DRAMATIC CONSEQUENCES OF FOOD HANDOUTS TO MACACA-THIBETANA AT MOUNT-EMEI, CHINA

Food handouts to Macaca thibetana at Mt. Emei have had dramatic consequences for both man and monkey as tourism has increased over the last decade. Food handouts and human submissive behaviour facilitate beg-robbing by the monkeys. which can be regarded as a mixed conditioning chain. Because of their lack of understanding of primate behaviour and resulting inappropriate responses, many visitors have lost possessions and have been severely frightened or even injured; in fact there have been 1 0 deaths as an indirect result over that past 8 years. The appropriate human response proved to be the display of dominance to maintain a distance from a beg-robbing monkey. Road-ranging macaques have also been injured or killed by visitors to obtain meat or bones or merely for amusement. Attempts should be made to eliminate the negative effects of food handouts by increasing visitors' awareness of behavioural and ecological aspects and through aversive conditioning of the macaques.

藏酋猴(Macaca thibetana)与猕猴(Macaca mulatta)长骨生长的比较研究

对15具不同年龄组的藏酋猴长骨的形态功能进行统计学处理和与猕猴比较分 析表明, 藏酋猴具有较长的躯干骨, 相对较短的前后肢和桡骨, 较高的肢间指 数(94.7)和低的臂指数(94.1)。在发育过程中, 其肢骨、体重和躯干骨的增长 速率均大于猕猴, 前后肢增长率相近。猕猴后肢的增长明显大于前肢, 具相对较 短的前肢。藏酋猴更适应于地栖生活。图3表5参15

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) over bar/(F) over bar) 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. (C) 1994 Wiley-Liss, Inc.

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.

黑白仰鼻猴（Rhinopithecus bieti）的种群现状、云南境内生境退化和藏猕猴（Macaca thibetana）的叫声

黑白仰鼻猴仅分布于滇西北和藏东南境内，金沙江和澜沧江间的一个狭小区域内。根据以前的研究结果，将暗针叶林和针阔混交林定义为其适宜生境（SH）黑白仰鼻猴的适宜生境由于当地人砍树烧树形成夏季高山牧场逐渐被侵蚀。为了解云南省境内黑白仰鼻猴适宜生境的现状，我们比较了1997年卫星图片和1958年以航片为基础绘制的地形图上夏季牧场和适宜生境的面积，得到以下结果：1）1997年黑白仰鼻猴的适宜生境面积为4169平方公里；2）夏季牧场面积为1923平方公里；3）过去的40年中，适宜生境面积减少了31％（1887平方公里），夏季牧场面积增加了204％（1291平方公里）；4）森林斑块的平均面积从15.6平方公．里减小到5.4平方公里。另外，夏季牧场的面积和当地人口成正相关（R2=053），这意味着黑白仰鼻猴生境的减少和破碎化是当地传统生产方式和人口增长共同作用的结果。目前在云南和西藏境内分别尚存11群和2群黑白仰鼻猴。

藏酋猴(Macaca thibetana)电刺激采精、精液冻存和精子活力的检测

本文研究中国特有的藏酋猴(Macaca thibetana)电刺激采精、精液冻存及精子活力检测。用电刺激方案[(DDR)_(17-c)V]：延搁(delay)17毫秒、间期(duration)17毫秒、刺激频率17脉冲／秒、单方波变电压连续刺激，对笼养藏酋猴进行电刺激采精，测定精液的各种特征参数，包括每次射精量、液化率、精子密度、运动精子的百分率、死活精子以及异常精子比例。并与其它非人灵长类进行了比较。在精液冻存实验中，通过对不同的防冻液、不同的降温程序及防冻液分散系的比较研究，从而确定了适于藏酋猴精液冻存的冻存方案：PSF-4%的甘油-TH-7.5%的小牛血清(PSF-TH/FBS-G)和PSF-MDM。用前一方案冻存藏酋猴精液，复苏运动度为63.58±0.06%；精子的存活率为90.14±0.03%(n=5)。冻存精液用金黄地鼠裸穿透分析法检测，结果表明具有相应新鲜精液穿透力的51.90±0.08%。后一冻存方案能保存88.00±1.03%的运动精子，但运动寿命较短。研究结果表明：1).刺激方案(DDR)_(17-c)V适于藏酋猴的电刺激采精。2).藏酋猴的精液量和精子数在已有过研究的非人灵长类动物中是最多的，藏酋猴有可能成为较理想的精子生物学研究用的非人灵长类动物。3).脂蛋白对维持精子膜的完整可能有重要作用。4).接近生理条件的防冻剂分散系有助于精液的冻存。5).两种冻存方案PSF-TH/FBS-G和PSF-MDM均适于藏酋猴的精液冻存。

MANDIBULAR MORPHOMETRICS AMONG MACAQUES - THE CASE OF MACACCA-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. thibetana corresponds 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.

藏酋猴的分类与分布

该文基于外部形态、毛色、头骨特征和地理分布对藏酋猴(Macaca thibetana)进行了分类整理, 认为藏酋猴在不同地理区域之间的差异已达到了亚种水平, 可分为４个亚种(包括两新亚种); M. thibetana thibetana, M. thibetana pullus, M.thibetana huangshanensis subsp.nov.和M. thibetana guizhouensis subsp. nov。

笼养藏猴交配行为的初步研究

对笼养藏猴交配活动的观察表明: (1)Macaca thibetana属多次跨爬及单次跨爬射精兼而有 之的类型, 而非典型的单次跨爬射精型; (2)在交配活动中, 特别是在性伙伴的选择上, 雌 性和雄性一样起着积极的作用; (3)M. thibetana雌性存在性干扰行为, 并且雌性的社会地 位可能受其交配活动影响。图3参18

峨眉山藏猕猴雄—婴社会行为的补充观察

藏猕猴 (Macaca thibetana) 雄—婴行为的补充数据收集于1997年1～9月、覆盖了整个出生季节和半个交配季节. 分别用1分钟点取样和5分钟事件取样收集雄婴照料(怀抱+接近)和雄—婴—雄三边作用 (triad) 的数据. 焦点动物为两个猴群中的8个婴儿. 数据处理结果表明: ①雄—婴照料集中在4.5～18周龄, 而20周龄内的婴儿平均有7.9％的时间受到雄性照料(雄9.6％, 雌5.6％); ②雄性婴儿较雌性接受的照料偏多, 但同性个体间无差异; ③雄性照料的强度随行为者社会等级的升高而增大;④婴儿介入 triad的频次与其接受照料的强度正相关. 考虑到藏猕猴繁殖单位中非优势雄猴的效配机会极少, 结果③、④支持了先前关于性/新选择介入了藏猕猴猴雄—婴行为进化的结论. 然而, 因为结果②不支持严格的“父—子”照料假设, 这种介入只具有模糊的统计学意义。

SEXUAL-BEHAVIOR OF TIBETAN MACAQUES AT MT EMEI, CHINA

Data on sexual behavior were collected in six groups of semi-commensal Macaca thibetana along the trail on the slope habitat between 1987 and 1989. Ignoring the common items such as mounting, presenting etc., 20 categories of sexual behavior were described. Most of the descriptions were likely to have enlarged the behavior repertoire reported in macaques, showing a great complexity of sociosexual interactions under the principally natural condition. A great diversity of grouping appeared in the mating season. The copulatory pattern was found to be the serial type contrary to previous speculation, and the mount-to-ejaculation ratio was higher in the central subgroup, as compared with the far-peripheral adult subgroup (FAS) with less male and female rivals. An age-class subdivision of sexually active males made it possible to show that the young adult male immigrants were the most active class in sexual activity. Subgrouping form FAS was a ''space-segregation'' tactic of mating for the losers of both sexes in the competition. Some parameters of copulation were also documented.

Male-infant-male interactions in Tibetan macaques

Event-sampling and scans were used for collecting data on male-infant-male triadic interactions, and their effects on member spacing respectively in a group of Macaca thibetana at Mt. Emei in 1989. The group was partially provisioned by human visitors in seasons other than winter, and could be observed closely. In addition, a stable linear male-hierarchy among five males existed for two years since the end of 1987, providing a good social condition for this topic. The triadic interactions were specific to the birth season, and recognized as three types being on a continuum functionally changing from passive ''agonistic buffering'' (4.8%) to active spatial cohesion, which resulted in a significant decline of intermale distances. Positive correlations were documented between the triad initiation rate and the number of females in consort with the males in the mating season (MS), and between the triad reception rate and the number of infants in proximity to the males in the MS when maternal care was significantly reduced. Thus the male's mating effort and kin/sexual selection may deeply be involved in the triad of this species. Considering that the two triad-species, M. sylvanus and M. thibetana, had different levels of paternity, but shared similar foraging conditions, and showed similar intensities of male-infant caretaking, the triad was very likely a byproduct of male-infant caretaking, which was probably shaped to compensate heavy maternal investment to young offspring in harsh conditions. Accordingly, the long-term arguments about the triad in M. sylvanus can be united to a model of the way in which ''male-infant caretaking'' hypothesis works ultimately, and ''regulating social relations'' hypothesis does proximately.

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, D-1, 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 Fl 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% bf 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. (C) 1997 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 mncaque'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 nor 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. Ir! addition, as the rate of behavioral interactions,was also related to the group density, I Waser's (1976) gas model probably applies to behavioral, as well as spatial, data on intergroup interactions.