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The presence of a large cercopithecine (cf. Theropithecus sp.) in the ‘Ubeidiya
formation (Early Pleistocene, Israel)
Department of Anthropology, Harvard University, 11 Divinity Ave, Cambridge MA 02138, USA
Received 25 June 2008
Accepted 20 August 2009
Out of Africa I
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 Para-
dolichopithecus, 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.
Ó2009 Elsevier Ltd. All rights reserved.
This study reports the discovery of a large cercopithecid calca-
neus from the Early Pleistocene site of ‘Ubeidiya, Israel. The spec-
imen, ‘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 repre-
sents a different taxon, probably Theropithecus sp. (Belmaker, 2002).
Taxonomic identiﬁcation of primate foot bones remains chal-
lenging. 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 terres-
triality vs. arboreality (Strasser, 1992; Yirga, 20 02; Youlatos, 2003).
At least three cercopithecine genera were present in Eurasia
during the Early Pleistocene: the Eurasian genera Macaca and Par-
adolichopithecus, and the African Theropithecus. Macaca sylvanus is
the most common cercopithecine recovered from European Early
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 Martı
´nez-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 cerco-
pithecids, 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 Pleisto-
cene, 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 furtherconﬁrm an African-Asian dispersal
route along the Levantine corridor (Tchernov, 1981) and shed light
on possible hominin dispersal routes during this time period.
The ‘Ubeidiya Formation lies about 3 km south of the Sea of
Galilee in Israel, on the ﬂanks of the western escarpment of the
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Journal of Human Evolution
journal homepage: www.elsevier.com/locate/jhevol
0047-2484/$ – see front matter Ó2009 Elsevier Ltd. All rights reserved.
Journal of Human Evolution 58 (2010) 79–89
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; Gue
´rin 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 (Tcher-
nov and Volokita,1986) as well as a small sample ofHomo 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
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 bio-
zone 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)(Bel-
maker, 2009). Furthermore, the large mammalian assemblage of
‘Ubeidiya is similar to the Farneta faunal unit (the sites of Selvella
and Pietraﬁtta, Italy) (Belmaker, 2006; Martı
´nez-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).
Figure 1. Location of the site of ‘Ubeidiya in the Southern Levant.
M. Belmaker / Journal of Human Evolution 58 (2010) 79–8980
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
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 Amer-
ican 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 unpub-
lished Macaca sylvanus from ‘Ubeidiya (UB 101), as well as two casts
of Theropithecus oswaldi (from Kanjera, Kenya), and a cast of Para-
colobus chemeroni (Chemeron Fm., Kenya) generously provided by
E. Delson. All extant comparative specimens (n¼146) 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
was divided by the geometric mean
, producing 14 size-adjusted
ratio variables (designated by the variable name followed by the
subscript ‘‘/GM,’’ e.g., Cal 1/
). 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 cer-
copithecid 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 Pvalue 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 probabili-
ties 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
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
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 geometric mean is calculated as the nth root of the product of n
M. Belmaker / Journal of Human Evolution 58 (2010) 79–89 81
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.,
All analyses were performed using SPSS (version 16.0) statistical
software. The jackknife DFA procedure was calculated using R.
Taxonomic comparisons and statistical results
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 Gal-
liari (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 morpho-
type is adapted to an increased ability in grasping and the supi-
nation 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 ever-
sion, 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).
Comparative species measured by sex.
Family Genus Species \_U Total
Colobinae Colobus Colobus guereza 221 5
Colobus polykomos 106 7
Colobus sp. 0 0 1 1
Procolobus Procolobus badius 211 4
Paracolobus Paracolobus chemeroni 001 1
Cercopithecinae Cercopithecus ascanius 130 4
Cercopithecus diana 012 3
Cercopithecus erythrotis 001 1
Cercopithecus hamlyni 011 2
Cercopithecus mitis 371 11
Cercopithecus mona 020 2
Cercopithecus nictitans 001 1
Macaca Macaca arctoides 100 1
Macaca assamensis 021 3
Macaca fascicularis 18 19 0 37
Macaca fuscata 030 3
Macaca maura 010 1
Macaca mulatta 030 3
Macaca nemestrina 76013
Macaca nigra 001 1
Macaca ochreata 100 1
Macaca sp. 0 1 0 1
Macaca sylvanus 202 4
Macaca thibetana 010 1
Macaca tonkeana 150 6
Mandrillus Mandrillus leucophaeus 010 1
Mandrillus sphinx 450 9
Papio Papio hamadryas
Theropithecus Theropithecus gelada 120 3
Theropithecus oswaldi 002 2
Papio hamadryas includes individuals assigned to ﬁve subspecies: P. h. ursinus, P. h. hamadryas, P. h. anubis, P. h. cynocephalus, and P. h. papio.
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 measure-
ments correspond to the descriptions of Cal 1–14 in Table S1.
M. Belmaker / Journal of Human Evolution 58 (2010) 79–8982
The specimen UB 330 has proportions and morphology similar
to that of Cercopithecidae in general, and terrestrial Cercopitheci-
nae in particular. The anterior articular facet and posterior articu-
lation for the talus possess several derived features typical of
cercopithecines: the anterior articular facet is fused into a single
facet and is not divided into two sub-facets; the posterior articu-
lation for the talus is well deﬁned, quadrilateral, medially deviated
relative to the long axis of the bone, and strongly arched; the
calcaneal tuberosity process is dorso–ventrally thick with
a rounded dorsal margin indicative of awell-developed insertion of
m. triceps surae; the posterior talar facet breadth is wide and short,
indicative of reduced movement along the calcaneo-astragalar
facet; and a weak peroneal process, which is a characteristic of
terrestrial Cercopithecinae as compared to the pronounced process
in arboreal or semi-arboreal taxa (Harrison, 1989).
One sample t-test
Table S3 presents the descriptive statistics (means and standard
deviations) for the raw measurements for the comparative sample
and UB 330, and Table S4 presents the descriptive statistics and the
results for the one sample t-test. Despite overall similarities in the
majority of measurements, the one sample t-test indicates that UB
330 differs from each of the modern genera by at least one size-
adjusted measurement, with the exception of Theropithecus and
Procolobus, which do not differ from UB 330 in any measurement.
However, both have the lowest sample sizes (5 and 4, respectively),
which may have affected the results of the t-test.
Subfamilial stepwise linear DFA
Three variables met the criteria for the stepwise linear DFA
criterion (in order of descending absolute size of correlation within
function): Cal 23, Cal 25, and Cal 4/
were used in this analysis
(Table 2A). Calcaneus morphology correctly distinguished between
Colobinae and Cercopithecinae in 89.5% of the specimens, ranging
from 88.9–89.6% for the two subfamilies. Leave-one-out cross
validation results indicate a robust classiﬁcation, as nearly identical
classiﬁcation rates were obtained with 89% of specimens correctly
classiﬁed. Misclassiﬁcation rates of each of the subfamilies were
nearly identical with 10.4% of the cercopithecines identiﬁed as
colobines and 11.1% of the colobines identiﬁed as cercopithecines.
Cross validation results were similar indicating a highly robust
classiﬁcation (Table 3).
Posterior probabilities of classiﬁcation to each subfamily were
obtained for each quartile, and indicated that UB 330 should be
assigned to the subfamily Cercopithecinae with a median proba-
bility of 97.6% (with an inter-quartile range of 97.2–98.2%), and
could be assigned to Colobinae with a median probability of 2.3%
(with an inter-quartile range of 1.79–2.79%).
The DFA produced a single eigenvector that explained 100%
of the variance (Table 2A). The three variables, Cal 23 (the index of
breadth of cuboid facet to distal length), Cal 25 (the index
of breadth to height of calcaneal tuberosity), and Cal 4/
talar facet breadth), are consistent with observed morphological
differences between cercopithecines and colobines. An increase in
the size of the cuboid facet relative to the size of the calcaneus has
been shown to secure the distal calcaneal joint (Langdon, 1986).
Similarly, a decrease in height-to-width ratio for the insertion of m.
triceps surae (Cal 25) and an increase in length of the posterior talar
facet (Cal 4/
) are indicative of a more ﬂexible distal calcaneal
joint and calcaneal-astragalar joint. These are adaptive for leaping,
bounding, and galloping performed during arboreal locomotion in
comparison to terrestrial locomotion (Langdon, 1986; Strasser,
These results are consistent with the qualitative morphological
observations that support an identiﬁcation of UB 330 as a terrestrial
Cercopithecine genera stepwise linear DFA
The variables which met the stepwise selection criterion (in
order of descending absolute size of correlation within function)
were: Cal 20, Cal 22, Cal 19, Cal 28, Cal 27, Cal 18, Cal 9/
, Cal 3/
Cal 25, Cal 17, and Cal 23. Stepwise linear discriminate function
analysis on size-adjusted ratios calculated four discriminate func-
tions. The ﬁrst function accounted for roughly 68.4% of the variance,
the remaining functions accounting for 22.1%, 7.3%, and 2.1%, in
descending order (Table 2B). Cercopithecine genera differed in
calcaneus morphology (Wilk’s
<0.001), and all variables signiﬁcantly affected genus differences.
The average correct classiﬁcation rate for cercopithecine genera
was 84.1%, ranging from 70–89% per genus (Table 3). Leave-one-out
cross validation rate indicates an overall correct classiﬁcation rate
of 75% with a range of 50–79.2%. The genera were ordered roughly
according to their level of arboreality, from Cercopithecus as the
most arboreal, and following in descending order to the most
terrestrial: Macaca,Mandrillus,Papio, and Theropithecus. Genera
tend to misclassify into genera that were most similar, or only ‘‘one
step over,’’ in locomotion pattern. However, there were two
exceptions. In addition to Cercopithecus and Mandrillus,Macaca is
misclassiﬁed (albeit in very low proportions) as the extremely
terrestrial Theropithecus. Similarly, Theropithecus is only mis-
classiﬁed as Macaca in the original reclassiﬁcation and misclassiﬁed
as both Macaca and Papio in the leave-one-out cross validation
reclassiﬁcation. This result appears contradictory to the expectation
derived from the behavioral position of the two genera, and
suggests that some other behavioral similarities exist between the
two genera. A closer examination reveals that the three Macaca
specimens that were misclassiﬁed as Theropithecus are M. fas-
cicularis. It has been hypothesized by Rodman (1979) that foraging
Pooled within-group correlations between discriminating variables and standard-
ized canonical discriminant functions*.
A. Subfamilial DFA
Cal 23 0.565
Cal 25 0.565
% of variance 100
Canonical Correlation 0.50
B. Cercopithecine genus DFA
Function 1 2 3 4
Cal 20 0.834 0.248 0.064 0.043
Cal 22 0.092 0.552 0.020 0.020
Cal 19 0.120 0.467 0.135 0.010
Cal 28 0.085 0.393 0.296 0.134
Cal 27 0.186 0.292 0.092 0.124
Cal 18 0.126 0.094 0.515 0.335
0.200 0.343 0.514 0.213
0.092 0.044 0.466 0.308
Cal 17 0.179 0.293 0.304 0.509
Cal 23 0.359 0.320 0.069 0.450
Eigenvalues 2.616 0.846 0.281 0.079
% of variance 68.4 22.1 7.3 2.1
Canonical Correlation 0.851 0.677 0.468 0.270
Pvalue <0.001 <0.001 0.001 0.217
*Variables ordered by absolute size of correlation within function.
M. Belmaker / Journal of Human Evolution 58 (2010) 79–89 83
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
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 indi-
cated 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
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
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.
M. Belmaker / Journal of Human Evolution 58 (2010) 79–8984
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 cer-
copithecine 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,Man-
drillus, 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 robus-
ticity to the calcaneus and suggests an adaptation to stresses in the
sagittal plane produced by m. triceps surae (Langdon, 1986; Youla-
tos, 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 posi-
tive 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
apronounced 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 ¼2.53,
S.D. ¼1.279; T. gelada mean ¼0.84, S.D. ¼0.645). However, the two
species do not differ signiﬁcantly along the second dimension (two-
tailed student t-test Pvalue >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 locomo-
tion 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 cercopi-
thecines 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 esti-
mate 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).
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,orMacaca 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
M. Belmaker / Journal of Human Evolution 58 (2010) 79–89 85
ones. Misclassiﬁcations of the DFA were very low between these
two groups. Therefore, the null hypothesis was rejectedwith a high
degree of probability, and UB 330 was assigned to a large Cerco-
pithecinae previously unidentiﬁed in ‘Ubeidiya. All previous Pleis-
tocene 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
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 <1%, and it differed signiﬁcantly
from Papio in the variable Cal 4/
. Therefore, the probability that
UB 330 may be assigned to Papio is very low. This is concordant
with analyses of the levels of genetic variation in P. h. hamadryas in
Southern Arabia compared to the population in the Horn of Africa,
which suggests a colonization date of ca. 200–400 ka (Wildman
et al., 2004; Winney et al., 2004). This coincides with the formation
of a land bridge in the Bab el Mandeb straits at 18 ka, 130 ka,
270 ka, 370 ka, and 440 ka (Rohling et al., 1998; Siddall et al.,
2003), which postdate the Early Pleistocene dispersal of African
species from East Africa into Eurasia ca. 1.8–1.6 Ma, which is
apparent in ‘Ubeidiya.
Do the data support an assignment of UB 330 to the genus
Comparison between UB 330 and fossil calcanei of Para-
dolichopithecus could not be done, because no Paradolichopithecus
calcanei are yet known; however, the estimated body size range
obtained for UB 330 falls within the estimated range for Para-
dolichopithecus males and females. Therefore, the possibility that
UB 330 may be attributed to this Eurasian taxon cannot be
excluded. Nonetheless, based on current information, this identi-
ﬁcation would appear less probable. A talus (PO 157F) from the site
of Vatera on the island of Lesvos, Greece, assigned to P. arvernensis
and dated to MN17 (St. Vallier faunal Unit) ca. 2.4–1.8 Ma, allows for
an indirect comparison of foot bone morphology. The morphology
of the talus is said to be unique among cercopithecine tali in that
the facet for the tibial malleolus in plantar view is ﬂattened rather
than bowl shaped; furthermore, the trochlea is only slightly
wedge–shaped, and the dorsal aspect of the ﬁbular articulation
protrudes laterally (Sondaar and Van der Geer, 2002; Van der Geer
and Sondaar, 2002; Sondaar et al., 2006). This would suggest that
UB 330, with its typical cercopithecine calcaneal morphology and
similarity to Theropithecus calcanei, could not articulate with the
unique talus of Paradolichopithecus. It is worth noting that this
morphological interpretation, while intriguing, is highly contro-
versial and a comparison by Delson and colleagues of the distal
tibia and talus of the Vatera specimen indicates no signiﬁcant
distinction between the postcranial elements of PO 157F and other
large papionins (E. Delson and W. Harcourt-Smith, pers. comm.),
suggesting that such conclusions may be premature. Nonetheless,
since the DFA results presented here suggest that cercopithecine
calcanei can be used to identify genera, and given the high posterior
probability obtained in this study of the assignment of UB 330 to
Theropithecus, while the possibility that UB 330 is a Para-
dolichopithecus cannot be excluded, it would seem less probable.
However, if future fossils were to conﬁrm that UB 330 is indeed
Paradolichopithecus, its recovery in ‘Ubeidiya would provide
a geographic expansion of this taxon westward and southward into
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
Macaca _4.9–17.5 Extant
Mandrillus _27–45 Extant
Papio _15–37.2 Extant
Theropithecus _16.5–20.25 Extant
Fossil species Paradolichopithecus avernensis _25–41 2.5–1.6 Ma
Theropithecus oswaldi oswaldi
_20–86 2.5–1.2 Ma
Macaca sylvanus (fossil) _10–17 Late Miocene – Early Pleistocene
Macaca sylvanus (from ‘Ubeidiya) \6.5–9.5 1.6–1.2 Ma
UB 330 ? 22.83–25.56 1.6–.2 Ma
Values for fossils are maximum estimated body mass ranges derived from cranial and postcranial material and include 20% estimated mass from Delson et al. (2000), as are
comparative extant data.
Measured specimens in this study are from Kanjera, Kenya, and dated to ca. 2 Ma.
M. Belmaker / Journal of Human Evolution 58 (2010) 79–8986
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 old-
uvaiensis,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 (Martı
´nez-Navarro and Palmqvist, 1995;
´nez-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
´nez Navaro, in press), Mirzapur (Gupta and Sahni, 1981), and
‘Ubeidiya (Belmaker, 2002). However, if future analysis of speci-
mens 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
Do the data support an assignment of UB 330 to the genus
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 discrim-
inant 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; Martı
´nez-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 Martı
´nez-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 Man-
deb 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 simula-
tion 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; Martı
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.
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 Pleisto-
cene 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
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
M. Belmaker / Journal of Human Evolution 58 (2010) 79–89 87
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 anon-
ymous reviewers for suggestions on earlier drafts of the
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