Postconflict Affiliation Between Former Opponents in Macaca thibetana on Mt. Huangshan, China

We describe basic patterns of postconflict affiliation between former opponents within a group of wild, provisioned Tibetan macaques Macaca thibetana on Mt. Huangshan, China. Like most primates studied to date, Tibetan macaques reconciled, i.e., overall they engaged in affiliative interaction with opponents at higher rates immediately after an aggressive conflict than at other times. Probabilities of affiliation were enhanced ≤30 s after the end of hostilities. However when we examined sex partner combinations separately, we found unequivocal evidence for reconciliation only for male-male dyads. Tolerant interaction among other partner combinations apparently was not disrupted after a conflict, presumably obviating the need to reconcile. One aspect of reconciliation among males was consistent with other indications of a despotic dominance style: aggressors initiated a higher proportion of affiliative interactions after a conflict than at other times. Another aspect of reconciliation was more typical of relaxed dominance styles: males used specialized behaviors (embraces and same-sex mounts) disproportionately to reconcile. We also found inconsistent evidence for the valuable relationship hypothesis; probabilities of reconciliation were enhanced for male-male dyads with the closest affiliative relationships, but not for those that displayed the most tolerance or mutual agonistic support. We discuss reconciliation and other aspects of conflict management among males in the context of a group with nearly even sex ratios and high male-male mating competition.     

Primate Tourism,Range Restriction,and Infant Risk Among <Emphasis Type="Italic">Macaca thibetana</Emphasis> at Mt. Huangshan,China

Primate tourism is a growing trend in habitat countries, but few assessments of its impact on primate groups are available. We compare infant mortality in a group of Tibetan macaques (Macaca thibetana) 6 yr before the government translocated the group and subsequently used them for tourism (1986–91), 12 yr during management for tourism (1992–2002, 2004), and 1 yr when management was temporarily suspended (2003). We also compare aggression rates among adults before and during management, and test several hypotheses about specific factors (numbers of tourists, degree of range restriction, demographic changes, changes in α-males) that may have harmed infants. Infant mortality was significantly higher during management than before, but it was similar before management vs. during its suspension. After management began, serious attacks on infants occurred shortly before they died, and many infant corpses had bite wounds. Typically, infants sustained wounds after aggression broke out among adults in the provisioning area used for tourist viewing. Adult aggression rates in the provisioning area correlated positively with infant mortality over time. Range restriction accounted for 54.5% of the variation in infant mortality, and was more closely associated with both mortality and aggression than any other specific factor examined. We hypothesize that range restriction led to increased infant mortality by raising aggression levels in the provisioning area. We conclude that infant mortality is useful as an indicator of the impact of tourism on primate groups, and that range restriction is an inappropriate tourism management practice.     

Ranging behavior ofMacaca thibetana at Mt. Emei,China

Food handouts from visitors to Mt. Emei, in The People’s Republic of China, have considerably increased the diversity of food available to an indigenous population of Macaca thibetana.Some 43% of the feeding time was spent at the trail area frequented by tourists. Ranging behavior was of two kinds: wandering around within the group’s most densely used areas and making peripheral excursions between the areas. Three kinds of trail-area use were observed: three-group overlapping, two-group overlapping, and exclusive. M. thibetanatended to use sheltered sites for sleeping, to ensure safety and/or to keep dry in a rainy habitat. Exclusively and successively used sleeping sites were noted. The average size of the home range per group was 3 km ²; the average population density for the entire range was 13/km², and the biomass was 109 kg/km². The population may be growing, a possibility that is also supported by previous analyses using data on group composition.     

Early mother-infant relationships ofMacaca thibetana at Mt. Emei China

Quantitative data on early mother-infant relationships in the Tibetan macaque was collected during the first 23 weeks of infant life in spring, 1987, at Mt. Emei, China. During the first week of life, infants spent 98.3% of their time in ventroventral contact with their mothers. This contact rapidly decreased to 33.8% by the 4th week and thereafter to 0.85% by the 23rd week. Nipple contact decreased relatively slowly from 89.7% to 62.9% within the first 4 weeks of infant life and to 19.8% by the 23rd week. Ventrolateral and ventrodorsal contact appeared by the 2nd week, meanwhile, maternal restraining behavior appeared, and reached a peak by the 3rd week. The mother neither encouraged nor discouraged her infant's independence during 4–8th weeks. Maternal rejection of the infant was first observed when the infant was 11 weeks old and continued thereafter.     

Mating competition and intergroup transfer of males in Tibetan macaques (Macaca thibetana) at Mt. Emei,China

In five groups of seasonally provisioned Tibetan macaques (Macaca thibetana) at Mt. Emei, males were sampled for wounds as an indicator of their competition for females during about 80 days in the 1987 mating season. Quantitative data on intergroup transfer were collected in a period between June 1986 and December 1987. The young adult (YA) males, the most active age-class in mating activity and intergroup transfer, received most of the wounds. Wounds tended to appear more in the front of body for YA and subadults (SA) than they did for middle-old aged (MO) males. This implies that some of the MO males were more active and aggressive in the fights. During the 1.5 year period, 5/6 of the YA and 5/17 of the MO males made intergroup shifts. Although YA males faced a high risk of receiving wounds at transfer, they usually rose in rank. On the other hand, the MO males transferred more smoothly but dropped in rank. The peripheral SA males, which rarely emigrated in the population, were an active component in determining the wounding rate, and the rate and direction of male migration. Three SA immigrants died of severe attacks made by resident males in 1988 and 1991. Adult sex ratios and their variations were considerably reduced with male nonrandom shifts and better conservation of the population.     

Impact of Translocation,Provisioning and Range Restriction on a Group of Macaca thibetana

We examined the possible long-term impact of translocation and subsequent provisioning and range restriction on recruitment rates in a group of rare Tibetan macaques (Macaca thibetana) in Anhui Province, China. We compared demographic data recorded in the group (and in its fission products) during the 6 years before and after translocation and in a group with no history of management. We also compared preliminary data related to nutritional condition, habitat characteristics, and group activities. Infant losses markedly increased following translocation and management. Estimated measures of nutritional condition varied more among adults in the managed group than among those in the never managed group, suggesting that members of the managed group may have experienced higher levels of intragroup competition. We hypothesize that increased levels of intragroup competition for provisioned food may have put infants at increased risk.     

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.     

Male mate choice in Tibetan macaques Macaca thibetana at Mt. Huangshan, China

Though females are generally more selective in mate choice, males may also benefit from mate choice if male reproductive success is limited by factors other than simply the number of female mates, and if females differ in short-term reproductive potential. We studied male mate choice in a free-ranging troop of Tibetan macaques Macaca thibetana at Mt. Huangshan,China, from August 2007 to April 2008. We employed focal animal sampling and all occurrence sampling to record sexual related behaviors. Eight adult females were divided into three female quality categories according to the females' age, rank and parity.Using male mating effort as a proxy for male mate choice, we found that males do distinguish female quality and show time-variant mating strategies. Specifically, females with dominant rank, high fecundity, and middle age attracted significantly more males. Our results suggest that female short-term reproductive potential appears to be an important variable in determining male mating effort. Male Tibetan macaques do exercise mate choice for higher quality females as well as reduce useless reproductive cost, which is consistent with the direct benefits theory of mate choice.     

Tourism and Infant-Directed Aggression in Tibetan Macaques (Macaca thibetana) at Mt. Huangshan,China

ABSTRACT

Previous studies of habituated Tibetan macaques (Macaca thibetana) at Mt. Huangshan, China demonstrated the negative impact that some tourism management strategies had on monkeys' annual infant mortality, but data on precise causes and perpetrators of infant injuries were not available. We worked at the same site and recorded rates of aggression that older monkeys directed toward group infants as a function of tourist numbers and proximity. We found a negative correlation between the number of people on the viewing platform and the frequency of monkeys' aggression toward the group's infants, but the effect is slight. We noted that infant-directed aggression (IDA) was more likely to occur in the provisioning zone where tourists bounded the monkeys on two sides, despite the fact that infants spent very little time in that location. The alpha and beta males engaged in more IDA than expected, while the other three adult males, adult females, and juveniles exhibited less IDA than expected. Males' IDA included biting, chasing, grabbing, and pushing. Adult females bit and grabbed mostly their own infants in the contexts of punishment and weaning. Juveniles' IDA was uncommon and consisted mostly of open-mouth threats. To minimize the severity and frequency of male-toinfant IDA, we recommend that sections of the tourist viewing platform be closed off so that tourists do not surround the monkeys, particularly when infants are near the viewing platforms and tourist numbers are few. The rule against tourists feeding monkeys should also be strictly enforced, so that altercations between the highest-ranked males and other group members, particularly infants, are less likely to occur.     

雄性黄山短尾猴颅骨变量的异速生长研究

对雄性黄山短尾猴颅骨标本的36项主要形态变量指标进行定点测量,并进行了初步的统计分析和异速生长分析。结果表明,大多数变量的统计值在雄性个体间的变化范围都比较大,2/3的变量表现为正的异速生长,1/3的变量表现为负异速生长。颅骨变量的生长存在年龄及部位间的差异,这些差异与雄性的不同生长发育时期、在社群中的不同地位和竞争压力以及取食习性有关。     

Macaca thibetana at Mt. Emei,China: III. Group composition

Data on group composition at the end of the 1986 birth season were collected from six groups of Macaca thibetana. All adult males, the members of group A, and some conspicuous animals were recognized individually. Fourhundred survey sessions were completed. The mean group size was 38.3 (SD = 13.8, range: 28–65); the number of adult females was the best correlate of total group size. The mean adult sex ratio (F:M) across groups was 3:1 (SD = 1.9, range: 1.5–6.5:1), which significantly deviated from 1:1. Sex ratios (F:M) in newborns, juveniles, and all members did not significantly deviate from 1. The ratio of immature animals to adults was 1.5 to 1 (average of groups); that is, 60% of the population was composed of immature animals, and the population was growing.     

Non-reproductive copulation behavior among Tibetan macaques (<Emphasis Type="Italic">Macaca thibetana</Emphasis>) at Huangshan,China

Non-reproductive copulation, which takes place outside of the mating season and does not result in conception and birth, was studied in a free-ranging group of Tibetan monkeys (Macaca thibetana) at Mt. Huangshan, China, in the birth seasons of 1992 and 1997. We employed all occurrence and focal animal samplings to record sexual and related behaviors and affiliation interactions, respectively. Compared with sexual behavior in mating season, non-reproductive copulation occurred at a lower frequency, with less frequent ejaculation, less harassment, shorter mount duration, and an absence of pause with vocalization. It often took place in a situation in which non-lactating females were involved in social conflict or approached males for mating. Neither pregnant nor lactating females were observed to mate in the birth seasons. Copulation during the birth season did not increase a sexually receptive female’s delivery the next year, nor was it associated with increased proximity, grooming, or agonistic aid for the mating pair. However, copulated pairs spent more time co-feeding, presumably reflecting an increased tolerance on the part of the male. Adolescent males, who rarely copulated in the mating season, engaged in mating activity in birth seasons as well. Therefore, though birth-season copulation had no reproductive functions, it may have fulfilled social functions for females, such as post-aggression appeasing by males or gaining access to resources. This also offered good opportunities for adolescent males and females to develop their sexual skills for later competition.     

雌性黄山短尾猴回避近亲交配

本研究分别在交配期(2006年9月-2006年12月)和产仔期(2007年1月-2007年4月)对黄山短尾猴鱼鳞坑YA1群中5只雄猴和5只雌猴成年个体采用目标动物法、随机取样法和连续记录法记录行为参数。研究期间记录交配行为336例:母子交配0例;母系兄妹交配7例(占2.1%),其中强行交配4例;非亲缘关系交配329例(占97.9%),非亲缘关系交配频次显著高于亲缘关系。在交配期,雄猴对亲缘雌猴跟随、性检查频次均低于非亲缘雌猴,接近指数(PMI)在亲缘和非亲缘雌猴间无显著性差异;雌猴对亲缘雄猴交配拒绝率显著高于非亲缘雄猴,接近指数在交配期显著低于非亲缘雄猴,产仔期接近指数在亲缘和非亲缘雄猴间无显著性差异。尽管雄猴在交配选择上趋于避免与亲缘雌猴交配,但某些雄猴仍会主动对有亲缘关系的雌猴邀配或强行交配,雌猴则主动回避。这些结果表明:黄山短尾猴母系亲属间可以通过行为倾向抑制近亲交配发生,且雌猴更主动回避交配,支持了近交回避的双亲投资理论。     

Seasonal changes in body weight of Macaca thibetana at Mt. Emei,China

This 1991–1992 study was designed to expand previous research on body weight (BW) in Tibetan macaques (Macaca thibetana) at Mt. Emei. Data on BW were collected in late autumn (LA) and late winter (LW) in groups ranging above 1,200 m. Over the winter, the BW fell significantly from a mean of 16.8 to 11.4 kg in females and from 19.5 to 17.0 kg in males. The previously reported BW means of 12.8 kg for females and 18.3 kg for males, measured in late spring, are near the center of the annual BW range for this species. In addition, with the sharper decline of female BW (?32% vs. ?13% seen in males), the sexual dimorphism (M /F ) in BW increased from 1.16 in LA to 1.49 in LW. This finding may be related to differential parental investment by two sexes. © 1994 Wiley-Liss, Inc.     

Southward extension of the range ofmacaca thibetana

Based on results of recent fieldwork, the known geographic range of Macaca thibetanais extended approximately 55 km southward, to 23°48’N, in Jinxiu County, Guangxi Province, China.     

Mandibular morphometrics among macaques: The case ofmacaca thibetana

Recent field studies suggest that Macaca thibetana,a large endemic Chinese macaque, may be quite folivorous, distinguishing it from most other macaque species, which tend to be primarily frugivorous. To understand how this diet affects its masticatory system, we conducted a comparative morphometric study of mandibular dimensions. We took linear measurements from male and female mandibles of this species as well as four other macaques—M. fascicularis, M. nemestrina, M. arctoides,and M. assamensis—and four species of Presbytis—P. obscura, P. rubicunda, P. cristata,and P. phayrei—and subjected to them to a variety of analyses. Based on analyses of variances and discriminant analyses on each sex individually, the mandible of M. thibetanacorresponds to expected patterns for folivorous primates with respect to its wide condyles and thick corpora. However, the height of the corpus and symphysis are lower, and the anteroposterior length of the condyle is longer than predicted for a folivore. In addition to interpretations specifically relating to M. thibetana,we also discuss the functional morphology of the other species in light of what is published about their diets.     

黄山短尾猴不同顺位等级雄性个体友好行为的比较研究

社会顺位等级制度在灵长类动物中普遍存在,它对动物个体的繁殖和社会行为有广泛的影响.本研究对黄山短尾猴鱼鳞坑YA2群6个不同顺位成年雄性个体,根据顺位高低,将它们分成3组:高顺位组(2只个体)、中等顺位组(2只个体)、低顺位组(2只个体),采用目标动物取样法对它们的社会行为进行研究,结果表明,3组之间(1)接收和发出架桥行为频次差异显著,高顺位组接收架桥次数最多;(2)接收和发出爬跨行为频次差异显著;低顺位组接收爬跨次数最多;(3)发出呈臀行为频次差异显著,但接收呈臀行为频次差异不显著;高顺位组是接收呈臀次数最多,低顺位组发出呈臀次数最多;(4)摇树行为频次差异显著,高顺位组发出摇树行为最多.因此雄性黄山短尾猴友好行为可以体现出社群顺位等级关系,这种社群顺位等级关系和这些友好行为在维持社群稳定上有重要作用.     

雄性黄山短尾猴(Macaca thibetana)攻击支持雌性以换取交配回报

生物市场理论认为动物个体之间通过某种协定交换有价值的商品,使双方均受益。该研究采用目标动物法、行为取样法和连续记录法,对浮溪黄山野生猴谷鱼鳞坑短尾猴(Macaca thibetana)A1群(YA1群)和A2群(YA2群)成年个体在非繁殖季节(2011年8月—12月)和繁殖季节(2012年2月—5月)的雄性攻击支持雌性行为和交配行为进行研究,探讨雄性攻击支持雌性与交配之间的关系。两猴群在繁殖季节和非繁殖季节雄性攻击支持雌性与交配行为均呈显著正相关;YA2群繁殖季节与非繁殖季节攻击支持后交配频次均显著高于随机交配;YA1群在繁殖季节攻击支持后交配频次与随机交配频次差异不显著,但在非繁殖季节攻击支持后交配频次显著高于随机交配,说明短尾猴成年雄性攻击支持雌性可以换取与该雌性个体的交配回报。本研究验证了生物市场理论中社会行为存在交换,首次证明了雄性攻击支持可以换取雌性的交配回报,为进一步研究雄性性竞争与雌性选择提供了实例。     

Macaca thibetana at Mt. Emei,China: I. A cross-sectional study of growth and development

Six free-ranging groups of Macaca thibetana were studied at Mt. Emei, in southwest China. Patterns of growth and development observed during the study are described for this species for the first time. Data were collected mainly during the birth season of 1986. Food handouts made possible the measurement of body weight and sitting height. Changes in fur color and growth of forehead hair were noted. Dark hair appeared on the broad white forehead of infants at the end of the third month. A triangular patch formed about 30 days later, and full cover developed within 4.5 to 5 months. For the first 1 or 2 weeks, the fur was blackish; it then became yellow, and by the age of 3.5–4.5 months, it was brown or blackish, i.e., adult color. Growth data on body weight and sitting height for different age-sex classes were collected. For adult males, body weight was 18 kg, sitting height (SH) 55 cm, and ponderal index 33. For adult female, body weight was 13 kg, sitting height 47 cm, and ponderal index 27. Females were considered to be adult at age 5 years.