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。     

白头叶猴日活动时间分配及其季节性变化

<正>活动节律和活动时间分配是动物行为学研究的两个重要方面,它们直接与动物的代谢和能量收支相关,但它们又会随环境不同不断调整。通过比较不同生态条件下动物的活动节律和活动时间分配,     

Habitat Quality and Activity Budgets of White-Headed Langurs in Fusui,China

Within a species, habitat quality may be a factor causing different activity budgets between populations. The habitat of white-headed langurs (Trachypithecus leucocephalus) has been seriously disturbed in Fusui Rare and Precious Animal Nature Reserve, China, where we carried out a study of their socioecology from September 1997 to September 1998. We collected data on langur activity budgets from the main population located in the central part of a group of limestone hills. We classified habitat quality into 4 grades according to the extent of human disturbance. We showed that the two main study groups of white-headed langurs spent on average 50% of time resting, 13% feeding, 18% moving (including foraging), 11% grooming, and 7% playing. Langur time budgets showed no significant seasonal change, but they differed among different sex-age classes. Infants and juveniles spent about 20.3% of time playing, whereas adults spent only 0.2% playing. The group in high quality habitat engaged less in feeding and more in playing than the group in low quality habitat did. Habitat quality influenced the playing time of young white-headed langurs and may be vital to their successful maturation.     

Ranging behavior of the François’ langur (<Emphasis Type="Italic">Trachypithecus francoisi</Emphasis>) in the Fusui Nature Reserve,China

Data on activity budgets and ranging patterns were collected from March to December 2001 for one group of François’ langurs (Trachypithecus francoisi) inhabiting a forested part of the Fusui Nature Reserve, Guangxi province, China. Our results indicate that the total size of the home range of the study group during the study period was 19 ha. The majority of their activities (52%) occurred within a small area, 22%, of their home range, and was concentrated in or near quadrats containing their sleeping sites, which may reduce the time and energetic cost of travel. The extent of the ranging behavior varied between months, with the smallest, 7 ha, recorded in July and the largest, 13.5 ha, in November. There was no significant difference between seasons. The monthly mean daily path lengths varied from 341 to 577 m. The daily path lengths showed significant seasonal changes: the path lengths were longer during the dry season than in the rainy season, which may be related to the scarcity of preferred food resources during the dry season.     

Sleeping Site Use by <Emphasis Type="Italic">Trachypithecus francoisi</Emphasis> at Nonggang Nature Reserve,China

We collected data on sleeping site use of the François’ langur (Trachypithecus francoisi) between August 2003 and July 2004 at Nonggang Nature Reserve, China. We tested hypotheses regarding possible ultimate causes of sleeping site selection in light of our results. Langurs selected the ledges and caves on cliffs as sleeping sites. Of 23 identified sleeping sites, 7 were more frequently used than the others (≥9 times each, accounting for 64% of total observed nights). Langurs used most sleeping sites repeatedly, and reused some of them on consecutive nights; 4 consecutive nights were the longest run. We suggest that langurs choose sleeping sites to make approach and attack difficult by predators, and to increase familiarity so as to improve chances for escape. Langurs’ cryptic behaviors before entering sleeping sites and the rapid movement toward sleeping sites (4 min on average) with an increased level of vigilance may help to decrease the possibility of detection by predators. Access to food appears to have a profound influence on sleeping site selection in François’ langurs, as demonstrated by the langurs’ tendency to select sleeping sites close to their current main feeding sites. The position of sleeping site relative to the last feeding site of the day and the first feeding site of the subsequent morning indicated a strategy closer to that of a multiple central place forager than of a central place forager. Our results do not support the influences of other factors, e.g., avoidance of parasites, seeking comfort, and range or resource defense, on sleeping site selection.     

环境和社会因素对喀斯特石山猕猴日活动节律和活动时间分配的影响

2007年9月至2008年8月,以广西弄岗国家级自然保护区内的一大群野生猕猴为研究对象进行连续的跟踪观察,采用瞬时扫描取样法收集其食物组成、活动节律和活动时间分配的数据,旨在探讨影响石山猕猴活动节律及活动时间分配的环境和社会因素。结果表明：研究猴群的日活动节律表现为上午和下午的觅食高峰以及中午的休息高峰,这与白天温度和日照强度的变化有关。日活动时间分配中,休息42.1%,移动34.5%,觅食15.1%,玩耍2.0%,理毛5.2%,其他行为1.3%。猴群依据食物的变化调整活动时间分配,其中用于移动的时间与食物中果实的比例成显著的正相关性。不同性别年龄组个体的活动时间分配表现出显著差异,主要表现为成年雄性花费更多的时间休息,而成年雌性花费更多的时间觅食和社会性理毛,未成年个体则花费更多的时间玩耍。与同域分布的小群猕猴相比,本研究群花费更多的时间休息,较少的时间觅食。这些差异可能与盗食农作物有关。     

The effect of urban and rural habitats and resource type on activity budgets of commensal rhesus macaques (Macaca mulatta) in Bangladesh

Macaques are characterized by their wide distribution and ability to adapt to a variety of habitats. Activity budgets are affected by habitat type, season, and food availability in relation to differing age–sex class and individual requirements. We conducted a comparative study on two commensal rhesus groups, one living in a rural village and the other in the center of urban Dhaka, Bangladesh. The study was conducted in three different seasons between 2007 and 2009 in order to evaluate how habitat type and season affects their behavioral activities. Differences in food type and its availability between these two habitats were mainly responsible for the variations in activity budgets between groups. Feeding time in the rural group was significantly longer than that in the urban group. In contrast, grooming and object manipulation/play were significantly greater in the urban than the rural group. Seasonal variations in all major behaviors were significantly affected by group, with more time spent feeding in summer than in winter/dry season, and more time spent grooming and moving in winter/dry season than summer in the rural group. In contrast, time spent resting was greater in the monsoon and summer seasons than the winter/dry season in the urban group. Grooming time was greater in the winter/dry season than the monsoon and summer seasons. In both groups, immature of both sexes spent significantly more time on feeding and object manipulation/playing and less time resting than adults. Adult females spent more time grooming than males and immatures, of both sexes, in both groups. Moreover, the rural group spent most of their time feeding on garden/crop produce and wild plant food resources, while the urban group spent more time feeding on provisioned foods. These results showed that differences in the activity budgets of rural and urban dwelling macaques were due largely to the differences in available food resources. Commensal rhesus macaques show a high degree of behavioral flexibility in response to habitat and resource variability, and knowledge of these differences is important for the conservation and management of highly commensal primates.     

广西扶绥黑叶猴活动节律和日活动时间分配

2002 年8 月至2003 年7 月,对生活在完全被周围农田隔离的石山中的一群黑叶猴的昼夜活动节律和日活动时间分配进行研究。采用以群为单位的焦点动物观察法和连续记录法,每月对猴群连续10 d 跟踪观察。结果表明:3 月至10 月,黑叶猴的昼夜活动节律由8 个环节组成,11 月至2 月由9 个环节组成,移动、觅食和休息在活动节律中重复出现,晒太阳仅出现在11 月至翌年2 月。休息占黑叶猴日活动时间的比例最大,其次是觅食、移动、晒太阳和理毛。黑叶猴的日活动时间分配具有明显的月变化规律,相关性分析表明,环境温度、栖息地的食物丰富度和光照强度是影响黑叶猴日活动时间分配的主要因素。进一步分析认为,黑叶猴的活动时间分配体现了叶食性灵长类的特点,更体现了黑叶猴对北热带喀斯特石山环境的适应特点。     

圈养阿拉伯狒狒日活动时间格局及季节差异

动物个体行为的生态效应主要反映在它们的活动节律和时间分配上.2012年11月至2013年10月,在杭州野生动物园,用瞬时扫描取样法对1群圈养阿拉伯狒狒的日活动节律和活动时间分配进行了调查研究.结果表明: 1)在活动时间分配中,主要活动(频次百分率≥5%)包括休息(42.5%)、游动(16.2%)、玩耍(10.0%)、摄食(9.8%)、理毛(9.4%)和观望(7.3%);频次百分率低于5%的称为次要活动,共4.8%.2)阿拉伯狒狒的活动时间分配具有明显的季节性变化,冬季摄食和观望时间增加,玩耍和理毛时间减少,但休息和游动行为没有显著季节差异.3)阿拉伯狒狒通常在上午和下午分别出现摄食高峰,并随着气温逐渐转冷,摄食活动逐渐增加,出现3个摄食高峰;这可能是动物在寒冷季节为抵御寒冷而采取的补充能量的一种策略.4)在温暖季节,阿拉伯狒狒只有中午一个休息高峰和一个理毛高峰,更多时间用于游动和玩耍,表现为多峰特征;在寒冷的冬季,休息行为也体现了多峰特征,但其游动、玩耍和张望时间节律并不与温暖季节有明显差异.因此,杭州地区圈养阿拉伯狒狒的冬季日活动节律与许多灵长类(如黑冠长臂猿、维氏冕狐猴和绒毛猴等)不一样,采取摄食增加能量,而非休息减少能量消耗的策略.总之,阿拉伯狒狒的行为体现出明显的节律性,且其活动节律具有季节性差异,这些可能是阿拉伯狒狒适应于圈养环境特有的食物资源和温度变化的影响而形成的.     

Sleeping Cave Selection,Activity Pattern and Time Budget of White-Headed Langurs

We describe the activity patterns and time budgets of white-headed langurs that were confined to about 4 km² of Longlin habitat in Fusui County, Guangxi Province, China. Between February and December 1996, we observed 6 langur groups monthly via group focal sampling and continuously recording the behavior of a focal group. Our results indicate that the langur groups selected stone caves on cliffs as sleeping sites. The daily activity pattern outside caves had 8 stages: (1) leaving the cave in the early morning; (2) moving and resting; (3) morning feeding; (4) moving a long distance; (5) resting at noon; (6) afternoon feeding; (7) moving back to the cave, and (8) entering the cave. Over the year, langurs spent a daily average of about 11.5 h outside caves and about 12.5 h inside caves. Moving accounted for 7% (spring), 7% (summer-autumn) and 13% (winter) of the time budget, and langurs spent 9% (summer-autumn) and 14% (spring) to 20% (winter) of their time feeding. Resting accounted for 79% of the time budget in spring, 84% in summer-autumn, and 57% in winter. Sunbathing only occurred in winter and accounted for about 10% of the time budget. One-way ANOVA and multiple range tests demonstrated that time budgets differed significantly among seasons. Langurs spent significantly more time feeding and moving in winter than in spring and summer-autumn, but significantly less time resting in winter than in spring and summer-autumn.     

Variation in gaze-following between two Asian colobine monkeys

Gaze-following is a basic cognitive ability found in numerous primate and nonprimate species. However, little is known about this ability and its variation in colobine monkeys. We compared gaze-following of two Asian colobines—François’ langurs (Trachypithecus francoisi) and golden snub-nosed monkeys (Rhinopithecus roxellana). Although both species live in small polygynous family units, units of the latter form multilevel societies with up to hundreds of individuals. François’ langurs (N = 15) were less sensitive to the gaze of a human experimenter than were golden snub-nosed monkeys (N = 12). We then tested the two species using two classic inhibitory control tasks—the cylinder test and the A-not-B test. We found no difference between species in inhibitory control, which called into question the nonsocial explanation for François’ langur’s weaker sensitivity to human gaze. These findings are consistent with the social intelligence hypothesis, which predicted that golden snub-nosed monkeys would outperform François’ langurs in gaze-following because of the greater size and complexity of their social groups. Furthermore, our results underscore the need for more comparative studies of cognition in colobines, which should provide valuable opportunities to test hypotheses of cognitive evolution.     

白头叶猴与黑叶猴觅食行为的比较

开展近缘物种觅食行为比较对理解动物的行为可塑性及适应性具有重要意义。白头叶猴（Trachypithecus leucocephalus）和黑叶猴（T.francoisi）是近缘物种,体形大小相近,社会结构和栖息环境相似,是广西崇左白头叶猴国家级自然保护区喀斯特季节性雨林中邻域分布的灵长类。为了探索两个物种在喀斯特生境中是否有相似的觅食策略,我们于2012年1-12月采用瞬时扫描取样法对两种叶猴的觅食行为进行研究。结果表明,白头叶猴与黑叶猴在不同时段均为叶食性,树叶是两种叶猴各个时段主要食物,其中白头叶猴日均取食树叶77.0%±4.4%,黑叶猴日均取食68.9%±8.3%,两者对树叶的采食比例均没有显著的日时段差异（白头叶猴：χ²=6.602,df=11,P=0.830;黑叶猴：χ²=11.393,df=11,P=0.411）。两种叶猴的觅食行为都在猴群清晨离开夜宿石洞后和进入过夜山洞前的时段中频繁发生。白头叶猴在09：00-10：59和16：00-17：59出现觅食高峰,时间占比分别为41.7%和46.3%;黑叶猴同样在09：00-10：59和16：00-17：59的时间段内发生高频率的觅食行为,时间占比分别为31.3%和38.0%。此外,两种叶猴的觅食时间在大部分时段中的差异并不明显。白头叶猴和黑叶猴在石灰岩生境中具有相似的觅食策略,这意味着两种叶猴在取食生态学上可能采取相似的保护措施。     

秦岭川金丝猴的昼间活动时间分配

1999年12月至2004年11月,通过野外直接跟踪观察,采用焦点取样和瞬时记录的方法,对秦岭川金丝猴西梁猴群的昼间活动时间分配进行了研究。将川金丝猴的昼间活动定义为摄食、休息、理毛、移动和其他行为5 种类型,结果发现：川金丝猴昼间活动中有两个摄食高峰,分别在每天的上午和下午,中午有一个较长时间的休息期。川金丝猴昼间的主要行为类型是摄食和休息（约占全部活动时间的62%）,各种行为依所占时间比例依次为：摄食（34.64 ± 1.70%）、休息（26.83 ± 1.69%）、理毛（11.58 ± 0.96%）、移动（18.93 ±1.08%）和其他行为（8.02 ± 0.82%）。在不同季节各种行为类型所占时间比例有明显的变化,表现为摄食和休息行为占昼间活动时间比例在夏（75.00%）、秋季（74.15%） 大于春（52.74%）、冬季（56.64%）;而移动、理毛和其他行为所占时间比例之和在夏（25.00%）、秋季（ 25.85%） 则明显小于冬（ 43.4%）、春季（47.3%）。不同季节间的活动时间分配经非参数检验（ Kruskal Wallis H test,N = 199）,理毛行为（ x² =11.438,df = 3,P < 0.05）有显著差异,摄食（x² = 20.469,df = 3,P < 0.01）、休息（x² = 27.235,df = 3,P < 0.01）、移动（x² = 24.110,df = 3,P < 0.01） 和其他行为（ x² = 60.810,df = 3,P < 0.01） 季节间差异非常显著;除摄食行为外,其余各行为类型的时间分配在不同性别-年龄组间也存在显著差异（P < 0.05）。     

干扰生境下黑叶猴(Trachypithecus francoisi francoisi)日活动时间的分配

1999年至2000年,采用瞬间采样法对贵州省麻阳河国家级自然保护区干扰生境中的野生成年黑叶猴的日活动规律进行了研究。结果表明：干扰生境中野生成年黑叶猴在全年的行为时间分配中表现为休息和摄食所占比例最多,分别为35．41％和31．67％;其次是游走、拥坐和嬉戏,分别为14．44％、9．61％和8．54％;仅将很少时间用于理毛行为,为0．33％。与未干扰生境相比,干扰生境下黑叶猴全年中除用于休息的时间下降外,用于其余活动类型的时间均有所提高。研究还表明野生成年黑叶猴的活动规律在不同季节存在一定差异性。休息和摄食行为在各季节日活动规律中都存在显著高峰期,但高峰期持续时间和强度不同。游走、嬉戏和拥坐行为在各季节的日活动规律所占比例均较低。理毛行为存在季节有或无的现象。     

Time budget of daily activity of Francois'' langur (Trachypithecus francoisi francoisi) in disturbance habitat

The daily activity pattern of wild Francois' langurs (Trachypithecus francoisi francisi) was observed in disturbed habitat at Mayanghe National Nature Reserve of Guizhou Province, China, between 1999 and 2000. The behaviors of wild adult Francois' langurs were recorded at a 5-min interval using an instantaneous sampling method. The results indicate that resting, foraging, traveling, huddling, playing and grooming activities account for 35.41%, 31.67%, 14.44%, 9.61%, 8.54% and 0.33%, respectively, in the daily time budget of these adult Francois' langurs during the study period. In comparison with the results in undisturbed habitat, the decreased food availability in disturbed habitat changes the foraging behavior of langurs, and also changes time budgets of other behaviors, such as the increase of the proportion of traveling and the decrease of the proportion of resting. At the same time, the higher frequency of human activity in the disturbed habitat influences the time budget of daily social behavior of the langurs including playing, huddling and grooming. The patterns of activity rhythms of wild adult Francois' langurs in the disturbed habitat show some seasonal differences which are also discussed.     

Diurnal activity budget of African buffalo (Syncerus caffer Sparrman, 1779) in Chebera Churchura National Park,Ethiopia

One of the fundamental questions in animal ecology concerns the activity pattern of animals and the environmental and intrinsic factors that influence such dynamics. This study tested the hypotheses that activity time budgets of the African buffalo appeared to vary by season and times of day and predicted that buffalo would express unequal proportion of time for different activity patterns during the wet and dry seasons in Chebera Churchura National Park (CCNP). An investigation on the diurnal activity budget of the African buffalo was carried out during the wet and dry seasons of 2012–2014 in the Chebera Churchura National Park, Ethiopia, using focal‐animal sampling method. Buffalo spent a greater proportion of the time in feeding and resting/ruminating activities in both the wet and dry seasons. Feeding and resting (lying down and standing) were the predominant activities (87.14% of the diurnal active period), 48.95% time spent feeding during the dry season and 44.91% during the wet season. There was a significant decrease in feeding and an increase in resting from dry seasons to wet seasons. Daytime grazing and resting periods during the wet season were estimated to be 5.39 h and 4.98 h, respectively. Morning and the late afternoon activity peaks were more pronounced during the dry season than the wet season. Therefore, feeding and resting time was influenced by the time of day and the seasons. But there were no significant difference in time allocation for other activities in both the wet and dry seasons. The study has implications for understanding animal activity budget across species, particularly relationships between temperature and season.     

Factors Influencing Interannual and Intersite Variability in the Diet of <Emphasis Type="Italic">Trachypithecus francoisi</Emphasis>

Comparative studies of the diet of the same species in different habitats, and over the long term, are essential to understanding a species’ behavioral and ecological plasticity. Moreover, such studies can help researchers and managers evaluate a species’ capacity to cope with changes in habitat quality resulting from natural processes or human disturbance, which is important for developing conservation strategies. We compared dietary data for François’ langur (Trachypithecus francoisi) collected at Nonggang and Fusui Nature Reserves, Guangxi Province, China, over 2 separate study periods to evaluate interannual and intersite variation in diet. Young leaves were the preferred staple foods for langurs, whereas mature leaves and seeds served as fallback foods in response to seasonal shortage in the abundance of young leaves. Species composition of the diets and the percentage of feeding records for plant species varied between the 2 study periods. The langurs at both study sites fed selectively, and they did not base their diet simply on the abundance of plant species in the habitat. However, the plant species eaten by langur groups inhabiting the 2 different reserves were markedly different, and the top 10 food species eaten by the Fusui group showed no overlap with those eaten by the Nonggang group. The variation may be related to differences in forest composition resulting from different level of human disturbance. In summary, our results indicate that François’ langurs exhibit a comparable dietary pattern both temporally and geographically, but there is marked interannual and intersite difference in species composition of the langur diet.     

邦亮东黑冠长臂猿日食性与活动节律的季节性变化

日食性变化和活动节律是动物行为的重要方面,通过对日食性变化与活动节律的研究,可以了解食物、温度和光照时间等环境因素的变化对动物行为的影响,以及动物在行为上的应对策略。2009 年1 -12 月,在广西靖西县邦亮东部黑冠长臂猿自治区级自然保护区内,采用瞬时扫描取样法对2 群东黑冠长臂猿觅食行为和活动节律进行了研究。结果显示,东黑冠长臂猿的活动和食性在旱季和雨季都具有明显的节律。鸣叫只发生在12:00之前,但旱季的鸣叫时间偏晚。在日节律上,东黑冠长臂猿在离开过夜树后和进入过夜树前各有一个取食高峰,07:00 喜欢取食果实和无花果,16:00 也选择更多的果实,但无花果较少,13:00 - 15:00 取食更多的叶和芽。在10:00 - 12:00,其用于休息和社会活动的时间增加。在雨季,东黑冠长臂猿以果实和无花果为主要食物,并且花大量时间觅食无脊椎动物;但在旱季,其食物以叶和芽为主。与之相适应,东黑冠长臂猿在旱季通过减少移动增加休息来节约能量开支和应对低温,同时它们增加取食的时间以获得等价的能量和营养。雨季时社会行为的比例明显高于旱季,而且其高峰出现在08:00 - 10:00,但旱季社会行为主要出现在11:00 - 14:00。在雨季东黑冠长臂猿没有出现休息高峰,相反在旱季10:00 左右出现一个明显的休息高峰。上述行为节律体现了东黑冠长臂猿对环境季节性变化的良好适应,使其能够在寒冷并且退化的喀斯特森林中生存繁衍。     

圈养白头叶猴春季昼夜活动节律

2008 年3 ～5 月,采用瞬时扫描法,对上海动物园5 只白头叶猴的行为进行24 h 昼夜连续观察。结果表明:春季圈养白头叶猴一天的活动起于06∶ 00 左右,结束于18∶ 00 或18∶ 30。白天白头叶猴的主要行为为休息、取食和移动,平均频次依次是13.79 次、4.75 次和2.18 次。夜间的主要行为为休息、移动和抓挠,平均频次依次是22.13 次、0.43 次和0.26 次。不同个体昼夜活动节律差异很大。其中,移动、理毛、玩耍和其它行为的差异显著。夜间无理毛和玩耍行为,移动行为在夜间21∶ 00, 00 ∶ 30, 03∶00 和04∶ 30 出现高峰。在22∶ 30, 01∶ 00, 02∶ 00, 03∶ 30, 04∶ 30 和05∶30 则出现抓挠高峰。这预示白头叶猴夜间休息时处于一个“轻睡眠”状态。同时,雌雄白头叶猴昼夜移动行为差异显著。     

Food diversity and choice of white-headed langur in fragmented limestone hill habitat in Guangxi,China

We studied the diet and food choice of three groups of white-headed langurs (Trachypithecus leucocephalus) from July 2002 to June 2003 at Fusui Rare Animal Reserve, Guangxi, China. Data were collected from via focus group sampling, with continuous recording. A total of 109 plant species (including 19 unidentified species) were used as food by the white-headed langur. They spent 91.6% of total feeding time feeding on leaves, 4.2% on fruits and 3.3% on stems. Flowers and other food items accounted for only 0.3% and 0.6% of total feeding time, respectively. There were significant seasonal variations in the langurs’ diets with changes in the total number of species and the diversity of food species consumed. The langurs used significantly more plant species as food in the rainy season than in the dry season (36 species vs. 26.5 species). Dietary diversity was also significantly higher in the rainy season than in the dry season (2.6 vs. 2.4). No significant seasonal variation was found in the percentage of various plant parts in their monthly diets. Though the white-headed langur fed on many plant species, 11 species made up 84.9% of their diet. There were no significant correlations between the percentage of feeding time for main food plant species and their corresponding abundance in the habitat, content of water, crude protein, crude fiber and the ratio of crude protein to crude fiber. These results suggest that food choice of the white-headed langur is not influenced by their abundance and nutritional content.