Food Choice of White-Headed Langurs in Fusui,China

We studied the feeding ecology of white-headed langurs (Trachypithecus leucocephalus) from September 1997 to September 1998 in Fusui Rare and Precious Animal Reserve, SW Guangxi, China. We collected data in the central part of a group of limestone hills where the main population pool of langurs was located. The langurs fed on 50 plant species (belonging to 42 genera from 28 families) out of 164 species in the habitat (belonging to 112 genera from 48 families). Most of the food plant species were not common in the habitat. There are 16 important food species, including 6 key species. White-headed langurs were extremely folivorous, feeding mainly on young leaves (75% of total feeding records) of a broad range of less common plant species, which were in less disturbed areas. Thus, langur groups in high quality habitat had greater access to preferred foods, and the future of langurs in the Reserve may depend on immediate cessation of tree felling.     

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,     

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.     

Seasonal Variation in the Activity Patterns and Time Budgets of <Emphasis Type="Italic">Trachypithecus francoisi</Emphasis> in the Nonggang Nature Reserve,China

Activity patterns and time budgets are 2 important aspects of animal behavior that researchers use to investigate ecological influences on individual behavior. We collected data on activity patterns and time budgets in 1 group of François’ langurs (Trachypithecus francoisi) from August 2003 to July 2004 in the Nonggang Nature Reserve, Guangxi Province, China, via instantaneous scan sampling method with 15-min intervals. The diurnal activity pattern of François’ langurs showed morning and afternoon feeding peaks, with a midday resting peak. Seasonal change was apparent in the activity pattern: 2 significant feeding peaks occurred in the dry season and only 1 significant feeding peak in the rainy season. The group spent an average of 51.5% of the daytime resting. Feeding and moving accounted on average for 23.1% and 17.3% of the activity budget, respectively. Subjects spent little time on social activities, averaging 2% for grooming and 5.5% for playing. Their time budgets showed significant seasonal variation: they spent a greater proportion of time on feeding and less time on resting and grooming in the dry season than in the rainy season. They also differed among different sex-age classes: immatures spent more time playing, whereas adults devoted more time to resting, feeding, and grooming. Correlations between time budgets and food items or food availability clearly indicated that François’ langurs might adopt an energy-maximizing strategy when preferred foods were scarce in the dry season.     

Are Limestone Hills a Refuge or Essential Habitat for White-Headed Langurs in Fusui,China?

We studied the ecology of white-headed langurs (Trachypithecus leucocephalus) in Fusui Precious Animal Reserve, Guangxi, China, in 1997/1998 and conducted surveys of them in 1996–1997 and 2003. We collected data on vegetation fragments and the activities of white-headed langurs at different levels on limestone hills. The langurs selected less fragmented habitats with less human disturbance. They spent ca. 60% of the day at lower levels of the hills during maintenance activities, including feeding. When humans were absent, the langurs even came to the ground. The hills themselves were used as a refuge from human disturbance but were not otherwise essential habitat. Results of this study have implications for improving conservation management for the langurs.     

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。     

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.     

广西扶绥白头叶猴对栖息地的选择与利用

为了解白头叶猴（Trachypithecus leucocephalus）的栖息地利用规律及其影响因素,2016年2月至2017年1月,采用瞬时扫描取样法对广西崇左白头叶猴国家级自然保护区一群白头叶猴的栖息地利用进行了研究。结果表明,白头叶猴对山体不同部位的利用存在显著性差异（χ² =39.467,df=3,P<0.001）,其中,对崖壁（56.75±9.55）%的利用比例最大,其次是对山坡（39.42±10.93）%和山顶（2.98±2.54）%的利用,而对山脚（0.84±1.47）%的利用频率最低。白头叶猴对不同微生境类型的利用存在差异（χ²=27.709,df=3,P<0.001）,其中对乔木（49.37±12.31）%的利用比例最大,其次是裸岩（24.05±13.61）%,随后依次为藤本（15.48±8.01）%和灌木（10.87±5.45）%。白头叶猴主要在山坡上觅食,利用崖壁移动、休息,进行社会活动;主要利用裸岩进行社会活动,觅食、移动、休息主要发生在乔木上。从整体来看,白头叶猴在雨季对乔木的利用频率显著大于旱季（Z=-2.680,n=12,P=0.007）;雨季在山坡觅食频率显著大于旱季（Z=-2.517,n=12,P=0.012）,而在崖壁觅食频率刚好相反（Z=-2.842,n=12,P=0.004）;白头叶猴雨季在乔木休息的频率显著大于旱季（Z=-2.355,n=12,P=0.019）。白头叶猴对栖息地的利用受到温度的影响。白头叶猴对乔木的总体利用频率随着平均温度的升高而增加（r=0.664,n=12,P=0.018）;觅食时,对崖壁、裸岩的利用频率均与平均温度成负相关关系（崖壁：r=-0.685,n=12,P=0.014;裸岩：r=-0.600,n=12,P=0.039）;休息时,对乔木的利用频率与平均温度呈正相关关系（r=0.650,n=12,P=0.022）。不同季节,白头叶猴对栖息地的利用方式不同。白头叶猴的栖息地利用模式可能是在觅食利益和捕食风险之间作出的权衡,并受到环境温度的影响。     

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。     

栖息地质量对黑叶猴活动时间分配的影响

2005 年9 月至2006 年8 月,在广西弄岗国家级自然保护区和扶绥珍贵动物保护区各选择一群黑叶猴作为研究对象,采用瞬时扫描取样法收集相关的行为数据。通过比较两个地理种群活动时间分配的数据来探讨栖息地质量对黑叶猴活动时间分配的影响。结果表明:生活在低质量栖息生境的黑叶猴猴群较生活在高质量栖息生境的猴群花费更多的时间休息和觅食,而用于移动和社会活动的时间相应减少。分析表明,栖息地质量的差异所造成的食物可获得性以及食物组成的差异可能是影响黑叶猴群活动时间分配差异的重要因素。虽然不同活动的峰值在日活动节律中出现的时间存在差异,但总的日活动节律在不同地理种群中表现出相同的趋势,即早晨和下午出现两个明显的觅食高峰,而中午则是长时间的休息;移动的高峰通常发生在觅食高峰之前。     

Lion-tailed macaques (Macaca silenus) in a disturbed forest fragment: Activity patterns and time budget

We describe the activity patterns and time budget of a feral group of lion-tailed macaques that were confined to a disturbed forest fragment of 65 ha and compare the results with those obtained for groups in protected forests. The degraded nature of the study site was reflected in low tree densities, low specific diversity, gaps in the girth distribution of trees, and frequent disturbance by humans. The study group of 43 subjects was twice as large as lion-tailed macaque groups in protected habitats. They spent the most time ranging (34.0%), followed by foraging (23.7%), feeding (17.9%), resting (16.0%), and other activities such as social interactions (8.4%). Monthly variations are significant for all activity categories except ranging. Times spent resting and foraging are negatively correlated (r = −0.5) and show significant seasonal differences. Foraging time was highest from September to November, when key food sources such asCullenia andArtocarpus were absent or marginally available. The study group spent most time (40.4%) at canopy levels between 21 and 30 m. They spent more time each day ranging than resting or feeding and more time terrestrially compared with groups in protected forests. Large group size, poor habitat quality, and seasonal variation in food availability were the major variables affecting their time budget, and these variables accounted for differences from the time budgets of groups in protected forests.     

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.     

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。     

Food Items Consumed by White-headed Langurs in Fusui, China

We studied the socioecology of white-headed langurs (Trachypithecus leucocephalus) from September 1997 to September 1998 in Fusui Precious Animal Reserve, SW Guangxi, China. We collected data on climate, phenology of food plant species, and foods consumed by langurs living in habitats with different levels of human disturbance. Feeding records showed that the most food is from less common plant species. Young leaves were the staple food item in all langur groups, and consumption correlates with availability. Consumption of mature leaves is not significantly correlated with their availability, but the langurs fed on them, and other food items such as fruits and seeds, when the availability of young leaves was low. Langurs in different areas had broadly similar diets, but they varied most in the proportion of supplementary items such as fruit. Groups in more disturbed habitat did not increase the overall proportion of mature leaves in their diet, but instead maintained their intake of young leaves from a greater diversity of species, and incorporated more immature fruit. Conservationists should direct efforts toward protection of rare plant species providing preferred food items, particularly supplementary foods at times when, and in areas where, preferred food items are less available.     

How does the white-headed langur (Trachypithecus leucocephalus) adapt locomotor behavior to its unique limestone hill habitat?

Limestone hill habitats pose unique challenges to langurs. One of the characteristics of this habitat is its cliffs, which account for about 10–20% of the total area. We have never observed langurs falling from the cliffs during our 10-year field study. Five patterns of locomotion were exhibited by the white-headed langur: (1) arboreal ascent and descent, (2) arboreal quadupedalism, (3) terrestrial quadrupadelism, (4) moving on cliffs and (5) leaping on cliffs. Locomotor patterns varied according to the substrate, but terrestrial quadrupedalism accounted for more than 50% of locomotion time. Moving on cliffs and leaping on cliffs may be modes of locomotion unique to the white-headed langur, at least in terms of frequency. White-headed langurs have an intermembral index of 76 and, compared to langurs with a similar intermembral index, are more terrestrial. Further analysis indicates that greater terrestrialism may be the result of adaptation to their limestone habitat. Interestingly, white-headed langurs select caves on the cliff as their sleeping sites, and they exhibit special behaviors for exiting and entering the cave very early in the morning and late in the evening.     

Comparison of locomotor behaviour between white-headed langurs Trachypithecus leucocephalus and Francois'''' langurs T.fran(c)oisi in Fusui, China

We studied the locomotor behaviour of white-headed langurs Trachypithecus leucocephalus and Fran(c)ois' langurs T.fran(c)oisi to test two hypotheses: (1) these monkeys have evolved locomotor ability to support their activities on limestone hills,and (2) Fran(c)ois' langurs have evolved more diverse locomotor skills than white-headed langurs. Data were collected from 1996 - 1998 and in 2005 in Fusui Nature Reserve,Guangxi,and showed that the two species had similar locomotor types,but Francois' langurs had more locomotor modes (26) than white-headed langurs (12). Quadrupedal walking and leaping were two major types,and white-headed langurs were more arboreal than Fran(c)ois' langurs. We suggest that,while keeping their ancestral locomotor types,the two species have evolved new types allowing them to live on limestone cliffs. Compared to white-headed langurs,Francois' langurs have more diverse locomotor modes that probably have allowed them to live in more habitat types. As an evolutionary outcome,Fran(c)ois' langurs have a larger distribution range.     

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。