<Emphasis Type="Italic">Macaca munzala</Emphasis>: A New Species from Western Arunachal Pradesh,Northeastern India<Superscript>*</Superscript>

Macaca, comprising 20 well-characterized species, represents the largest and one of the most ecologically and socially diverse of all the nonhuman primate genera. We report the discovery of a macaque that is new to science from the high altitudes of western Arunachal Pradesh, a biodiversity-rich state in northeastern India. We propose the scientific name Macaca munzala and the vernacular name Arunachal macaque for the species. It shares morphological characteristics independently with the Assamese macaque (Macaca assamensis) and with the Tibetan macaque M. thibetana; like them, it appears to belong to the sinica species-group of the genus. However, the species is distinctive in relative tail length, which is intermediate between those of Tibetan and Western Assamese macaques, the subspecies with which it is sympatric. It is also unique in its altitudinal distribution, occurring largely at altitudes between 2000 and 3500 m. We provide a morphological characterization of the species, report preliminary data on its field biology and discuss possible taxonomic identity in relation to the other closely-related species of Macaca.^*This paper is dedicated to Dr. A. J. T. Johnsingh for his immense contributions to the study and conservation of India’s wildlife and for being an inspiration to a whole generation of students.     

Body Size and Proportions and Pelage Color of Free-Ranging Macaca mulatta from a Zone of Hybridization in Northeastern Thailand

Rhesus (Macaca mulatta) and long-tailed (M. fascicularis) macaques belong to the same species, and are parapatric within a zone that lies between 15° and 20° N on the Indochinese peninsula. Researchers have reported probable hybrids between the 2 species from that zone, but have not studied the extent of introgression. To test for phenotypic evidence of hybridization, we collected body mass, morphometrics (body size and proportions), and pelage color readings from free-ranging rhesus living close to the zonal boundary at Wat Tham Pa Mak Ho (WTPMH), Wang Saphun district, Loei province, northeastern Thailand (17°14′N, 101°47′E). Female WTPMH rhesus macaques (n =12) were 10–20% smaller, but with a greater relative tail length than the captive Chinese or Indian female rhesus. Female WTPMH were larger than the free-ranging long-tailed macaques, but with similar limb proportions and a shorter relative tail length. The WTPMH rhesus macaques also displayed the bipartite pelage color pattern typical of Macaca mulatta . The evidence suggests slight contribution of long-tailed macaques to the gene pool of the WTPMH population. Further sampling of other macaque populations within the zone and genetic analysis are essential to address better the question of hybridization. Determination of the distribution and range of biobehavioral variation of macaques within the zone is urgently needed, because their habitat is being rapidly destroyed by deforestation, and their demography and social structure are threatened by artificial disturbance.     

Winter ecology of the Arunachal macaque Macaca munzala in Pangchen Valley,western Arunachal Pradesh,northeastern India

The newly described Arunachal macaque Macaca munzala occurs largely in sub‐tropical to temperate environments at elevations of c. 1,800–3,000 m in Arunachal Pradesh, northeastern India. We studied its over‐wintering strategy by comparing the diet, ranging, and behavior of a troop of 24 individuals during winter and spring (December 2005 to May 2006) through instantaneous scan sampling (3,002 records, 448 scans, 112 hr of observation). We also monitored the phenology of food plants. The macaques spent more time (41–66%) feeding in the winter than in spring (33–51%), whereas time spent moving and resting was greater in spring. The diet composed largely of plants, with animal matter being eaten rarely. The number of plant species in the diet increased from 18 to 25 whereas food types rose from 18 to 36 from winter to spring, respectively. Although only two species formed 75% of the winter diet, seven species comprised this proportion in spring. Availability of fruits and young leaves increased in spring; the troop moved more and utilized a larger part of its range during this time. Seasonal changes in behavior could be explained by the scarcity of food and the costs of thermoregulation in winter. Our study suggests that the Arunachal macaque inhabits a highly seasonal environment and has an over‐wintering strategy that includes subsisting on a high‐fiber diet by increasing the time spent feeding, and minimizing energy expenditure by reducing the time spent moving. Am. J. Primatol. 71:939–947, 2009. © 2009 Wiley‐Liss, Inc.     

海南猕猴的形态学特征及其适应性

本文通过测量猕猴海南亚种(Macaca mulatta brevicaudus)的形态学指标,讨论了该亚种的形态特点及其适应性意义。于2015年7-8月在海南省陵水县猕猴自然保护区对8个群的猕猴进行了形态学指标测量,共测得253只个体的体质量,132只个体的头长、顶臀长、尾长、前脚掌长、后脚掌长、前臂长、后臂长、大腿长、小腿长等体长数据。研究结果发现:1)猕猴海南亚种的体型明显小于位于更高纬度地区的川西亚种、福建亚种和指名亚种,也小于相近纬度的大陆猕猴,结果支持贝格曼法则和岛屿法则。2)猕猴海南亚种雄性的生长速度大于雌性,且前者体型显著大于后者。3)群大小对群内个体的体型无显著影响,无显著群间竞争。4)个体的体质量、头长、顶臀长随等级增加而增加,但是尾长、臂长、腿长和脚掌长与等级无显著相关关系。本文为探讨猕猴属的形态演化提供基础资料,同时也为濒危灵长类保护和管理提供科学依据。     

Niche separation of sympatric macaques, Macaca assamensis and M. mulatta, in limestone habitats of Nonggang, China

Comparative studies of sympatric species are essential in understanding those species’ behavioral and ecological adaptations as well as the mechanisms that can reduce resource competition enough to allow coexistence. We collected data on diet, activity budget and habitat use from two sympatric macaque species, the Assamese macaque (Macaca assamensis) and the rhesus macaque (M. mulatta), in a limestone seasonal rainforest of Nonggang Nature Reserve, southwestern Guangxi, China. Our results show that the two sympatric macaques differ in diet, activity budget, and habitat use: (1) out of the 131 plant species that were used by both macaque species as food over the year, only 15 plant species (11 %) were shared. Rhesus macaques used more plant species as major foods, and had higher dietary diversity and evenness indexes than Assamese macaques. (2) Assamese macaques fed predominantly on leaves, whereas rhesus macaques fed more selectively on fruits. The rhesus macaques’ diet varied according to season, and was significantly correlated to season fluctuation in fruit availability. (3) Assamese macaques devoted more time to resting, and less time to feeding than rhesus macaques (4) Assamese macaques were present mostly on the cliff, and tended to stay on the ground, whereas rhesus macaques were present mostly on the hillside, and showed preference to lower and middle canopy. The observed differences in diet and habitat use between the two macaque species represent behavioral patterns enabling their coexistence.     

Morphological and Body Color Variation in Thai <Emphasis Type="Italic">Macaca fascicularis fascicularis</Emphasis> North and South of the Isthmus of Kra

Long-tailed macaques (Macaca fascicularis fascicularis) are widely distributed in Southeast Asia and are morphologically and genetically (Tosi et al. in International Journal of Primatology 23:161–178, 2002) distinguishable on either side of the Isthmus of Kra (ca. 10.5°N). We compared the somatometry and body color of 15 local populations of long-tailed macaques in Thailand distributed over areas from 6.5°N to 16.3°N and also a Thai rhesus macaque population at 17.2°N. Limb proportions and body color variation follow the geographical trend. However, contrary to a previous report, body size does not decrease with latitude in the northern group and also in the southern (southerly distributed) rhesus macaque. Relative tail length (RTL) and color contrast in yellow between the back and thigh are the sole traits that distinctively separate the 2 groups: the southern group has a long relative tail length (RTL >125%) and small color contrast, whereas the northern group has a short RTL (<120%) and large color contrast. The southern rhesus macaques appear to have somatometric and body color traits that follow the geographical trend in long-tailed macaques, though they maintain their distinctive species-specific traits of shorter RTL (ca. 55%), shorter relative facial length, and a bipartite body color pattern. Researchers assume that the northern group of long-tailed macaques and the southern rhesus macaques had undergone partial introgression with each other. Montane refugia present during the glacial period are localities in which introgression occurred in long-tailed macaques.     

Distribution of Assamese macaques (Macaca assamensis) in the Inner Himalayan region of Bhutan and their mtDNA diversity

The distributions of Assamese macaques (Macaca assamensis) and rhesus macaques (M. mulatta) in Bhutan have been only partially documented. In order to investigate the distribution patterns of these species, we conducted field observation and genetic assessment with mitochondrial DNA (mtDNA) typing of macaques in the Inner Himalayas of Bhutan. There were 24 sightings of macaque groups, and all were visually identified as Assamese macaques. No groups of rhesus macaques were sighted in this survey area, in contrast with the survey results in the Nepalese Himalayas. Molecular phylogenetic analysis revealed that the Bhutan macaques are closer in proximity to their counterparts in the Indo-Chinese region (Thailand and Laos) than to rhesus macaques in China, Laos and India. However, clustering results suggested the marked differentiation of the macaques in Bhutan from the Assamese macaques in Indo-China. We tentatively conclude that the macaques of the Inner Himalayan regions in Bhutan are Assamese macaques and that they appear to be of a lineage distinct from Assamese macaques in the Indo-Chinese region (subspecies M. a. assamensis). The degree of mtDNA diversity suggests that the Assamese macaques in Bhutan are of a more ancient ancestry than M. a. assamensis, thereby supporting the speciation hypothesis of the expansion of a sinica-group of macaques from South Asia to Southeast and then to East Asia (Fooden; Fieldiana Zool 45:1–44, 1988). Assignment of Assamese macaques in Bhutan to M. a. pelops is premature due to the lack of molecular data and recent taxonomic controversy. The mtDNA diversity of Assamese macaques was greater than that of rhesus macaques, suggesting the earlier speciation of Assamese macaques. The significance of the ecogeographic segregation model of macaque distribution is discussed in relation to the evolutionary range expansion into the Himalayan regions in South Asia.     

Evaluating Dominance Styles in Assamese and Rhesus Macaques

Researchers have suggested that several types of agonistic and affiliative behavior covary as a set of species-specific traits, and have used the term dominance style to describe the covariation. We compared measures of dominance style between a group of Assamese macaques (Macaca assamensis) and a group of rhesus macaques (M. mulatta), though kinship information was unknown. Assamese and rhesus female-female dyads each showed a low proportion of counter aggression and a low conciliatory tendency, suggesting that they have despotic social relationships. They also showed a despotic pattern on several other types of agonistic and affiliative behavior, such as approach outcomes and grooming distributions, which is consistent with the covariation of dominance style traits. Assamese male-male dyads showed relatively high levels of reconciliation and counter aggression versus other macaque males portrayed in the literature, suggesting that Assamese males have a tolerant dominance style. Insofar as macaque dominance style depends on the behavior of females, we suggest that Assamese macaques, like rhesus macaques, have despotic social relationships, which contrasts with evidence of a strong correlation between phylogeny and dominance style in macaques. Further, our results indicate that strong male bonding and tolerant dominance relationships among males are independent of female dominance style. Lastly, some measures of agonistic behavior, such as rate of aggression or proportion of bites, are likely altered in competitive environments and thus are not useful indicators of dominance style.     

Distribution and Abundance of Primates in Rain Forests of the Western Ghats,Karnataka, India and the Conservation of Macaca silenus

We assessed the distribution and abundance of 4 species of diurnal primates viz. lion-tailed macaque, bonnet macaque, Nilgiri langur and Hanuman langur, in 2 areas—Brahmagiri-Makut and Sirsi-Honnavara—in rain forests of the Western Ghats in the state of Karnataka, India. The Nilgiri langurs in Brahmagiri-Makut and the lion-tailed macaques in Sirsi-Honnavara are the northernmost populations of the 2 species in the Western Ghats. The 2 regions represent changes in ecological zones in the Western Ghats. In Brahmagiri-Makut, Hanuman langurs and bonnet macaques occupy lower elevations whereas Nilgiri langurs live in relatively higher altitudes. Only one group of lion-tailed macaques was in Brahmagiri-Makut. In the forests of Sirsi-Honnavara, 3 species of primates viz. lion-tailed macaque, bonnet macaque and Hanuman langur were in larger numbers throughout the forest. A self-sustainable single population of 32 groups of lion-tailed macaques occurred in Sirsi-Honnavara. Few subspecies of Hanuman langurs exist in the study regions. Due to local hunting practices, the relative abundance of primates in Brahmagiri-Makut is lower than that in Sirsi-Honnavara.     

三种笼养灵长类活动时间分配的比较研究

采用焦点动物法和连续记录法对3种笼养灵长类活动时间分配进行比较研究。结果表明,川金丝猴Rhi-nopithecus roxellanae和熊猴Macaca assamensis用于休息的时间多于猕猴Macaca mulatta,移动时间则相反;川金丝猴和猕猴用于理毛的时间多于熊猴。不同性别年龄组之间的活动时间分配有差异,主要表现在川金丝猴母亲用于理毛的时间明显多于成年儿子,用于玩耍的时间则相反;猕猴成年雄性的移动时间和理毛时间多于成年雌性,玩耍时间则少于成年雌性;熊猴成年雌性的移动时间多于成年雄性,与成年个体相比,幼体花更多的时间用于玩耍,而相应的减少了其休息时间。     

Boundary zone between northern and southern pig-tailed macaques and their morphological differences

Based on previous conflicting reports that the two forms of pig-tailed macaque (northern and southern) exist as separate species, subspecies, or forms, and that their boundary zone lies in Thailand, a survey of the distribution range and morphology of pig-tailed macaques in Thailand was conducted during 2003–2010. We first conducted a questionnaire survey. Questionnaires were sent to 7,410 subdistricts throughout Thailand. We then traveled to 72 of the 123 subdistricts reporting the presence of pig-tailed macaques. However, due to a lack of reports of the presence of free-ranging pig-tailed macaques living south of the Isthmus of Kra, a survey of pet pig-tailed macaques was also conducted during 16–24 September 2011. Furthermore, 35 wild northern pig-tailed macaques inhabiting northern Thailand (13°13′N, 101°03′E) were temporarily caught and their morphological characters were measured and then compared to those of the southern form captured from Sumatra, Indonesia. Although largely allopatric, the ranges of the northern and southern pig-tailed macaques in Thailand were found to have a partially sympatric boundary at the Surat Thani–Krabi depression (8–9°30′N). Morphologically, these two forms were very distinctive, with different morphological characters such as the crown patch, the white color of the triangle above the eyes, the red streak at the external rim of the eyes, pelage color, ischial callosity, tail length and carriage, facial height, and limb length in both sexes, and patterns of sex skin swelling and reddening in females. These differences in morphological characters between the northern and southern forms should help settle the problems of their taxonomy.     

A Voucher Specimen for <Emphasis Type="Italic">Macaca munzala</Emphasis>: Interspecific Affinities,Evolution, and Conservation of a Newly Discovered Primate

Sinha, A., Datta, A., Madhusudan, M. D., & Mishra, C. (2005. Macaca munzala: A new species from western Arunachal Pradesh, northeastern India. International Journal of Primatology, 26, 977–989) discovered Arunachal macaques (Macaca munzala), a species new to science, in the eastern Himalaya of Arunachal Pradesh in northeastern India. They depicted the holotype and paratypes of the species in photographs, and a specimen of the species had been unavailable for preservation and examination. In March 2005, we obtained an entire specimen of an adult male Macaca munzala, which we propose as a voucher specimen for the species. We provide detailed morphological and anatomical measurements of the specimen and examine its affinities with other macaques. Macaca munzala appears to be unique among macaques in craniodental size and structure, baculum, and aspects of caudal structure, while exhibiting affinities with the other members of the sinica-group to which it belongs. We summarize our insights on the origins and phylogeny of Macaca munzala. Finally, we review the current conservation status of the macaques, which are threatened by extensive hunting in the only 2 districts of Arunachal Pradesh where they are documented to occur.     

What makes a long tail short? Testing Allen's rule in the toque macaques of Sri Lanka

Allen's rule (1877) predicts ecogeographical anatomical variation in appendage proportions as a function of body temperature regulation. This phenomenon has been tested in a variety of animal species. In macaques, relative tail length (RTL) is one of the most frequently measured appendages to test Allen's rule. These studies have relied on museum specimens or the invasive and time-consuming capturing of free-ranging individuals. To augment sample size and lessen these logistical limitations, we designed and validated a novel noninvasive technique using digitalized photographs processed using LibreCAD, an open-source 2D-computer-aided design (CAD) application. This was used to generate pixelated measurements to calculate an RTL equivalent, the Tail to Trunk Index (TTI) = (tail [tail base to anterior tip] pixel count/trunk [neck to tail base] pixel count). The TTI of 259 adult free-ranging toque macaques (Macaca sinica) from 36 locations between 7 and 2,087 m above sea level (m.a.s.l.) was used in the analysis. Samples were collected from all three putative subspecies (M. s. sinica, aurifrons, and opisthomelas), at locations representing all altitudinal climatic zones where they are naturally distributed. These data were used to test whether toque macaque tail length variation across elevation follows Allen's rule, predicting that RTL decreases with increasing elevation and lower temperature. Our results strongly supported this prediction. There was also a statistically significant, negative correlation between elevation and annual average temperature. The best predictor for the TTI index was elevation. Significant subspecies differences in RTL are linked in part to their ecological and altitudinal niche separation, but overall the variation is seen as the species' adaptation to climate. The method developed for the quick morphometric assessment of relative body proportions, applicable for use on unhabituated free-ranging animals, widens the range of materials available for research studying morphological characteristics and their evolution in primates.     

弄岗熊猴的活动节律和活动时间分配

2005年9月至2006年9月,采用瞬时扫描取样法,在弄岗国家级自然保护区对野生熊猴(Macaca assamensis)进行了连续跟踪观察,收集相关的活动节律和活动时间分配的数据,通过分析其与气候及食物组成的关系,从中探讨影响熊猴活动节律及活动时间分配的因素。研究结果发现:在熊猴的日活动节律中,觅食活动表现为逐渐增强的趋势,最高峰值出现在下午15:00,休息活动在中午出现一个小的高峰;其日活动节律表现出明显的季节性差异,主要表现为旱季大部分时间段的觅食强度均高于雨季,且早上7:00-10:00出现一段长时间的休息,而雨季里长时间的休息则发生在中午12:00-14:00;在活动时间分配上,熊猴平均花费39.6%时间用于休息,33.2%用于移动,18.3%用于摄食,5.1%用于理毛,2.4%用于玩耍,1.4%用于其他行为,其活动时间分配也表现出明显的季节性差异;与雨季相比,在旱季熊猴明显增加用于觅食的时间,而相应地减少用于休息的时间,活动时间分配的季节性变化明显与食物组成的变化有关;在不同年龄组个体间,青少年猴花费更多的时间用于玩耍,而用于休息和理毛的时间明显少于成年猴。     

四种猕猴属动物精母细胞联会复合体的比较研究

本工作采用去污剂微铺展——硝酸银染色技术研究熊猴、平顶猴、藏酋猴、恒河猴及其亚种毛耳猴的精母细胞联会复合体（SC）核型、SC的结构及其在减数分裂中的行为。结果表明这几种动物的SC核型以及SC的发育过程基本一致。SC的形成开始于偶线期,成熟于粗线期,解体于双线期。在减数分裂前期,性染色体轴呈强嗜银性,配对明显落后于常染色体。根据减数分裂前期性染色体的形态和行为,性染色体的配对可分为五种类型。此外,本文还对XY染色体的同源性和侧轴加粗等现象进行了讨论。     

太行山猕猴研究进展与展望

猕猴广泛分布于从阿富汗到泰国和中国南部的亚洲地区.分布于中国的猕猴有6个亚种.太行山猕猴为猕猴华北亚种Macaca mulatta tcheliensis,系中国所特有,目前仅分布于河南、山西两省交界的太行山和中条山南端地区,为世界野生猕猴分布最北的种群.自20世纪80年代初期开始,学者们在数量与分布、形态学(包括普通形态学、骨骼系统形态学)、生态学、微观生物学、疾病、实验技术等方面,对太行山猕猴展开了研究.本文对已有的研究成果进行了综述,并提出了今后亟待开展的研究工作,以期能有助于对这一珍稀物种的有效保护.     

Macaque–human interactions and the societal perceptions of macaques in Singapore

Humans and long‐tailed macaques (Macaca fascicularis) interface in several locations in Singapore. We investigated six of these interface zones to assess the level of conflict between the two species. We observed macaque‐to‐human interactions and distributed questionnaires to residents and visitors of nature reserves. We observed an average of two macaque‐to‐human interactions per hour at the sites, which included affiliative or submissive behaviors (46.9%), aggression (19.1%), taking food and other items (18.5%) searching bins, cars, and houses (13.4%), and nonaggressive contact (2.1%). Two‐thirds of interactions occurred when a human was carrying food or food cues, and one‐quarter occurred when a human provoked macaques. Only 8% of interactions occurred without a clear human‐triggered context. Our interview showed one‐third of respondents experienced nuisance problems from macaques. They had items taken from them (50.5%) and received threats (31.9%). Residents reported more nuisance problems than visitors, and their perceptions toward macaques differed. Residents were more aware of the consequences of food provisioning and that there were regulations against feeding. Residents fed macaques less and held more negative sentiments toward macaques. Nearly half of the interviewed people held neutral attitudes toward macaques and only 26.2% of respondents thought conflict with macaques warranted urgent action. Nearly two‐thirds of the respondents supported education programs to ameliorate human–macaque conflict, and less than 15% supported removing or eradicating macaques. 87.6% felt that it is importance to conserve and protect macaques. Our results show that human–macaque conflict exists in Singapore, but that it may not be severe. Human behavior is largely responsible for macaque‐to‐human interactions, and thus could be lessened with management of human behavior in interface zones (i.e. restrict food carrying and provocation). Moreover, our interviews shows people living in Singapore value macaques, do not wish them entirely removed, prefer education‐based solutions, and consider conservation and protection of them important. Am. J. Primatol. 71:825–839, 2009. © 2009 Wiley‐Liss, Inc.     

Counter aggression and reconciliation in Assamese macaques (Macaca assamensis)

Patterns of aggressive and affiliative behavior, such as counter aggression and reconciliation, are said to covary in the genus Macaca; this is referred to as the systematic variation hypothesis. These behavior patterns constitute a species dominance style. Van Schaik's [1989] socioecological model explains dominance style in macaques in terms of within- and between-group contest competition. Dominance style is also said to correlate with phylogeny in macaques. The present study was undertaken to examine phylogenetic and socioecological explanations of dominance style, as well as the systematic variation hypothesis. We collected data on counter aggression and reconciliation from a habituated group of Assamese macaques (Macaca assamensis) at the Tukeswari Temple in Assam, India. The proportion of agonistic episodes that involved counter aggression was relatively low. Counter aggression, however, occurred more often among males than among females, and it was most common when females initiated aggression against males. The conciliatory tendency for this group of Assamese macaques was 11.2%. The frequency of reconciliation was low for fights among males and for fights among females, but reconciliation was particularly rare for opposite-sexed opponents. Female social relationships were consistent with the systematic variation hypothesis, and suggest a despotic dominance style. A despotic dominance style in Assamese macaques weakens the correlation between dominance style and phylogeny in macaques, but it is not inconsistent with the socioecological model. Male-female relationships were not well explained by the despotic-egalitarian framework, and males may well have more tolerant social relationships than do females. Sex differences need to be considered when categorizing species according to dominance style.     

The effect of island area on body size in a primate species from the Sunda Shelf Islands

Aim  We examine the effect of island area on body dimensions in a single species of primate endemic to Southeast Asia, the long-tailed macaque ( Macaca fascicularis ). In addition, we test Allen's rule and a within-species or intraspecific equivalent of Bergmann's rule (i.e. Rensch's rule) to evaluate body size and shape evolution in this sample of insular macaques.
Location  The Sunda Shelf islands of Southeast Asia.
Methods  Body size measurements of insular macaques gathered from the literature were analysed relative to island area, latitude, maximum altitude, isolation from the mainland and other islands, and various climatic variables using linear regression.
Results  We found no statistically significant relationship between island area and body length or head length in our sample of insular long-tailed macaques. Tail length correlated negatively with island area. Head length and body length exhibited increases corresponding to increasing latitude, a finding seemingly consistent with the expression of Bergmann's rule within a single species. These variables, however, were not correlated with temperature, indicating that Bergmann's rule is not in effect. Tail length was not correlated with either temperature or increasing latitude, contrary to that predicted by Allen's rule.
Main conclusions  The island rule dictating that body size will covary with island area does not apply to this particular species of primate. Our study is consistent with results presented in the literature by demonstrating that skull and body length in insular long-tailed macaques do not, strictly speaking, conform to Rensch's rule. Unlike previous studies, however, our findings suggest that tail-length variation in insular macaques does not support Allen's rule.     

Molecular phylogeny of macaques: implications of nucleotide sequences from an 896-base pair region of mitochondrial DNA

We determined the nucleotide sequences of an 896-base pair region of mitochondrial DNA (mtDNA) from 20 primates representing 13 species of macaques, a baboon, and a patas. We compared these sequences and the homologous sequences from four macaques and a human against each other and deduced the phylogenetic relationships of macaques. The results from the phylogenetic analyses revealed five groups among the macaques: (1) Barbary macaque, (2) two species of Sulawesi macaques, (3) Japanese, rhesus, Taiwanese, crab-eating, and stump-tailed macaques, (4) toque, pig-tailed, and lion-tailed macaques, and (5) Assamese and bonnet macaques. The phylogenetic position of Tibetan macaque remains ambiguous as to whether it belongs to the fourth or fifth group. Phylogenetic trees revealed that Barbary macaque diverged first from the other Asian macaques. Subsequently, the four groups of Asian macaques diverged from one another in a relatively short period of time. Within each group, most of the species diverged in a relatively short period of time following the divergence of the groups. Assuming that the Asian macaques diverged from the outgroup Barbary macaque three million years ago (MYA), the divergence times among groups of Asian macaques were estimated at 2.1-2.5 MYA and within groups at 1.4- 2.2 MYA. The intraspecific nucleotide diversity observed among three rhesus macaques was so large that they did not form a monophyletic cluster in the phylogenetic trees. Instead, one of them formed a cluster with Japanese and Taiwanese macaques, whereas the other two formed a separate cluster. This implies that either polymorphisms of mtDNA sequences that existed before the divergence of these three species (ca. 700,000 years ago) have been retained in rhesus macaques or introgression has occurred among the three species.