Characterizing human-macaque interactions in Singapore

Previous studies have noted substantial human-macaque interactions involving physical contact in Bali, Indonesia; Gibraltar; and Mt. Emei, China [Fuentes, American Journal of Primatology 68:880-896, 2006; Zhao, Tibetan macaques, visitors, and local people at Mt. Emei: problems and countermeasures. In: Paterson and Wallis, editor. Commensalism and conflict: the human-primate interface. Norman, OK: American Society of Primatologists. p 376-399, 2005]. The aim of this study was to conduct preliminary observations in order to begin to characterize interaction patterns between humans and long-tailed macaques (Macaca fascicularis) in Singapore. Unlike Bali, Gibraltar, and Mt. Emei, Singapore occasionally enforces fines and penalties and engages in an education campaign in an effort to minimize physical contact between humans and macaques. Observers stationed at two sites in Singapore conducted 92 5 hr of observation that included 730 human-macaque interactions over 16 days. Data recorded include interaction characteristics, demographic and behavioral variables, presence of feeding by humans, and presence of automobiles. Although feeding by humans was relatively infrequent overall, it generally occurred most often by individuals in cars and when human children were present. Data analysis suggests that interactions involving physical contact between macaques and humans are rare in Singapore, in contrast to the findings from Bali, Gibraltar, and Mt. Emei. This low level of physical contact suggests a low risk of macaque-human pathogen transmission in Singapore.     

Status of the long-tailed macaque Macaca fascicularis in Singapore and implications for management

The long-tailed macaque (Macaca fascicularis) population of the island-state of Singapore consists of ca. 1,218–1,454 individuals. About seventy percent of the population (ca. 1,027 individuals) is concentrated in both Bukit Timah and Central Catchment Nature Reserves, a system of reservoirs and forest reserves located in the center of Singapore. This core population resides mainly along perimeter forest areas of the reserve system, which is bordered by residential and recreational areas (e.g., parks and golf courses) and encircled by expressways. Periphery sub-populations (ca. 427 individuals) persist in forest fragments throughout Singapore mainland and on 5 offshore islands. Much of the Singaporean macaque population overlaps with human settlement and these commensal groups are mainly distributed close to roads, parks and residential areas. At least 70% of these groups are habituated to human presence and at least 50% to food provisioning. Moreover, commensal groups have more individuals and have higher infant:adult female ratios than non-commensal groups. The close association of habituated macaque groups living in human environments has led to increasing human-macaque conflict in Singapore. The overlap is also associated with human-induced population loss resulting from road accidents (2.4%); and trapping efforts (14%) aimed at ameliorating conflict issues. Consequently, it is important to better understand how humans are affecting macaque populations. In order to mitigate human-macaque conflict and at the same maintain a sustainable macaque population in Singapore, there is an urgent need for wildlife management strategies aimed at minimizing the extent of human–macaque conflict. Such strategies should include designing appropriate buffers around reserve areas, revised urban development plans, and managing the behavior of people interfacing with macaques.     

Human–wildlife conflict: Proximate predictors of aggression between humans and rhesus macaques in India

Macaques live in close contact with humans across South and Southeast Asia, and direct interaction is frequent. Aggressive contact is a concern in many locations, particularly among populations of rhesus and longtail macaques that co‐inhabit urbanized cities and towns with humans. We investigated the proximate factors influencing the occurrence of macaque aggression toward humans as well as human aggression toward macaques to determine the extent to which human behavior elicits macaque aggression and vice versa. We conducted a 3‐month study of four free‐ranging populations of rhesus macaques in Dehradun, India from October–December 2012, using event sampling to record all instances of human‐macaque interaction (N = 3120). Our results show that while human aggression was predicted by the potential for economic losses or damage, macaque aggression was influenced by aggressive or intimidating behavior by humans as well as recent rates of conspecific aggression. Further, adult female macaques participated in aggression more frequently than expected, whereas adult and subadult males participated as frequently as expected. Our analyses demonstrate that neither human nor macaque aggression is unprovoked. Rather, both humans and macaques are responding to one another's behavior. Mitigation of human‐primate conflict, and indeed other types of human‐wildlife conflict in such coupled systems, will require a holistic investigation of the ways in which each participant is responding to, and consequently altering, the behavior of the other. Am J Phys Anthropol 156:286–294, 2015. © 2014 Wiley Periodicals, Inc.     

Differences in Activity Budgets and Diet Between Semiprovisioned and Wild‐Feeding Groups of the Endangered Barbary Macaque (Macaca sylvanus) in the Central High Atlas Mountains,Morocco

The Barbary macaque, Macaca sylvanus is a very adaptable primate species occupying a wide range of habitats in Morocco and Algeria. Several groups of this endangered macaque can be found in tourist sites, where they are affected by the presence of visitors providing food to them. We compare the activity budgets and the diet of semiprovisioned and wild‐feeding groups of Barbary macaques in the central High Atlas Mountains of Morocco from February to August 2008. We used instantaneous scan sampling at 15‐min intervals. The behaviors included in the activity budget were feeding, moving, foraging, resting, and aggressive display. Food items were grouped into seven categories. We found no differences between the two groups in the daily percentages of records attributed to feeding. The semiprovisioned group spent significantly more time engaged in resting and aggressive behavior, and foraged and moved significantly less than the wild‐feeding group. There was no significant difference between the two groups in time spent eating leaves, fruits, or roots and bark. The semiprovisioned group, however, spent significantly less time per day feeding on herbs, seeds, and acorns than the wild‐feeding group. Human food accounted for 26% of the daily feeding records for the semiprovisioned group and 1% for the wild‐feeding group. Our findings agree with previous studies and indicate that in the tourist site, where food is highly clumped, macaques decreased foraging time yet showed higher levels of contest competition. Our results support the common claim that the diet of the Barbary macaque is highly flexible, differing among its varied habitats. Conservation efforts for the Barbary macaques should take into account the changes in behavior that human‐modified environments may cause.     

Interactions between visitors and Formosan macaques (Macaca cyclopis) at Shou‐Shan Nature Park,Taiwan

Ecotourism involving feeding wildlife has raised public attention and is a controversial issue, especially concerning nonhuman primates. Between July 2002 and April 2005, the behavior of monkeys and tourists was collected through scan samplings, focal samplings and behavior samplings at the Shou‐Shan Nature Park located in Taiwan's second largest city—Kaohsiung. In addition, the number of tourists and monkeys was counted in different hours and places within the park. Four hundred visitors were interviewed using a questionnaire to gather data on sex, age, purpose and frequency of visit to the park. The number of tourists was significantly higher during weekends than in weekdays in all locations. Humans dominated in the initiation of interspecies interactions—the overall ratio of human‐initiated and monkey‐initiated interactions was 2.44:1. Human–monkey conflicts accounted for only 16.4% of the total interactions (n=2,166), and adult human males and adult male macaques participated in higher rates than other age/sex groups in these conflicts. Visitors showed more affiliative behavior (15.9%) than agonistic behavior (8%) toward the macaques. In response to visitors' threat or attack, the Formosan macaques mostly showed submissive behavior with bared teeth, squealed or ran away to avoid confrontation (69.1%)—only few responded with counteraggression (18.7%). This study for the first time provided evidence that food provisioning increased both the frequency and duration of aggression among Formosan macaques (P<0.001). During food provisioning, the average frequency and the duration of agonistic events of macaques were more than 4 times higher compared with those without food provisioning. The average frequency of food provision by tourists was 0.73 times/hr—more than twice the incident that monkeys grabbed the food from tourists (0.34 times/hr). If people refrain from feeding monkeys and destroying the city park's natural vegetation, monkeys can be used to educate public about nature conservation in an urban setting. Am. J. Primatol. 71:214–222, 2009. © 2008 Wiley‐Liss, Inc.     

Influence of Fruit Availability on Fruit Consumption in a Generalist Primate,the Rhesus Macaque <Emphasis Type="Italic">Macaca mulatta</Emphasis>

Fluctuations in resource availability occur in all ecosystems. To survive, species must alter their foraging strategies according to the quantity, quality, and distribution of available food. The rhesus macaque (Macaca mulatta), a commensal primate, is considered a generalist omnivore and very few studies have addressed how its feeding strategies change with respect to resource availability. We examined dietary diversity and frugivory levels in a group of rhesus macaques at the Buxa Tiger Reserve in northern India across one year. Using behavioural observations of diet and phenological monitoring, we found that although rhesus macaques fed on 107 food items including leaves, flowers, fruits, seeds, and insects, fruits made up ca. 74% of their diet. Fruit consumption correlated positively with fruit availability, but fruit preference appeared to play an important role; 16% of all the fruit species they fed on accounted for >50% of all fruit feeding observations. We suggest that afforestation programs involving preferred fruit species at the agricultural land–forest interface would prevent forest groups of rhesus macaques from gravitating toward human habitations and reduce conflict over anthropogenic resources. We further propose that the movement of certain primates in the direction of human habitations may be contingent on resource availability and food preference rather than an inherent propensity to gravitate to anthropogenic areas.     

Visitor Perceptions of Captive,Endangered Barbary Macaques (Macaca sylvanus) Used as Photo Props in Jemaa El Fna Square,Marrakech, Morocco

ABSTRACT

Visitors to Jemaa El Fna Square can pay to have their photograph taken with Barbary macaques. Our aim was to characterize visitors’ perceptions of such photo props, enabling demand-reducing interventions to be targeted appropriately and destination managers to maintain or enhance the visitor experience. Visitors to Jemaa El Fna (n = 513) were surveyed using a 25-item questionnaire including closed and open questions. Most visitors (88%) neither intended to use macaque photo props nor did; 7% either intended to or did use photo props; while 5% both used photo props and had intended to do so. Moroccans were more likely than foreigners to use macaque photo props or intend to do so. Among international visitors, those who had their photo taken or intended to were younger, with a lower level of education and lower income than those who neither had their photo taken nor intended to. Visitors who did not use photo props pitied the animals’ treatment, disapproved of captivity or exploitation, had concerns over hygiene or safety, and disliked trader harassment. Visitors who did use photo props valued novelty and contact with the animal, although half of them also described negative experiences including trader harassment and animal mistreatment. While 16% felt the presence of macaques makes Marrakech lively and interesting, 40% recognized risks to health and safety, and 57% thought Marrakech would be a better place without macaque photo props. Although 66% agreed the practice should be illegal, 80% were unaware that it is illegal. Only 25% correctly identified the Barbary macaque as endangered. Macaque photo props undermine current conservation objectives and legislation, contribute to Disneyfication of macaques, and may threaten the image of the destination; however, their appeal to a minority of visitors indicates a desire to interact with animals, which visitor education might divert toward more responsible tourism.     

Status of urban populations of the long-tailed macaque (<Emphasis Type="Italic">Macaca fascicularis</Emphasis>) in West Sumatra,Indonesia

We studied long-tailed macaque (Macaca fascicularis) populations in Padang, West Sumatra, Indonesia, focusing on the effect of human provisioning on their demography and dietary composition. We conducted a field survey at three sites in the city: Gunung Meru, Gunung Padang, and Gunung Panggilun. Mean troop size (range 28–68) and infant ratio (range 0.38–1.00) were greater in Gunung Meru, where the macaques have been highly provisioned, than at the other two study sites (troop size 10–15; infant ratio 0.00–0.33). The macaques at all sites consumed both natural and human foods, but dependence on the latter differed among sites: three-quarters of the diet of macaques in Gunung Meru consisted of human foods, while human foods comprised less than 5% of the macaque diet at the other sites. The ability of macaques to modify the proportion of human food is a behavioral flexibility that facilitates the survival of the long-tailed macaque in urban habitats. Without restrictions on provisioning, the degree of dependence of macaques on human foods and population size could increase, especially in Gunung Meru, and human–macaque conflict could escalate. In order to create an effective management policy for urbanized monkeys, long-term quantitative data on macaque behavior and monitoring of population parameters are required.     

Male–Male Relationships in Lion-tailed Macaques (<Emphasis Type="Italic">Macaca silenus</Emphasis>) and Bonnet Macaques (<Emphasis Type="Italic">Macaca radiata</Emphasis>)

Socioecology suggests that female distribution in space is determined by the distribution of food resources and the male distribution is influenced by female distribution. Though studies have traditionally focused on females, males have received increasing attention in recent years. We compared male–male relationships in lion-tailed macaques and bonnet macaques. Because bonnet macaques have a high adult male:female sex ratio and are seasonal breeders whereas lion-tailed macaques have a low adult male:female sex ratio and are largely aseasonal breeders, we predicted that bonnet macaque males would be spatially and socially more tolerant of each other and would have less linear dominance relationships than lion-tailed macaques. We recorded male–male and male–female relationships in 1 group of wild macaques of each species via scan sampling and 1–0 sampling. The results revealed that lion-tailed macaque males largely remained at a distance from each other whereas bonnet macaque males remained in close proximity to one another. Lion-tailed macaque males were more agonistic toward each other whereas bonnet macaque males showed more affiliative interactions. The dominance hierarchy among lion-tailed macaque males was more linear than among bonnet macaque males. Our data support the hypothesis that the study of spatial structuring, temporality of interactions, and linearity of social relationships may contribute to a better understanding of macaque social systems.     

Within-group female-female agonistic interactions in Taiwanese macaques (Macaca cyclopis)

Feeding-related agonism among wild female Taiwanese macaques was investigated in two study groups at ecologically diverse sites (Fu-shan and Ken-ting) to determine whether contest-feeding competition was present in these groups. Females that contest for food within a primate group are hypothesized to form dominance hierarchies and tend to be philopatric. In this study we tested 1) whether Taiwanese macaque females show higher agonism in a feeding context, 2) whether they exhibit stronger agonism over higher-quality foods, and 3) whether higher agonism rates occur in smaller food patches. Female Taiwanese macaques at both study sites showed similar agonism rates in a feeding context (0.30 events/hr). They exhibited higher agonism rates inside food patches than outside food patches in the spring. Higher agonism rates occurred during seasons of higher fruit availability, and a lower agonism rate occurred in winter when the macaques switched to feeding on fallback foods. Females in the Fu-shan group exhibited higher proportions of aggressive interactions over higher-quality foods, such as animal matter and the reproductive parts of plants. In the Ken-ting group, 95.8% of feeding-related agonistic interactions among females occurred over fruits. Agonistic interactions that occurred in small food patches tended to result in the agonism recipient leaving the food patch. We conclude that female Taiwanese macaques show contest feeding competition in certain contexts. The patterns we observed have also been documented in other primate species in which females are philopatric and form linear dominance hierarchies.     

Mother-infant interactions of wild-born,individually-caged cynomolgus monkeys (Macaca fascicularis) during the first 14 weeks of infant life

This study documents age-related changes in the interactions of wild-born cynomolgus macaque mothers and their infants living in individual cages during the first 14 weeks of infant life. Body contact between mother and infant, maternal holding, and infant sucking were found to decrease, and the mothers showed an increased frequency of aggression toward their infants with age. These results were broadly similar to those reported for mother-infant interactions in other macaques living in social groups. Nevertheless, a clear difference between the present cynomolgus macaques and other macaques in social groups was apparent. The cynomolgus macaque mothers tended to permit their infants to move about freely without displaying maternal protectiveness such as restraint or retrieval, unlike other macaque mothers in social groups. Such maternal behaviors might derive from the experience of living in individual cages for many years and the relative safety of living in individual cages. The lack of maternal restraint and retrieval could be responsible for the observed sex differences in behavior: male infants moved more actively, and broke, and made contact with their mothers more frequently than did female infants. Moreover, mothers of female infants held and groomed them more frequently and were less aggressive toward them.     

海南南湾猴岛景区内猕猴与游客接触行为的研究

在世界范围内,观赏和投喂野生猿猴是最常见的野生动物旅游形式之一。其中游客与动物的关系越来越成为动物学、人类学和社会学学者研究关注的热点。游客在娱乐的过程中,存在被猿猴威胁甚至咬伤等问题。对人猴接触行为的客观分析,是提高旅游服务管理和保障游客安全的必要措施。本研究记录了2012-2014年海南南湾猴岛景区内游客与猕猴的接触行为,并分析了景区医务室近两年的游客就诊记录,主要探讨：（1）哪些游客会参与人猴接触,及其游客的认识误区;（2）哪些猕猴会频繁参与人猴接触;（3）哪些原因会引发人猴接触行为。本文旨在了解景区内游客与猿猴的互动关系,探讨游客受伤原因及其解决办法,为完善野生动物旅游的健康发展提供基础资料。     

The Hand That Feeds the Monkey: Mutual Influence of Humans and Rhesus Macaques (<Emphasis Type="Italic">Macaca mulatta</Emphasis>) in the Context of Provisioning

Historically, humans and other primates (primates henceforth) have coexisted across cultures and contexts, and many primate populations use anthropogenic food sources as their main or supplementary food. While primates may actively forage for such food, they are also directly provisioned by humans in many regions. Ethnoprimatology views humans and primates as cohabitants of integrated socioecological spaces who mutually influence each other’s ecologies and social lives. We contextualized provisioning of primates by humans within an ethnoprimatological framework and examined if the availability of anthropogenic food affected primate diets or the amount of time primates spent in anthropogenic habitats and whether primates influenced the human act of provisioning. To this end, we used scan sampling on a group of rhesus macaques across a year, and conducted interviews with 86 people who paused at a nearby tea shop for refreshments. We found that the macaques’ consumption of natural resources and dietary diversity decreased, and they spent more time in human-modified habitats when provisioned food was available. We also found that particular behaviors of the provisioned macaques stimulated provisioning by humans. Our findings show that provisioning influences macaque feeding ecology and habitat use, and that the behavior of the macaques themselves drives people to provide them with food subsidies, illustrating a complex web of interactions between the sympatric species.     

Community Perceptions of the Crop-Feeding Buton Macaque (<Emphasis Type="Italic">Macaca ochreata brunnescens</Emphasis>): an Ethnoprimatological Study on Buton Island,Sulawesi

Human–wildlife overlap is increasing worldwide as a result of agricultural expansion. This can reduce human tolerance of wildlife, especially if wildlife threatens human food sources. The greatest threat to the declining populations of the endemic Buton macaque (Macaca ochreata brunnescens) is habitat destruction, but as a common crop-feeding species, there is also an additional risk of retaliation killings from farmers. Finding means of reducing this risk will thus help secure the long-term future of this range-restricted subspecies. Here, we investigate variability in farmers’ perceptions of primate crop-feeding and mitigation techniques in three farming communities on Buton Island, Indonesia, which differ in wealth and agricultural resources. We employ a mixed methodology, collecting qualitative social data from focus groups and quantitative observational data to measure macaque crop-feeding occurrences. Our findings indicate that the least wealthy community used lethal control methods more frequently than the comparatively wealthier communities, even when the crop-feeding problem was less severe. The least wealthy community also expressed high levels of fear of macaques, and had the most negative perceptions of them. This community also had no knowledge of the macaques’ conservation status or their ecological roles. We recommend that efforts to protect Buton macaques focus on education and the use of effective nonlethal mitigation techniques, such as electric fencing. We also suggest that initiatives to support such measures may be most effectively directed toward communities with relatively low economic wealth and high reliance on subsistence agriculture, especially where crop-feeding wildlife is feared, even when such communities do not experience the highest losses from crop-feeding wildlife.     

The Human–Macaque Interface: Conservation Implications of Current and Future Overlap and Conflict in Lore Lindu National Park, Sulawesi, Indonesia

Here, I examine overlapping resource use of forest and cultivated resources by villagers and tonkean macaques (Macaca tonkeana) in Lore Lindu National Park, Sulawesi, Indonesia. An integrative research design was employed, using methods from primatology and cultural anthropology, in conjunction with quantitative measurements of cacao crop loss. The results indicate that the current patterns of overlapping resource use may not be severely affecting the tonkean macaques or villagers in Lindu. The research does, however, point to possible future difficulty if cacao crop raiding by macaques increases, and as changing socioeconomic trends and loss of traditional folklore result in further modification of macaque habitat. Successful strategies to ameliorate human–macaque conflict may require efforts aimed at the adoption of alternative buffer zone crops that use shade-management systems, the deliberate protection of important macaque food species, and increasing local tolerance to crop raiding by exploring the role of macaques in forest regeneration.     

Disproportionate participation by age/sex classes in aggressive interactions between long-tailed macaques (Macaca fascicularis) and human tourists at Padangtegal monkey forest, Bali, Indonesia

We observed 420 aggressive interactions between tourists and Macaca fascicularis at the Padangtegal Wanara Wana Temple forest site in Bali, Indonesia, during the months of June and July 2001. The data collected included patterns of aggression, presence or absence of food, and demographic information on resident macaques and human visitors. Analyses of the interactions suggest that macaques respond differentially to humans according to the age/sex classes involved. Additionally, adult and subadult male macaques participated in more aggressive behaviors than expected, while adult female macaques and immatures participated in such behaviors less than expected. These variations in interaction patterns between macaques and tourists may have substantial implications for management issues and the potential for pathogen transmission.     

Qualitative Assessment of Macaque Tourist Sites in Padangtegal,Bali, Indonesia,and the Upper Rock Nature Reserve,Gibraltar

Human tourists share space with, touch, feed, and otherwise interact with Macaca at multiple locations. Across Asia and stretching to Gibraltar and Northern Africa, macaques form a substantial tourist attraction as their ranges become increasingly coincident with human use zones. Residing in or as tourist attractions, macaques frequently generate economic benefits to local humans and compete with them for habitat and specific resources. In addition, health conflicts may emerge from increased overlap and interactions between humans and macaques. There is only a handful of studies on the impact, structure, context, and cultural ecology of macaque tourist sites. We provide a general overview of 2 macaque tourist sites, one in Padangtegal, Bali, Indonesia and one in Gibraltar. Qualitative assessment reveals variability in behavioral, ecological, economic, and politically relevant facets of macaque tourism. Specifically, differences in aggression during interactions, potential health dangers, economic factors, and local human perceptions between Padangtegal and Gibraltar suggest that qualitative assessments can assist in our construction of broader models and conceptualizations of the interaction context.     

Post-Conflict Affiliation in Barbary Macaques is Influenced by Conflict Characteristics and Relationship Quality, but Does Not Diminish Short-Term Renewed Aggression

Many group living primates have evolved mechanisms to repair their social relationships after conflicts ('reconciliation'). We analysed the post-conflict behaviour of female Barbary macaques, Macaca sylvanus , living in the enclosure 'La Forêt des Singes' at Rocamadour, France. Based on a sample of 914 conflicts, we investigated whether relationship (kinship, rank, affiliation, support and sex) and conflict characteristics (conflict intensity, context and duration) affected the quality and frequency of affiliative post-conflict interactions. Thirty-two per cent of all conflicts were followed by post-conflict affiliation. Rates of socio-positive interactions and support were better predictors of post-conflict affiliation than kinship or rank. Short conflicts were followed by post-conflict affiliation relatively more frequently, after a shorter latency, but only briefly, and such interactions were initiated by both parties equally frequently. The majority of affiliative post-conflict interactions occurred immediately after the end of the conflict. In sum, female Barbary macaques invest more in post-conflict affiliation with valuable partners, and they modulate their post-conflict behaviour in relation to conflict characteristics. Remarkably, affiliative post-conflict interactions increased the short-term probability of renewed aggression by the former aggressor to 16% compared with 9% for conflicts that were not followed by affiliative behaviour. Such renewed aggression after post-conflict affiliation occurred particularly frequently among females and after conflicts over food, suggesting that post-conflict affiliation sometimes falsely lures the former victim to stay in the vicinity, even at the risk of receiving renewed aggression.     

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.      

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.