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.     

Macaca thibetana at Mt. Emei,China: II. Birth seasonality

Birth censuses were conducted every 2 or 3 days for each of six groups of Macaca thibetana along trails at Mt. Emei in southwest China from March 7 to June 15, 1986. Based on direct observations and the timetable of forehead hair growth and behavior, each of 32 infants could be placed in one of sixteen 14-day periods of the 1986 birth season. The mean estimated birth date was March 27 (SD = 39 days); the median estimated birth date was March 14. Sex ratios in newborns and yearlings did not deviate significantly from 1:1. Seasonal birth timing was correlated with the altitude of the range (r = ?0.84, P < .05); that is, infants were born earlier in the season at higher altitudes.     

Dominance Style Among Macaca thibetana on Mt. Huangshan, China

The dominance style concept has proven useful for understanding covariation patterns in relationship qualities, particularly among macaques. However, the dominance styles of many macaques, including Tibetan macaques (Macaca thibetana), have not been examined in detail. We describe patterns of bidirectionality of aggression, postconflict affiliation and kin bias in a group of wild, but provisioned Tibetan macaques over a 2-yr period in order make an initial assessment of their dominance style. Bidirectional aggression, including percentage of counteraggression (1.9%), and conciliatory tendencies (6.4%) were consistently low across partner combinations, seasons and locations (forest vs. provisioning area). In addition, females consistently displayed high levels of kin bias in affiliation and tolerance. Compared with macaque species with better known dominance styles, the Tibetan data generally fell within the range for despotic species and outside the range for relaxed species. Although other researchers have tentatively classified them as tolerant or relaxed, we conclude that Tibetan macaques display a despotic dominance style. This conclusion poses complications to explanations based both on phylogenetic inertia and socio-ecological models.     

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.     

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.     

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.     

Responses to Seasonal Changes in Nutrient Quality and Patchiness of Food in a Multigroup Community of Tibetan Macaques at Mt. Emei

Provisioning along pedestrian trails by tourists much increased the nutrient quality and patchiness of food (NqPF) for Tibetan macaques (Macaca thibetana) at Mt Emei in spring and summer. In the habitat at a temperate-subtropical transition zone, the macaque's NqPF could be ordered in a decreasing rank from spring-summer to autumn to winter. With the aid of a radio-tracking system, I collected ranging data on a multigroup community in three 70-day periods representing the different seasons in 1991–92. Rank-order correlation on the data show that with the decline of NqPF, the groups tended to increase days away from the trail, their effective range size (ERS), their exclusive area (EA) and the number of days spent in the EA, and reduced their group/community density and the ratio of the overlapped range to the seasonal range (ROR). In icy/snowy winter, the macaques searched for mature leaves slowly and carefully in the largest seasonal range with a considerable portion that was not used in other seasons. Of the responses, the ROR decreased with the reduction in group/community density; and the ERS was the function of both group size (+) and intergroup rank (–) when favorite food was highly clumped. All above responses were clearly bound to maximize foraging effectiveness and minimize energy expenditure, and their integration in term of changes in time and space leads to better understanding macaque ecological adaptability. Based on this study and previous work on behavioral and physiological factors, I suggest a unifying theory of intergroup interactions. In addition, as the rate of behavioral interactions was also related to the group density, Waser's (1976) gas model probably applies to behavioral, as well as spatial, data on intergroup interactions.     

黄山短尾猴（Macaca thibetana）的主要食性分析

2011年11月至2012年10月,作者采用粪便显微分析法研究了黄山短尾猴的四季食性.结果表明：黄山短尾猴共取食植物26科50种;不同季节间黄山短尾猴食性具有明显变化,冬季取食13科25种植物,春季取食23科46种植物,夏季取食14科32种植物,秋季取食17科37种植物;壳斗科、樟科是黄山短尾猴全年的主要食物,占短尾猴总取食量的51.26%～59.75%,其他科属植物也占有重要地位;甜槠在四季均为黄山短尾猴的主要取食食物,豹皮樟在冬季取食量相对较高,秋季最低,红楠、杉木在冬春季节的取食比例差异最大;黄山短尾猴采取与其他灵长类相似的觅食策略;黄山短尾猴四季食物生态位宽度差异不明显,受人为影响较为明显.     

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.     

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.     

Intergroup interactions in Tibetan macaques at Mt. Emei,China

Data on intergroup-interactions (I-I) were collected in 5 seasonally provisioned groups (A, B, D, D₁, and E) of Tibetan macaques (Macaca Thibetana) at Mt. Emei in three 70-day periods between 1991 April–June (P1), September–November (P2), December–1992 February (P3). The I-I were categorized as forewarning made by high-ranking males (including Branch Shaking and/or Loud Calls), long-distance interactions in space (specified by changes in their foraging movements), and close encounters (with Affinitive Behavior, Male's Herding Female, Sexual Interaction, Severe Conflict, Adult Male-male Conflict, Opportunistic Advance and Retreat, etc. performed by different age-sex classes). From periods P1 to P3, the I-I rate decreased with reduction in population density as a positive correlate of food clumpedness or the number of potential feeders along a pedestrian trail. On the other hand, from the birth season (BS, represented by P1 and P3) to the mating season (MS, represented by P2) the dominance relation between groups, which produced a winner and a loser in the encounters, became obscure; the proportion of close encounters in the I-I increased; the asymmetry (local groups over intruders) of forewarning signals disappeared; the rate of branch shaking decreased; and sometimes intergroup cohesion appeared. Considering that sexual interactions also occurred between the encountering groups, above changes in intergroup behaviors may be explained with a model of the way in which the competition for food (exclusion) and the sexual attractiveness between opposite sexes were in a dynamic equilibrium among the groups, with the former outweighing the latter in the BS, and conversely in the MS. Females made 93% of severe conflicts, which occurred in 18% of close encounters. Groups fissioned in the recent past shared the same home range, and showed the highest hostility to each other by females. In conspicuous contrast with females' great interest in intergroup food/range competition, adult male-male conflicts that were normally without body contact occurred in 66% of close encounters; high-ranking male herding of females, which is typical in baboons, appeared in 83% of close encounters, and showed no changes with season and sexual weight-dimorphism; peripheral juvenile and subadult males were the main performers of the affinitive behaviors, opportunistic advance and retreat, and guarding at the border. In brief, all males appeared to “sit on the fence” at the border, likely holding out hope of gaining the favor of females both within and outside the group. Thus, females and males attempted to maximize reproductive values in different ways, just as expected by Darwin–Trivers' theory of sexual selection. In addition, group fission was observed in the largest and highest-ranking group for two times (both in the MS) when its size increased to a certain level, and the mother group kept their dominant position in size and rank among the groups that might encounter, suggesting that fission takes a way of discarding the “superfluous part” in order to balance the cost of competition for food and mates within a group, and the benefit of cooperation to access the resources for animals in the mother group. Am J Phys Anthropol 104:459–470, 1997. © 1997 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.     

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.     

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.     

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.     

Gastro-intestinal helminths in a natural population ofMacaca sinica andPresbytis spp. at Polonnaruwa,Sri Lanka

Helminth parasites were identified in a natural population of primates at Polonnaruwa, Sri Lanka. Five fatally wounded or recently deceased toque macaques (Macaca sinica) and three langurs (Presbytis senex andP. entellus) were autopsied. The following nematodes were found:Oesophagostomum aculeatum (Chabertiidae),Streptopharagus pigmentata (Spirocercidae),Physaloptera sp. (Spiruridae),Enterobius vermicularis (Oxyuridae), andTrichuris trichiura (Trichuridae). One cestode,Hymenolepis was also recovered fromP. entellus. Among fresh faecal samples of 210M. sinica, worm eggs ofOesophagostomum andStrongyloides were most abundant, followed byTrichostrongylus. Trichuris andStreptopharagus eggs were found occasionally.     

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.     

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。