Diet of <Emphasis Type="Italic">Pygathrix nigripes</Emphasis> in Southern Vietnam

Black-shanked douc langurs (Pygathrix nigripes) are Southeast Asian colobines about which primatologists know very little, but they are classed as endangered because of population decline due to habitat loss. Two preliminary studies have shown that this monkey is primarily folivorous, but there are few details of plant selection or seasonal changes in diet. We set out to observe douc langurs directly in the wild to quantify the diet during wet and dry seasons. We confirmed that the species relies on foliage throughout the year, but it includes significant proportions of fruit and flowers in its diet when those items are available. The douc langurs selected various parts from 152 species of plants in 2 national parks, but there did not appear to be favorite species that were heavily selected over others. In both the wet and dry seasons, the black-shanked douc langurs ate mostly leaves, but the diversity of plants consumed increased and the proportion of fruit eaten almost doubled in the wet season. There were diurnal shifts in food selection, but we found no evidence that the douc langurs were foraging strategically to maximize their protein or energy intake.     

Karyotype abnormalities in two primate species, Pygathrix nemaeus and Lemur coronatus.

The karyotypes of 7 douc langurs (Pygathrix nemaeus) and 3 crowned lemurs (Lemur coronatus) were examined. Abnormalities in 23.5% of the karyotypes of 1 male douc langur were associated with a history of fathering stillborns and abortuses (38%). Karyotype analysis of an apparently normal female lemur revealed three differing karyotypes, one normal and two abnormal.     

Seasonality of group size,feeding, and breeding in wild red‐shanked douc langurs (Lao PDR)

In Asian colobines, small one‐male groups (OMG) seem to predominate alongside all‐male groups (AMG), while larger multimale groups (MMG) are rare, but are reported for Hanuman langurs and red‐shanked douc langurs. Recently, however, it has been speculated that the genus Pygathrix could have multilevel societies based on (1) a theoretical extension of the multilevel societies found in Rhinopithecus to all odd‐nosed colobines and (2) first data for black‐shanked douc langurs. This assumes bands composed of small OMG with a skewed adult sex ratio. Band size may vary with seasonal food availability resulting in smaller bands when feeding competition is increased. To investigate the social organization of red‐shanked douc langurs and potential seasonal influences, we observed 2 unhabituated groups at Hin Namno National Protected Area, Lao PDR from March 2007 to August 2008 for 803 hr. We recorded births and performed group counts and scan sampling of feeding behavior. Most births (79% of N = 15) occurred from June–September, indicating a 4‐month peak conception season from November to February. Group size averaged 24.5 individuals (range 17–45) with 2.45 adult males (range 1–4). Although the smaller group remained at a stable size (about 18 individuals), the larger group reduced from about 45 to 25 individuals during the 7‐months long lean season, when less than 50% of the feeding time was spent on fruits. This suggests feeding competition as a potential cause of seasonal variation in group size. With 1.9 females per male the skew in adult sex ratio was much lower compared with Rhinopithecus, indicating MMG rather than multilevel societies. However, data on the spacing and interaction patterns between recognized individuals need to be collected and analyzed before the social organization can be determined. Detailed ecological data are furthermore required to investigate the basis for the seasonal changes in group size found. Am. J. Primatol. 73:1134–1144, 2011. © 2011 Wiley Periodicals, Inc.     

A Phylogeny of Chinese Leaf Monkeys Using Mitochondrial ND3-ND4 Gene Sequences

The phylogeny of Chinese leaf monkeys, especially the snub-nosed monkeys (Rhinopithecus), has not been thoroughly investigated using molecular sequence data, perhaps due to their rarity in the wild and their poor representation in institutional collections. Despite several proposed classifications, systematic relationships of these species remain poorly defined and this has hindered their conservation. To clarify the phylogenetic relationships of the leaf monkey clade in China, we sequenced the mitochondrial ND3, ND4L, ND4, tRNA ^Arg , tRNA ^His , tRNA ^Ser , and tRNA ^Leu genes for Rhinopithecus bieti, R. roxellana, Trachypithecus francoisi, T. f. leucocephalus, and T. phayrei as well as Pygathrix nemaeus and Colobus guereza. We included a total of 2252 characters for each individual, excluding gaps in primary sequences. Our interpretation of the results from character-and distance-based phylogenetic analyses suggest that (1) Pygathrix nemaeus is sister to Rhinopithecus rather than to Trachypithecus though it is quite divergent from the former; (2) the Yunnan snub-nosed monkey, Rhinopithecus bieti, represents a valid species; (3) the white-headed leaf monkey is not a distinct species, but instead is a subspecies of Trachypithecus francoisi (T. f. leucocephalus), though it should still be considered a separate evolutionary significant unit (ESU); and (4) because two individuals of the Phayrei's leaf monkey, T. phayrei, are genetically distinct from one another, a more extensive revision of the taxonomy of this putative species in China is needed. These results, plus ongoing work on the molecular systematics of the entire Asian leaf monkey radiation, can provide a sound basis for identifying the appropriate units of conservation for this endangered group of primates.     

Ecological Niche Conservatism in Doucs (Genus Pygathrix)

The impressive diversity and unique distribution of primate taxa in Southeast Asia creates a region of interesting biogeography and evolutionary history that remains poorly understood. The three species in the Asian colobine genus Pygathrix (P. cinerea, P. nemaeus, and P. nigripes) appear to follow the unique distribution pattern, replacing one another along the north–south axis. However, the evolutionary history and taxonomic status within Pygathrix is currently debated. We constructed climate-based ecological niche models (ENMs) for the three species, using six environmental variables, to confirm that the bioclimate envelopes of Pygathrix cinerea, P. nemaeus, and P. nigripes follow the north–south gradient. We also used the ENMs to test whether the three species lack ecological exchangeability, meaning differentiation among factors that define the fundamental niche, and whether they exhibit ecological niche conservatism, or the tendency for related species to differ less ecologically than might be expected by the climate available to each species. Our ENMs for Pygathrix followed a north–south gradient as expected, with P. nemaeus extending from ca. 14°N to 21°N, P. cinerea from 14°N to 16°N, and P. nigripes from 11°N to 13.5°N. This study presents the first use of ENMs for doucs that yield significant predictive ability based only on climatic variables. The models are significantly different for all three species, supporting a lack of ecological exchangeability between them, as well as the recent elevation of Pygathrix cinerea to species status. Importantly, Pygathrix cinerea and P. nemaeus show evidence of ecological niche conservatism, which lends support to the occurrence of an allopatric speciation event. This result also suggests that the current overlapping distribution of Pygathrix cinerea and P. nemaeus is the result of secondary contact. The north–south distribution gradient, which exists in similar form among other groups of sister taxa in the region, may be attributed to a zoogeographic barrier, vicariance, or current or historical climatic shifts.     

Comparisons of Suspensory Behaviors Among <Emphasis Type="Italic">Pygathrix cinerea</Emphasis>, <Emphasis Type="Italic">P. nemaeus</Emphasis>, and <Emphasis Type="Italic">Nomascus leucogenys</Emphasis> in Cuc Phuong National Park,Vietnam

In our study at the Endangered Primate Rescue Center of Cuc Phuong National Park, Vietnam, we aimed first to assemble a positional behavioral profile of captive gray-shanked (Pygathrix cinerea) and red-shanked (P. nemaeus) doucs that relates to the use of forelimb suspensory postures and arm-swinging locomotion. The profile is of interest because researchers have documented that red-shanked doucs more frequently use suspensory postures and locomotions than other colobines do. We confirmed that red-shanked doucs commonly use suspensory positional behaviors and also that gray-shanked doucs use suspensory behaviors at similar or even higher frequencies than those of red-shanked doucs. Our second goal was to assemble a preliminary kinematic profile of suspensory locomotion in Pygathrix within the context of the arm-swinging locomotion exhibited by northern white-cheeked gibbons, Nomascus leucogenys. Mean forelimb angles at initial contact and release of arm-swinging behaviors were remarkably consistent among gibbons and doucs despite the fact that gibbons typically used more continuous brachiation. Doucs also exhibit a greater range of forelimb angles than gibbons do. In addition, trunk orientation tends to be less vertical at initial contact for doucs than for gibbons, perhaps owing to the frequent use of quadrupedal sequences directly before or after forelimb suspension. Our behavioral and kinematic analyses add to the emerging realization that Pygathrix is capable of, and frequently expresses, a range of suspensory positional behaviors, including brachiation.     

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.     

Cranial morphology of the golden monkey (Rhinopithecus) and douc langur (Pygathrix nemaeus)

Because they have been less studied than most other non-human primates (partly due to the difficulties in accessing their habitat) the origins and phylogenetic relationships ofR. roxellana andR. bieti are controversial. These controversies may be clarified to some degree by adding information on the cranium. To this end, ten cranial dimensions analysed morphometrically here provide data about cranial differences among species of the genusRhinopithecus, and between species ofRhinopithecus andPygathrix nemaeus. Though more similar to each other than to any others in the same genus, the results show a significant separation betweenR. roxellana andR. bieti to the degree that they may be regarded as two different species. This confirms the conclusions of prior studies of external features, qualitative morphological characteristics and biochemical evidence (Yeet al, 1987; Zhang and Ryder, 1995; Jablonski, 1998; Penget al., 1988). The differences between these two species are mainly size-related, being highly correlated with cranial length. Other differences, probably non-size related shape differences, however, are highly correlated with cranial width. Sexual dimorphism plays a part in these findings. In relation to the other species, however, the results show that the Vietnam golden monkey (R. avunculus) has closer craniometric relationships with the douc langur (Pygathrix nemaeus) rather than with the three Chinese golden monkey species. Of these, the Gouzhou species (R. brelichi) shares more similarity withR. avunculus and is more separate fromRhinopithecus roxellana and R. bieti. The smaller differentiation between the two latter species could be due to their more recent separation following the dramatic elevation of Qinghai-Tibetan Plateau after the Middle Pleistocene.     

Airborne Mycobacterium avium infection in a group of red‐shanked douc langurs (Pygathrix nemaeus nemaeus)

Background This report describes an airborne Mycobacterium avium (MA)‐infection in two red‐shanked douc langurs (Pygathrix nemaeus nemaeus) from Cologne zoo. Methods The two individuals and their tissues were investigated clinically (including x‐rays), in pathology, in pathohistology, in classical bacteriology and by polymerase chain reaction (PCR). Results Clinically, one individual displayed emaciation and a positive reaction in an intrapalpebral testing for M. bovis/MA. The other individual was without any symptoms and did not show any reaction in the intrapalpebral test. In x‐ray photos of the lungs, calcified nodules were detected. In pathology, calcified and necrotic nodules were observed within the lungs and the bronchial lymph nodes. In pathohistology, both classical tuberculous granulomas, and few acid fast rods were seen in Ziehl‐Neelsen‐stain. However, classical bacteriology could not demonstrate mycobacteria. In PCR, MA‐infection could be confirmed in one individual using the bronchial lymph nodes. Conclusions It was an airborne infection; however, the definite source of infection in these cases remained unclear. Animals in contact to the langurs (house sparrows and mice) as well as water used in the building are the most promising candidates. The risk for a zoonotic transmission in these cases has been calculated to be low.     

Learning the ropes: The ontogeny of locomotion in red-shanked douc (Pygathrix nemaeus), Delacour's (Trachypithecus delacouri), and Hatinh langurs (Trachypithecus hatinhensis) I. positional behavior

Recent studies at the Endangered Primate Rescue Center (EPRC) in the Socialist Republic of Vietnam's Cuc Phuong National Park by Byron et al. ([2002] Am. J. Phys. Anthropol. [Suppl.] 34:51) and Covert and Byron ([2002] Caring for Primates) revealed unexpected locomotor and postural behaviors in the red-shanked douc langur (Pygathrix nemaeus). This paper further elucidates the question of red-shanked douc suspensory behavior, and provides initial positional behavior data for two other rare Asian colobines, by comparing the ontogenetic positional behavior of red-shanked douc langurs, Delacour's langurs (Trachypithecus delacouri), and Hatinh langurs (Trachypithecus hatinhensis) at the EPRC. Two hundred and seven hours of positional behavior data were collected, with approximately equal amounts of data on each species, and equal amounts on adults and those less than 18 months in age. All young animals were more active than adults, used a wider repertoire of locomotor behaviors, and expressed suspensory behaviors more frequently than did the adults. Young animals also "invented" one new locomotor and two new postural behaviors. These differences are due to both play and explorative behavior, as well as to the youths' changing musculoskeletal systems. The number of positional behaviors utilized by the adults of these species is quite similar to one another (23-32), as is that utilized by the young (51-56). Douc langurs in both age categories used suspensory behaviors more frequently than did Delacour's and Hatinh langurs. Because the uniformity of enclosures offers a control, the results of this study generate hypotheses regarding adaptive radiations and niche partitioning in wild populations.     

Distribution and Population Densities of Diurnal Primates in the Karst Forests of Phong Nha – Ke Bang National Park,Quang Binh Province,Central Vietnam

We studied the status and distribution of the diurnal primates in the Phong Nha – Ke Bang National Park (PNKB NP) from April to August 2007. In the past, researchers reported 9 primate species and subspecies for the karst forests of PNKB NP, which constitutes the most important protected area for the endangered Hatinh langur (Trachypithecus hatinhensis) in Vietnam. All 9 primate taxa are threatened due to hunting pressure and habitat loss. We applied line transect sampling in 4 areas. During a total of 117 transect inspections along 12 different transect routes, we confirmed 5 primate taxa and the analysis of population densities resulted in 2143 (±467) Hatinh langurs, 1316 (±871) red-shanked douc langurs (Pygathrix nemaeus), 930 (±489) stump-tailed macaques (Macaca arctoides), 986 (±883) eastern Assamese macaques (M. a. assamensis), and 18 (±18) southern white-cheeked crested gibbons (Nomascus siki) in the whole PNKB NP, which covers an area of ca. 85,000 ha. We could not detect the 2 nocturnal lorises, Bengal slow lorises (Nycticebus bengalensis) and pygmy slow lorises (N. pygmaeus), as well as rhesus macaques (Macaca mulatta) and northern pig-tail macaques (M. leonina). The distribution of the primates predominantly depended on human impact. We could not recognize a correlation between habitat constitution and abundance of primates. The population density estimates showed a much higher density of the Hatinh langur than previously assumed. Thus the importance of the PNKB NP for the conservation of this endangered langur increased significantly.     

Status and Conservation of Douc Langurs (Pygathrix nemaeus) in Laos

A recent systematic survey program of protected areas in Laos documented douc langurs (Pygathrix nemaeus) widely between 14°25N and 18°38N. Their precise northern limit is unclear as security constraints prevented survey there, but the species is unlikely to occur in Laos north of 20°. All our records are of the nominate, red-shanked form, P. n. nemaeus, though in the far south, greatly reduced red on the legs suggests intergradation with another form. Our records are overwhelmingly from evergreen or semievergreen forest and from a wide altitudinal range (200–1600 m). In most sites habitat was little degraded, but whether this reflects choice for such habitats, or the generally lower hunting pressure within them, is unclear. The largest population occurs over 3000 km ² of the Nam Theun basin and surroundings, encompassing the protected areas of Nakai-Nam Theun, Hin Namno and the proposed Nam Theun Extension. This is the most important population of red-shanked langurs in the world. Laos supports globally the vast majority of their conservable population. Douc langurs are threatened in Laos by habitat fragmentation increasing their vulnerability to the existing high hunting levels. They are a favored target for local consumption and for local and Vietnamese hunters and traders. Protective measures should center around implementing habitat conservation and antihunting measures in the recently-declared National Biodiversity Conservation Areas. A recent government initiative to reduce gun use in rural areas is likely to have a strong positive effect on the population. The best protection for some time will remain the inaccessibility of their habitat. Accordingly, road-building into remote forest and piecemeal clearance at its edges should thus be minimized. Cross-border attention should be given to wildlife trading, including measures to reduce market demand.     

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).     

Chromosome Painting Shows That Pygathrix nemaeus Has the Most Basal Karyotype Among Asian Colobinae

We mapped the chromosomal homology of Pygathrix namaeus (douc) with human and other primates by in situ hybridization of human chromosome paints. The synteny of 3 human chromosomes (1, 2, 19) is fragmented in the douc karyotype and the 23 human probes (autosomes plus X) provided 26 signals. There are associations between human chromosomes 14/15, 21/22, and 1/19. Human chromosomes 1 and 19 are divided in two segments and associated on douc chromosomes 8 and 10. The fragmentation and association of human chromosomes 1 and 19 is best explained as the result of a reciprocal translocation, which occurs in all documented Asian colobines studied, but not in the African species Colobus guereza. However, the homologs to douc chromosome 10 in all other Asian documented colobines show an additional pericentric inversion. Our results indicate that Pygathrix nemeus is karyologically the most conservative colobine species yet studied and that this species probably diverged early after the separation of Asian and African Colobinae. The data reinforce the monophyly of the Colobinae and their division into an African and an Asian clade.     

Sexual dimorphism in the snub-nosed langurs (Colobinae: Rhinopithecus)

Sexual dimorphism in the dentition and skeleton of the four extant species of snub-nosed langurs, Rhinopithecus (R.) bieti, R. (R.) brelichi, R. (R.) roxellana and R. (Presbytiscus) avunculus, was studied. The species shared a similar general pattern of sexual dimorphism, but were found to differ in respects that appear to reflect the influence of disparate socioecological and environmental factors. All the species showed marked canine dimorphism, but the very high degree of canine dimorphism in R. bieti appeared to be due to the intensity of intermale competition for mates during a temporally restricted breeding season, and possibly also to the intensity of competition between males for other resources during other times of the year. Sexual dimorphism in the postcranial skeleton of Rhinopithecus species was also most pronounced in R. bieti and may be related to the relatively higher frequency of terrestrial locomotion in males of the species. © 1995 Wiley-Liss, Inc.     

Phylogeny of Snub-Nosed Monkeys Inferred from Mitochondrial DNA,Cytochrome B,and 12S rRNA Sequences

Snub-nosed monkeys (Rhinopithecus spp.) are confined to isolated mountainous regions in China and North Vietnam. Their systematic classification and phylogenetic relationship has been controversial. The structures of mitochondrial DNA cytochrome b and 12S rRNA show that the 4 species of Rhinopithecus are quite different from other colobines. It is reasonable to regard them as an independent genus, as determined by external features, morphometric characters and behavior. However, whether or not there should be a subdivision between the Vietnamese and Chinese species at the subgeneric level remains to be clarified; more evidence from a large range of Asian colobine species is needed. The Guizhou species, Rhinopithecus brelichi, is a valid species, which is more closely related to Pygathrix than the other species ( R. roxellana, R. bieti and R. avunculus) are. Results also indicate that 3 species—Rhinopithecus roxellana, R. bieti and R. avunculus—might have diverged from R. brelichi, but the phylogenetic relationship of R. avunculus is not clear.     

It’s Tough Out There: Variation in the Toughness of Ingested Leaves and Feeding Behavior Among Four Colobinae in Vietnam

Colobines are similar in their exploitation of a high percentage of leaf matter. However, this observation obfuscates interesting differences among genera of Southeast Asian colobines in morphology and behavior that may be reflected in the degree to which they rely on mastication or gut volume and gut retention time when ingesting and digesting leaves. We detail the use of a laboratory-based method to measure the mechanical properties of foods selected and processed by 4 captive species of Southeast Asian Colobinae —Pygathrix nemaeus, Pygathrix cinerea, Trachypithecus delacouri, and Trachypithecus laotum hatinhensis— at the Endangered Primate Rescue Center (EPRC), Vietnam. We also detail a field method that quantifies chewing rates and chewing behavior via a consumer-grade video camera and laptop computer. Observations in the captive setting permit a degree of experimental control that is not possible in the wild, and the location of the EPRC in the primates’ habitat country permitted us to provide leaves that they encounter and eat in the wild. We collected toughness data with a portable tester designed by Lucas et al. The average toughness of selected leaves does not differ among the taxa, nor does the length of time spent chewing foods. However, there are differences in feeding rate, with Trachypithecus spp. chewing foods twice as fast as Pygathrix spp. Our findings suggest that Trachypithecus spp. emphasize comminution of food by mastication, while Pygathrix spp. emphasize the comminution of leaf matter in the stomach. The hypothesis is supported by data on molar size, gut mass, and gut morphology. We provide new insights into dietary variation among primate species and detail methods that are typically conducted only in a laboratory setting. We augment the findings with additional data on activity, feeding rates, and tooth morphology.     

Zoogeography of Vietnamese primates

Based on geographic complementarity of congeneric taxa, the 22 species or well-defined subspecies of primates that inhabit Vietnam may be allocated to nine zoogeographic groups (lorises, two; macaques, three; langurs, three; gibbons, one). Analysis of group distributions reveals two major patterns: (1) four of the six non-macaque groups are essentially restricted to the eastern part of the Indochinese Peninsula, east of the Mekong River; and (2) species and subspecies in seven of the nine groups reach their northern or southern limits of distribution in central Vietnam, at 14–17‡N. The first pattern suggests that the Mekong River has been an important barrier to westward dispersal of nonmacaque primates in continental Southeast Asia. The second pattern suggests that a Zoogeographic barrier formerly extended east and west across Vietnam at ca. 14–17‡N.     

Pygathrix nemaeus in Hainan? new evidence,no resolution

The douc, Pygathrix nemaeus, isnow restricted to the Indochinese Peninsula. An enigmatic report of this species in Hainan, China, is based on a specimen preserved in the Staatliches Museum für Tierkunde, Dresden. Examination of this specimen and relevant museum records reveals (a) that the specimen belongs to the subspecies P. nemaeus nemaeusand (b) that the specimen was received from the Dresden Zoological Garden, which apparently provided information that it originated in Hainan. Evaluation of the reliability of this geographic information is now impossible, because relevant zoo records no longer exist. The provenance of the problematic specimen — Hainan or Indochinese Peninsula-remains an open question.     

Sequence evolution of the CCR5 chemokine receptor gene in primates.

The chemokine receptor CCR5 can serve as a coreceptor for M-tropic HIV-1 infection and both M-tropic and T-tropic SIV infection. We sequenced the entire CCR5 gene from 10 nonhuman primates: Pongo pygmaeus, Hylobates leucogenys, Trachypithecus francoisi, Trachypithecus phayrei, Pygathrix nemaeus, Rhinopithecus roxellanae, Rhinopithecus bieti, Rhinopithecus avunculus, Macaca assamensis, and Macaca arctoides. When compared with CCR5 sequences from humans and other primates, our results demonstrate that: (1) nucleotide and amino acid sequences of CCR5 among primates are highly homologous, with variations slightly concentrated on the amino and carboxyl termini; and (2) site Asp13, which is critical for CD4-independent binding of SIV gp120 to Macaca mulatta CCR5, was also present in all other nonhuman primates tested here, suggesting that those nonhuman primate CCR5s might also bind SIV gp120 without the presence of CD4. The topologies of CCR5 gene trees constructed here conflict with the putative opinion that the snub-nosed langurs compose a monophyletic group, suggesting that the CCR5 gene may not be a good genetic marker for low-level phylogenetic analysis. The evolutionary rate of CCR5 was calculated, and our results suggest a slowdown in primates after they diverged from rodents. The synonymous mutation rate of CCR5 in primates is constant, about 1.1 x 10(-9) synonymous mutations per site per year. Comparisons of Ka and Ks suggest that the CCR5 genes have undergone negative or purifying selection. Ka/Ks ratios from cercopithecines and colobines are significantly different, implying that selective pressures have played different roles in the two lineages.