Seed-eating by red leaf monkeys (Presbytis rubicunda) in dipterocarp forest of northern borneo

Data are presented on the feeding behavior ofPresbytis rubicunda at Sepilok, north Borneo. Emphasis is given to describing the fruit-eating behavior of this small colobine monkey, which specializes in eating seeds from large, dull-colored, and fleshless fruits. The seed predation is conspicuously different from the seed dispersal effected by sympatric monogastric primates and is characteristic of colobine frugivory at other forest sites. Most seed-eating occurred during the period of maximum fruit production and fewer small-seeded, animal-dispersed fruits were eaten at other times.     

Dental Eruption Sequences in Fossil Colobines and the Evolution of Primate Life Histories

Unlike other catarrhines, colobines show early molar eruption relative to that of the anterior dentition. The pattern is variable, with Asian genera (Presbytina) showing a greater variability than the African genera (Colobina). The polarity of early relative molar eruption, as well as the degree to which it is related to phylogeny, are unclear. Schultz (1935) suggested that the trend reflects phylogeny and is primitive for catarrhines. More recently, however, researchers have proposed that life history and dietary hypotheses account for early relative molar eruption. If the colobine eruption pattern is primitive for catarrhines, it implies that cercopithecines and hominoids converged on delayed relative molar eruption. Alternatively, if the colobine condition is derived, factors such as diet and mortality patterns probably shaped colobine eruption patterns. Here we update our knowledge on eruption sequences of living colobines, and explore the evolutionary history of the colobine dental eruption pattern by examining fossil colobine taxa from Eurasia (Mesopithecus) and Africa (Kuseracolobus aramisi and Colobus sp.) and the basal cercopithecoid Victoriapithecus macinnesi. We scored specimens per Harvati (2000). The Late Miocene-Early Pliocene Mesopithecus erupts the second molar early relative to the incisors, while the Early Pliocene Kuseracolobus aramisi does not. These results demonstrate that the common colobine tendency for early molar eruption relative to the anterior dentition had appeared by the Late Miocene, and that some of the diversity observed among living colobines was already established in the Late Miocene/Early Pliocene. We discuss the implications of these results for phylogenetic, life history, and dietary hypotheses of dental development.     

The feeding ecology and activity budget of proboscis monkeys

A group of proboscis monkeys (Nasalis larvatus) consisting of an alpha‐male, six adult females, and several immatures was observed from May 2005–2006. We collected over 1,968 hr of focal data on the adult male and 1,539 hr of focal data on the six females in a forest along the Menanggul River, Sabah, Malaysia. Availability and seasonal changes in plant species consumed by the focal monkeys were determined by vegetation surveys carried out across an area of 2.15 ha along 200–500 m trails in riverine forest. A total of 188 plant species were consumed by the focal monkeys. The activity budget of members of our study group was 76.5% resting, 19.5% feeding, and 3.5% moving. Young leaves (65.9%) and fruits (25.9%) accounted for the majority of feeding time. Over 90% of fruit feeding involved the consumption of unripe fruits and in the majority of case both the fruit flesh and seeds were eaten. Although fruit eating was rare in some months, during other times of the year time fruit feeding exceeded the time devoted to young leaves. We found that monthly fruit availability was positively related to monthly fruit eating and feeding activity, and seasonal fluctuations in dietary diversity were significantly affected by fruit eating. These results suggest that fruit availability and fruit‐eating behaviors are key factors that influence the activity budget of proboscis monkeys. Earlier assumptions that colobine monkeys are obligate folivores do not apply well to proboscis monkeys and certain other colobines. Our findings may help contribute to a better understanding of the dietary adaptations and feeding ecology of Asian colobines. Am. J. Primatol. 71:478–492, 2009. © 2009 Wiley‐Liss, Inc.     

The nomenclature and taxonomy of the colobine monkeys of Java

The current terminology for the colobine monkeys of Java is incorrect. The appropriate term for the West Javan surili is Presbytis comataDesmarest,1822, and that for the Javan and Bali/Lombok lutung is Trachypithecus auratusE. Geoffroy St. Hilaire, 1812.     

Diets of two sympatric colobines in Zaire: More evidence on seed-eating in forests on poor soils

Recent studies have shown that the so- called folivorous African colobines eat a significant amount of seeds. There is disagreement as to whether seed-eating is due to the poor quality of tree foliage, due in turn to poor soils, or to the fact that seeds are a normal part of colobine diets. To test these hypotheses, we studied feeding of red colobus monkeys, Colobus badius tholloni,and black-and-white colobus monkeys, Colobus angolensis angolensis,in a tropical rain forest of Central Zaire (Salonga National Park). We conducted studies on soil properties, vegetation composition, and the availability and chemical contents of food plants simultaneously. Soils were very acid, with a high percentage of sand, very low cation exchange capacity, and very low exchangeable bases. The forest was dominated by legumes (45.6% of trees), among which the Caesalpinioideae were the best represented (85%). C. badiusfed mostly on leaves (61%) and seeds (33%), legumes making up 65% of their diet. C. angolensisfed mostly on seeds (50%) and leaves (27%); 39% of their diet came from legumes. The two species tended to select items richer in crude protein or lipids or both. Total phenolics and condensed tannins were abundant in the foliage and seeds but were poor predictors of colobine choice of food. Intersite comparisons show that colobines in Zaire ate a higher proportion of seeds than all other related species so far studied in Africa and that the Salonga forest had among the poorest soils and harbored the highest percentage of Caesalpinioideae. Our results confirm that seed- eating is more common among colobines living in areas where soils are poor. They strongly suggest that this link is mediated through forest composition, especially the abundance of legumes, and that the development of seed- eating results both from the high availability of nutrient-rich seeds and from the poor quality of mature tree foliage.     

Dental eruption sequence among colobine primates

Dental development is one aspect of growth that is linked to diet and to life history but has not been investigated among colobines since the work of Schultz [1935]. This study establishes the dental eruption sequence for several colobine species and compares it to that of other catarrhines. The mandibles and maxillae of two hundred and four juvenile colobine specimens were scored for presence or absence of permanent teeth and for stages of partial eruption. Eruption was defined as ranging between tooth emergence (any part of a tooth crown above the alveolar margin) and full occlusion, with three intermediate levels manifest between these boundaries. In African colobines, represented by C. guereza, C. angolensis and P. badius, M2 erupts before I2, and in C. angolensis it also erupts before I1. The canine is delayed, erupting after the premolars in females and after M3 in males. Asian colobines show greater diversity in eruption sequences. Nasalis shows no early eruption of the molars and is very similar to Macaca. In Trachypithecus and Pygathrix M(2) erupts before I(2). The canine in Trachypithecus is delayed, erupting after the premolars and, in some males, after M3. In Presbytis M2 erupts before both incisors; M3 erupts before C in both sexes, and often before both premolars. Although the actual timing of eruption is unknown, all colobine species examined except N. larvatus showed some degree of relatively early eruption of M2 and M3. The lack of this tendency in Nasalis sets this genus apart from all other colobines represented in this study. Dental eruption sequence is thought to reflect life history patterns. Early molar eruption in colobines was thought by Schultz (1935) to be a primitive character reflecting shorter life history. Faster growth rates found in folivorous primates have been interpreted as being related to an adaptation to folivory (Leigh 1994), and early eruption of molars may be part of this dietary specialization. The relationships between dental development and both diet and life history are investigated.     

Jaw morphology and function in living and fossil old world monkeys

This allometric investigation on a sample of 29 cercopithecine and 22 colobine taxa augments the data and implications of prior work on subfamilial variation in mandibular form and function in recent Cercopithecidae. To increase the size range encompassed by living cercopithecines and colobines, I included many of the larger-bodied fossil specimens. These analyses serve to fill a gap in our understanding of size-related changes in masticatory function and symphyseal morphology and curvature in extant and extinct Old World monkeys. Results of subfamilial scaling comparisons indicate that for a given jaw length, colobines possess significantly more robust corpora and symphyses than those of cercopithecines, especially at smaller sizes. Following from previous work, the most plausible explanation for why the subfamilies differ in relative corporeal and symphyseal dimensions is that colobine mandibles experience elevated loads and greater repetitive loading during mastication due, on average, to processing a diet of tough leaves and/or seeds. Although colobines have relatively larger symphyses, subfamilial analyses of symphyseal curvature demonstrate that they evince less symphyseal curvature vis-à-vis cercopithecines of a common size. Moreover, both subfamilies exhibit similar allometric changes in the degree of curvature, such that larger-bodied Old World monkeys have more curved symphyses than those of smaller taxa. Subfamilial scaling analyses also indicate that colobines possess a shorter M2 bite-point length relative to masseter lever-arm length, but not versus temporalis lever-arm length. Thus, as compared to cercopithecines, colobines can recruit less masseter-muscle force to produce similar bite forces during mastication. In both clades, M² bite-point length scales with positive allometry relative to masseter lever-arm length, such that larger species are less efficient at generating molar bite forces. This seems especially important due to the lack of subfamily difference in M² bite-point:temporalis lever-arm scaling (which is isometric across cercopithecids). A consideration of extinct cercopithecids indicates that many of the large-bodied papionins have more robust corpora, due perhaps to a diet which was of similar toughness to that of extant and extinct colobines. However, the biomechanical arrangements of the masseter and temporalis in all but one fossil cercopithecine and all of the fossil colobine specimens are much as predicted for a subfamilial member of its skull size. That most large-bodied papionins with tougher diets nonetheless maintain a less efficient jaw-muscle configuration may be due to stronger offsetting selection for increased relative canine size and gape.     

Mapping homology between human and black and white colobine monkey chromosomes by fluorescent in situ hybridization

We used in situ hybridization of chromosome specific DNA probes (“chromosome painting”) of all human chromosomes to establish homologies between the human and the white and black colobus (Colobus guereza 2n = 44). The 24 human paints gave 31 signals on the autosomes (haploid male chromosome set). Robertsonian translocations between chromosomes homologus to human 14 and 15, 21 and 22, form colobine chromosomes 6 and 16, respectively. Reciprocal translocations were found between human chromosomes 1 and 10, 1 and 17, as well as 3 and 19. The alternating hybridization signals between human 3 and 19 on Colobus chromosome 12 show that in this case a reciprocal translocation was followed by a pericentric inversion. The hybridization data show that in spite of the same diploid number and similar Fundamental Numbers, the black and white colobine monkey differs from Presbytis cristata, an Asian colobine, by 6 reciprocal translocations. Comparisons with the hybridization patterns in other primates show that some Asian colobines have a more derived karyotype with respect to African colobines, macaques, great apes, and humans. Chromosome painting also clearly shows that similarities in diploid number and chromosome morphology both between colobines and gibbons are due to convergence. Am. J. Primatol. 42:289–298, 1997. © 1997 Wiley-Liss, Inc.     

Underbite and the scaling of facial dimensions in colobine monkeys.

This investigation was concerned with the dimensions of the facial skeleton and the incidence of underbite in colobine monkeys. Standard craniofacial dimensions and the notation of the presence or absence of underbite were recorded for skulls of 308 adult, wild-caught colobines belonging to 10 species. Underbite was noted in all species examined, ranging in frequency from 10% in Colobus polykomos to 100% in 2 species of Presbytis. Univariate statistics indicate that species with high frequencies of underbite tend to have shorter, broader maxillary dental arches, shorter faces and rostrums, and longer mandibular dental arches. A multivariate discriminant analysis supported the univariate statistics. Interspecific allometric analysis of facial measurements against a constructed size variable indicated that maxillary dimensions scale in a strongly positive manner in relation to size. However, mandibular dimensions tend to scale isometrically. This pattern of facial scaling indicates that the tendency toward underbite should increase as overall size decreases. This would explain why smaller species have higher frequencies of underbite than larger species and why females often have a higher incidence of underbite than conspecific males. Zingeser has suggested that the high incidence of underbite in colobine and some cebid monkeys is an adaptation to folivory. The results of this study support an alternative hypothesis that the high incidence of underbite in colobine monkeys is related to the pattern of facial scaling with size in combination with relaxed selection on the anterior dentition.     

Fallback Foods of Red Leaf Monkeys (Presbytis rubicunda) in Danum Valley, Borneo

Animals in Southeast Asia must cope with long periods of fruit scarcity of unpredictable duration between irregular mast fruiting events. Long-term data are necessary to examine the effect of mast fruiting on diet, and particularly on the selection of fallback foods during periods of fruit scarcity. No such data is available for colobine monkeys, which may consume substantial amounts of fruits and seeds when available. We studied the diet of red leaf monkeys (Presbytis rubicunda, Colobinae) in Danum Valley, Sabah, northern Borneo, using 25 mo of behavioral observation, phenology and vegetation surveys, and chemical analysis to compare leaves eaten with nonfood leaves. The monkeys spent 46% of their feeding time on young leaves, 38% on seeds, 12% on whole fruits, 2.0% on flowers, 1.0% on bark, and 1.2% on pith. They spent more time feeding on seeds and whole fruit when fruit availability was high and fed on young leaves of Spatholobus macropterus (liana, Leguminosae) as fallback foods. This species was by far the most important food, constituting 27.9% of the total feeding time, and the feeding time on this species negatively correlated with fruit availability. Consumed leaves contained more protein than nonconsumed leaves, and variation in time spent feeding on different leaves was explained by their abundance. These results suggest that red leaf monkeys show essentially the same response to the supra-annual increase in fruit availability as sympatric monogastric primates, increasing their seed and whole-fruit consumption. However, they depended more on young leaves, in particular Spatholobus macropterus, as fallback foods during fruit-scarce periods than did gibbons or orangutans. Their selection of fallback food appeared to be due to both nutrition and abundance.     

Craniofacial variation and dietary adaptations of African colobines

African colobine monkeys show considerable craniofacial variation among species, although the evolutionary causes of this diversity are unclear. In light of growing evidence that diet varies considerably among colobine species, we investigated whether colobine craniofacial morphology varies as a function of their diet. We compared craniofacial morphology among five African species: Colobus angolensis, C. guereza, C. polykomos, Piliocolobus badius, and P. verus. Matrix correlation analysis indicated a significant correlation between species-specific morphological distance and dietary distance matrices. The mechanical advantage of the masseter muscle was higher in seed-eaters (C. angolensis and C. polykomos) and lower in those that eat mainly young leaves (C. guereza, P. badius, and P. verus). Canonical correspondence analysis revealed that the durophagous colobines possess relatively wider bigonial breadths, anteroposteriorly shorter faces, shorter postcanine tooth rows, more medially positioned dental batteries, wider bizygomatic arches, and anteroposteriorly longer zygomatic arches. Under the constrained lever model, these morphological features suggest that durophagous colobines have the capacity to generate relatively greater maximum bite forces. However, no consistent relationship was observed between diet and variation in the mandibular corpus and symphysis, implying that robust mandibles are not necessarily adaptations for stress resistance. Factors that may influence mandibular robusticity include allometry of symphyseal curvature and canine tooth support. Finally, linear measures of mandibular robusticity may suffer from error.     

Positional Behavior of Delacour’s Langurs (Trachypithecus delacouri) in Northern Vietnam

Information on positional behavior can help elucidate relationships between a species’ morphology, behavior, and environment. Delacour’s langurs (Trachypithecus delacouri) are similar to other colobines in body mass and intermembral index, yet inhabit a limestone karst environment. From August 2007 to July 2008, we collected 372 h of positional behavior and substrate use data from 8 groups of Trachypithecus delacouri in Van Long Nature Reserve, Vietnam to address questions about how the distinct —and potentially dangerous— features of karst influence colobine positional behavior. Results show that Trachypithecus delacouri is predominantly quadrupedal (66%). However, they exhibit differences that set them apart from many other colobines. Nearly 80% of locomotor and postural behaviors were performed on rocks. Leaping was remarkably infrequent, representing only 6% of overall locomotion. They leapt 3 times more frequently on trees (13%) than on rocks (4%) and more frequently used trees as a landing substrate than rocks (38% vs. 23%), both significant differences. We argue that rock (and cliff) travel is altogether different from terrestrial and arboreal travel and propose using the term petrous to indicate the substrate and incorporate the implications of its precise sharpness and verticality that lead to the complexities and risk of locomotion on rock surfaces. Trachypithecus delacouri does not show specific adaptations for limestone karst. Instead they appear to be a behaviorally flexible species and, owing to the generalized locomotor capabilities that characterize cercopithecids, capable of locomoting through and living in the limestone rock environment to which they have relatively recently been restricted.     

The morphology and wear of the lingual notch in macaques and langurs

Although interspecific differences in the relative development of molar occlusal features have been documented for most primate species, wear-related changes in these features are only beginning to be understood. In the present study, the notch between the metaconid and entoconid of M₂ was examined in a skeletal sample of Macaca fascicularis, Presbytis cristatus, and Presbytis rubicunda from the Museum of Comparative Zoology at Harvard University. Photogrammetric analyses yielded X, Y, and Z coordinates that were used to compute three angular measurements and two length measurements at the lingual notch. Statistical analyses of measurements from unworn dentitions indicated that the mesial slope of the entoconid is steeper than the distal slope of the metaconid in all three species. In addition, interspecific comparisons demonstrated that P. cristatus has the steepest lingual notch bordered by the largest molar shearing crests, while P. rubicunda and M. fascicularis have molar shearing crests of similar size, with P. rubicunda merely having higher cusps that are closer together than those of M. fascicularis. Fanally, P. cristatus shows a smaller decrease in lingual cusp relief with wear than M. fascicularis, while P. cristatus and P. rubicunda probably show similar decreases in lingual cusp relief with wear. The differences in wear-related changes in lingual cusp relief between M. fascicularis and P. cristatus may well be related to the frugivorous/folivorous dietary differences between these species.     

Patterns of reproduction in Malayan silvered leaf monkeys at the Bronx Zoo

Within phylogenetic limits reproductive characteristics of a given species may vary between populations in response to ecological and social factors. For instance, in environments where high quality nutrition is readily available, the onset and speed of reproduction are often accelerated. Other influencing factors might be maternal experience or the sex of the infant. Here we present data on reproductive characteristics for the silvered leaf monkey (Trachypithecus cristatus), a medium‐sized Asian colobine housed at the Wildlife Conservation Society's Bronx Zoo as a one‐male group. To place the species into an appropriate phylogenetic context, we limited our comparison to other colobine species. Demographic data span 21.4 years (October 1985 to March 2007) and derive from 30 adult females (128.0 female years). Detailed behavioral data stem from a 2.2 years study (November 2002 to January 2005; 734 days, 4,225 hr). As in other Asian colobines, receptive periods were short (mean=4.3 days, n=68). This is expected for one‐male groups where receptivity likely indicates, rather than conceals, ovulation. Gestation length was estimated based on a change in the pattern of sexual behavior (mean=194.6 days, n=7). It fell within the range reported for the taxon. Births occurred year round, at an early age (mean=2.9 years, n=8), at short intervals (mean=14.9 months, n=59) in combination with a short lactation (mean 12.1 months, n=9) likely due to the nearly unlimited availability of nutrition in this zoo setting. Primiparous females tended to have a longer first interbirth interval but infant survival rates were similar to multipara possibly due to the absence of predators. Maternal investment was independent of the infant's sex and birth sex ratio was even. Our results emphasize that when interpreted with caution, zoo populations yield realistic reproductive characteristics that can help fill the gap in our knowledge about colobine life history. Am. J. Primatol. 71:852–859, 2009. © 2009 Wiley‐Liss, Inc.     

Dietary and dental adaptations in the Pitheciinae.

Since Mivart (1865), Cacajao, Chiropotes, and Pithecia have been grouped into a single taxon, which he called the subfamily Pitheciinae but which I, following Rosenberger (this issue), refer to as the living members of the tribe Pitheciini. While few today doubt the association of these three living genera, not all would place them together with Aotus and Callicebus in the subfamily Pitheciinae. This is an attempt to sort out the behavioral and morphological features of feeding and dental morphology in these taxa. Extant members of the tribe Pitheciini are adapted for sclerocarpic foraging, morphological evidence for which is found in the fossils of Soriacebus and Cebupithecia. Sclerocarpic foraging in living pitheciins is a two-stage process of seed predation involving 1) specialized features of the anterior dentition that allow removal of a hard pericarp that protects a seed or seeds, followed by 2) mastication by the posterior dentition having low cusp relief to triturate nutritious seeds of a relatively soft and uniformly pliable consistency. The dentitions of fossil pitheciins, Soriacebus and Cebupithecia, demonstrate that the hypertrophy of lower incisors plus the robustness and flaring of the canine precede development of low cusp relief on molars and premolars in the evolution of morphological features associated with sclerocarpic foraging. Features of sclerocarpic foraging are found less uniformly in the other two pitheciines, Callicebus and Aotus.     

Mitochondrial cytochrome b gene sequences of old world monkeys: With special reference on evolution of Asian colobines

The classification and phylogenetic relationships of the Old World monkeys are still controversial. For Asian colobines, from three to nine genera were recognized by different primatologists. In the present study, we have sequenced a 424 bp mitochondrial tRNA^Thr gene and cytochrome b gene fragment fromMacaca mulatta, Mandrillus sphinx, Mandrillus leucophaeus, Semnopithecus entellus, Trachypithecus vetulus, T. johnii, T. phayrei, T. francoisi, Pygathrix nemaeus, Rhinopithecus roxellanae, R. bieti, R. avunculus, Nasalis larvatus, andColobus polykomos in order to gain independent information on the classification and phylogenetic relationships of those species. Phylogenetic trees were constructed with parsimony analysis by weighting transversions 5 or 10 fold greater than transitions. Our results support the following conclusions: (1) the Old World monkeys are divided into two subfamilies; (2) that among the colobines,Colobus, the African group, diverged first, andNasalis andRhinopithecus form a sister clade toPygathrix; (3) that there are two clades within leaf monkeys, i.e. 1)S. entellus, T. johnii, andT. vetulus, and 2)T. phayrei andT. francoisi; (4) thatRhinopithecus avunculus, R. roxellanae, andR. bieti are closely related to each other, and they should be placed into the same subgenus; (5) thatRhinopithecus is a distinct genus; and (6) that the ancestors of Asian colobines migrated from Africa to Asia during the late Pliocene or early Pleistocene.     

Changes in Baboon Feeding Behavior: Maturity-dependent Fruit and Seed Size Selection within a Food Plant Species

Despite considerable inter- and intraindividual variation in fruit and seed size in many plant species, researchers have given little attention to the relevance of the traits for primate fruit choice within a food plant species and its implications for tree regeneration. We studied feeding behavior and selectivity of olive baboons (Papio anubis) in the African locust bean (Parkia biglobosa, Mimosaceae), via direct observations of habituated groups and indirect evidence from leftovers of pods after feeding events. Olive baboons acted as both seed predators and dispersers for Parkia biglobosa. They fed on and destroyed unripe seeds, and swallowed intact ripe seeds when consuming mature fruit pulp. Predation rate was high, and only 10% of the seeds were dispersed. Predation and dispersal of seeds is linked to seed number and size. Digestible unripe seeds accounted for 10% of the unripe fruit mass, while indigestible ripe seeds made up 28% of the mature fruit mass. With these constraints, olive baboons increased food gain per fruit by selecting unripe pods containing a high number of large and heavy seeds. Consequently, only pods with fewer and smaller seeds remained for maturation. Thereafter, baboons fed on mature pods containing the smallest seeds, and exploited pods with more seeds to a greater extent than those with fewer seeds. Thus, fruits with small seeds and an intermediate seed number contributed the most to dispersal by baboons.     

The history and chronology of siwalik cercopithecids

Cercopithecid monkeys make a comparatively late appearance in the Miocene Siwalik formations of the Indian Subcontinent. The oldest well-dated specimen is a 6.3 MY old colobine, ?Presbytis sivalensis, and it is doubtful that any cercopithecids were present in the Siwalik faunas prior to 7.0 MY, a date that is considerably later than estimates for their appearance elsewhere in Eurasia. Cercopithecines appear later than colobines. Although their temporal ranges are uncertain, cercopithecines evidently were not present before 3.2 MY and possibly not until 2.5 MY. They are represented by three species,?Macaca palaeindica, Procynocephalus subhimalayanus, andTheropithecus delsoni. In the Siwaliks the appearances of colobines and cercopithecines coincide with or follow shortly after major faunal changes that also bring other northern Eurasian and African taxa into the region.     

Systematic classification of Asian colobines

In order to study the differentiation of Asian colobines, fourteen variables were analysed in one way, on 123 skulls, includingRhinopithecus, Presbytis, Presbytiscus, Pygathrix, andNasalis with both cluster and differentiated functions tests. Information on paleoenvironment changes in China and South-East Asia since late Tertiary have been used to examine the influences of migratory habits and the distribution range in Asian colobines. The cladogram among different Asian colobines genera was made from the results of various analysis. Some new points or revisions were suggested: 1. Following the second migratory way, ancient species of Asian colobines perhaps passed through Xizang along the northern bank of Tethis sea and Heng-Duan Shan regions, across Yunnan into Vietnam, since the ancient continent between Yunnan and Xizang was already located in on eastern bank of Tethis sea. Thus, during the evolution, Asian colobines must have had two original centres, i.e. “Sundaland” and Heng-Duan Shan Chinese regions; 2. Pygatrix possesses a lot of cranial features more similar toPresbytiscus than toRhinopithecus. The small difference from the modification combinesPygatrix with other two genera as shown by Groves (1970), but it is better to putPygatrix andPresbytiscus together as one genus; 3.Nasalis (2n=48) may be the most primitive genus within Asian colobines. Some features shared withRhinopithecus, for example body size, terrestrial activities, limb proportion etc. ...seem to be considered as a common inheritance of symlesiomorphus characters; 4.Rhinopithecus, with reference to cranioface and cranium or to its origin, is a special genus of Asian colobine. It may represent the highest level of evolutionary position within various genera (Peng et al., 1985).