Dental Morphometric Variation Between African and
Asian Colobines, With Special Reference to the Other
Old World Monkeys
Ruliang Pan*
School of Anatomical Sciences, University of Witwatersrand, Johannesburg, Parktown,
Johannesburg 2193, South Africa
ABSTRACT In order to reveal differences between
Asian and African colobines (nonhuman primates) in
terms of dietary adaptation and evolutionary develop-
ment, a large number of the species of the cercopithecoids
(Old World monkeys) was analyzed with morphometric
methods. In addition to the raw data and ratios, devia-
tions of the colobines, predicted from allometric baselines
derived from the cercopithecines, were analyzed by uni-
variate and multivariate analyses. Some results indicated
that there exists significant differentiation between colo-
bines and cercopithecines; the latter exhibit a larger den-
tal scale relative to body size, and the less developed front
teeth of the colobines may be related to their fewer fre-
quent incisal use, compared with those of the cerco-
pithecines. With regard to variations between the two
subtribes of the colobines, which are found in African
(Colobina) and Asia (Presbytina), the Asian subtribe dis-
plays a larger scale of postcanine teeth, referring to the
results of the raw data. This may correspond to their
larger body size. While ratios were considered, the varia-
tion between Presbytina and Colobina was diminished
greatly. This implies that, unlike the scenario postulated
to reflect the relationships between colobines and
cercopithecines—which supposes that their differentia-
tion is in both size and shape—the variation between the
two subtribes is principally size associated: relative to
body size Colobina exhibits more emphasized incisors, but
less developed postcanine teeth—with the exception of
width of M3s. In other words, the relationships between
teeth and body size of the African colobines are more
similar to those of the cercopithecines. This implies, com-
pared to their Asian partners, that African colobines share
more similarities with cercopithecines. This may be re-
lated to the episodes in which African species underwent a
longer period of sharing environmental and climatic pat-
terns with the cercopithecines that moved to Asia about 5
or 8 million years ago. Canines were found to be important
in distinguishing colobines from cercopithecines and in
separating one subtribe from the other. J. Morphol. 267:
1087–1098, 2006. © 2006 Wiley-Liss, Inc.
KEY WORDS: colobines; Africa; Asia; Colobina; Presby-
tina; cercopithecines; dental morphometric variation; di-
etary adaptation; evolution
Colobines are primates now widely distributed in
Asia and Africa. The great diversities among them
in many aspects have made it difficult to understand
their biological issues clearly, and thus the relation-
ships between taxa (genera, species, and subspecies)
and their systematic position among Old World
monkeys still remain difficult to determine. This is
more impressive if the historical alternations re-
garding their classification and phylogeny are re-
viewed: they were once regarded as an independent
family, Colobidae (Ye et al., 1987; Groves, 1989); a
subfamily, Colobinae, in Cercopithecidae (Corbet
and Hill, 1987; Fleagle, 1988; Groves, 2001); and one
family that contained two subfamilies, Nasalinae
and Colobinae (Groves, 1989). They were also re-
garded as two groups, or subtribes, in a subfamily—
the Presbytina including Asian species and the Colo-
bina containing African taxa (Strasser and Delson,
1987; Delson, 1994). Thus, more studies to clarify
those controversies from different fields are ex-
tremely necessary.
Colobines now inhabit the forests and the wood-
lands of African tropical regions and southern and
eastern Asia. They are now found in habitats around
the equator—the forest gallery by the Nile River
(Colobus guereza) in Africa and the coastal swamp
forests in Asia (Nasalis larvatus) (Rowe, 1996); con-
fined to the plateaus—the Qinghai–Tibet Plateau—
adjacent to the Himalayas, where, in a habitat cov-
ered by snow all year round and with dominant
coniferous forest and lichen, one of the snub-nosed
monkey species (Rhinopithecus bieti) is found. The
niches occupied by this species reach an altitude of
4,700 m above sea level (Li et al., 1981, 2002).
Although colobines are generally regarded as fo-
livorous primates—feeding primarily on leaves,
Contract grant sponsors: Paleontology Scientific Trust; the Na-
tional Research Foundation, South Africa.
*Correspondence to: Dr. Ruliang Pan, School of Anatomical Sci-
ences, University of Witwatersrand, Johannesburg, 7 York Rd., Park-
town, Johannesburg 2193, South Africa.
E-mail: ruliang.pan@wits.ac.za
Published online 2 June 2006 in
Wiley InterScience (www.interscience.wiley.com)
DOI: 10.1002/jmor.10463
JOURNAL OF MORPHOLOGY 267:1087–1098 (2006)
© 2006 WILEY-LISS, INC.