distinction from Macaca is of particular importance. Macaca 

has been previously found in ‘Ubeidiya (Tchernov and Volokita, 

1986). Given the difference in size between UB 330 and other 

Macaca specimens at the site, specifically the Macaca calcaneus UB 

101, the null hypothesis was that UB 330 represents a large male 

Macaca sylvanus. The DFA provides the most significant distinction 

along the first function, which separates the smaller-bodied taxa 

(Macaca and Cercopithecus) from the larger and more terrestrial 


j.jhevol.2009.08.004

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? 


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? 


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 


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 


j.jhevol.2009.08.004

10 

Chupasko, and Eileen Westwig for access to collections. I am 

grateful to Philip Rightmire, Eric Delson, W.E.H. Harcourt-Smith, 

and Tina Warinner for useful comments during the process of 

identification of the specimen, and for suggestions during the 

preparation of this manuscript, and to Sarah Elton and two anonymous 

reviewers for suggestions on earlier drafts of the 

manuscripts. 

Supplementary data 

Supplementary data associated with this article can be found in 

online version, at doi:10.1016/j.jhevol.2009.08.004. 

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