中国石山叶猴生态学研究进展

喀斯特石山是一类特殊环境,生存在喀斯特石山的动物形成特殊的行为机制以适应这一特殊的环境。石山叶猴是仅分布于喀斯特石山地区的珍稀濒危灵长类动物,属疣猴亚科乌叶猴属的一个种组,包括黑叶猴（Trachypithecus francoisi）、白头叶猴（T.leucocephalus）、德氏叶猴（T.delacouri）、金头叶猴（卡巴叶猴,T.poliocephalus）、越南乌叶猴（T.hatinhensis）、印支黑叶猴（T.ebenus）、老挝乌叶猴（T.laotum）7种。迄今为止,对分布于中国的黑叶猴和白头叶猴的生态学已有较多研究,而对其他石山叶猴的研究较缺乏。本文对中国黑叶猴和白头叶猴的种群生态学、行为生态学、保护生物学研究进行综述,以促进叶猴生态学理论的发展和完善,为我国特有珍稀濒危动物保护提供基础资料。未来研究应关注石山叶猴的种群历史以及对栖息地破碎化和气候变化的适应机制等问题。     

白头叶猴食谱与觅食时间分配的研究

白头叶猴生活在喀斯特石山环境 ,以树叶为食。对其进行跟踪观察表明白头叶猴有食物 42种 ,其中乔木占 42 .8% ,灌木占 2 6.1 % ,藤本占 2 1 .5% ,草本占 9.6% ,食物组成中树叶占63%— 95% ,果实占 5%— 35% ,花占 0— 6%。白头叶猴有两种觅食方式 ,每天猴群有觅食高峰 2次。白头叶猴觅食的持续时间占日活动时间分配的比例在一年中有季节性变化 ,表现为夏天低冬天高 ,分别为 1 0 .0 3% (6月份 )— 2 3.2 1 % (1 2月份 ) ,这与食物的丰盛状况有关。     

基于MaxEnt模式的白头叶猴栖息地评价——以广西崇左白头叶猴保护区为例

针对物种分布格局与其环境变量关系的研究,对于生态廊道规划与环境恢复研究具有重要意义。本文以白头叶猴（Trachypithecus leucocephalus）为研究对象,针对广西崇左白头叶猴国家级自然保护区51个白头叶猴分布点和11个环境变量数据,利用MaxEnt模型（maximum entropy modeling）对栖息地适宜性进行综合评价,通过模型生成栖息地适宜度指数（habitat suitability index,HSI）阈值,划定研究区域为低适生区、适生区和高适生区3种类型,其中低适生区面积为5 061.43 km²,适生区和高适生区面积分别为42.80 km²和20.63 km²。主要环境变量分析表明：年均降水量、土地利用分类、坡度和年平均气温的综合贡献值分别为54.6%、17.4%、11.8%和9.5%,4项环境变量累积贡献值达到93.3%,是影响白头叶猴分布的主要环境因素。为更有效保护白头叶猴及其栖息地,建议在保护区规划生态廊道,扩大恢复区内白头叶猴适宜栖息地面积,以促进白头叶猴种群间基因交流。     

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。     

鄱阳湖围垦区藕塘越冬白鹤的时间分配与行为节律

2016年12月—2017年3月,采用瞬时扫描法研究了鄱阳湖围垦区藕塘生境中白鹤Grus leucogeranus越冬期的行为,共扫描2560次,23219只次,包括18031只次成鹤和5188只次幼鹤。结果表明,觅食(41.78%)、警戒(25.02%)、修整(15.00%)和休息(10.84%)是白鹤越冬期的主要行为。与自然生境相比,藕塘生境中白鹤主要采取多休息和修整的策略来节省能量支出。成鹤的觅食行为(35.29%)显著低于幼鹤(62.42%)(F_(1,12)=45.977,P0.01),警戒行为(28.66%)则显著高于幼鹤(10.26%)(F_(1,12)=38.975,P0.01)。家庭群成鹤觅食行为(43.96%)极显著高于非家庭群成鹤(27.04%)(F_(1,12)=60.169,P0.01)。家庭群成鹤需要喂食幼鹤,它们花费更多的时间觅食弥补能量的消耗。行为节律上,白鹤各时段觅食行为占总行为的比例均较高,11:00—11:59出现明显高峰,占总行为的48.64%。警戒行为无明显的低谷和高峰。成鹤在各时段的觅食行为比例均明显低于幼鹤,成鹤和幼鹤的觅食曲线变化趋势相似,即觅食比例的升高和降低比较同步。成鹤在各时段的警戒行为和修整行为比例均明显高于幼鹤,成幼鹤的休息行为比例在各时段差别不大,均在14:00—14:59有个明显的高峰。家庭群成鹤的觅食行为比例几乎在各时段均明显高于非家庭群,家庭群成鹤的修整行为和休息行为在大部分时段均低于非家庭群成鹤。因此家庭群成鹤采取多取食,少休息和修整的策略提高自身的适合度,同时保证对后代的抚育。     

云南高黎贡山赧亢白眉长臂猿春季食谱及活动时间分配初探

从2007年3月9日至4月14日,调查了高黎贡山赧亢的白眉长臂猿的食性和时间分配,旨在为该物种的栖息地保护和食物资源的管理提供指导.观察时,将白眉长臂猿的行为分为取食、移动、休息3种活动类型,利用瞬时扫描取样法取样记录,并记录取食种类及部位.白眉长臂猿春季取食植物种类共有10种,其中果实类植物3种,果汁类植物1种,嫩叶类植物6种.相对其它灵长类而言,其春季的食物多样性较低.白眉长臂猿移动、休息和取食的时间分配分别为33%、35%和32%,较叶猴属动物休息少,移动多.赧亢的白眉长臂猿1天中有3个觅食高峰期,9:00～10:00为第1个觅食高峰;11:00～13:00为第2个觅食高峰;16:00～17:00为第3个觅食高峰.而孟加拉国 Lawachara 的白眉长臂猿仅在早上有两次觅食高峰.这种差异可能是不同区域的白眉长臂猿对环境的适应结果.     

重庆金佛山国家级自然保护区黑叶猴春季食物选择

2008年3～5月,采用焦点猴群连续跟踪观察和记录法对金佛山黑叶猴的食性进行了研究.黑叶猴春季采食的植物主要有18种,分属8科17属.黑叶猴嗜食灯台树、香叶树、毛叶木姜子、山梨、构树和木姜子,喜食山樱桃、山桃、山桑和野桐.取食部位有叶、芽、嫩枝、花和果实.在日活动节律中,觅食行为有两个高峰期,分别是7:30～9:30和17:00～18:30.在黑叶猴每天的时间分配中,觅食所占的时间比例为21.56%±8.13%.     

夏季可可西里雌性藏原羚行为时间分配及活动节律

2005和2010两年的6—8月,在可可西里采用目标动物观察法记录藏原羚的行为时间分配和活动节律。所有行为被划分为觅食、警戒、躺卧、移动和其它5种类型。研究结果表明,觅食和躺卧是藏原羚的主要行为类型,分别占总行为时间的(42.02±2.22)%和(31.21±2.71)%,警戒、移动和其它3种行为类型时间比例依次减少。藏原羚的集群大小为1—10和12只,随着集群规模的增加,藏原羚的觅食时间增加,警戒时间减少,但其余3种行为类型与集群规模无显著相关性。与无羔藏原羚个体相比,有羔藏原羚的移动行为所占比例较高,而躺卧比例较低,觅食、警戒和其它3种行为类型差异不显著。一天中,藏原羚的觅食行为存在3个高峰,分别为8:00—8:59,10:00—12:59和16:00—19:59;躺卧行为的变化趋势与觅食行为相反;警戒行为仅有1个高峰,为14:00—14:59。5种行为类型在各时段间均存在显著差异。     

黑叶猴食物组成的地域性差异比较

研究同一个物种在不同生境觅食对策的差异不仅有助于了解其行为和生态的可塑性,还有助于评价其对自然或人为干扰导致的生境质量的改变的适应能力,从而为物种的有效保护和管理提供科学依据。本文以广西弄岗国家级自然保护区和扶绥保护区的黑叶猴为研究对象,采用瞬时扫描取样法收集相关的食性数据,并结合贵州麻阳河国家级自然保护区黑叶猴的食性研究结果,比较分析黑叶猴食物组成的区域性差异以探讨黑叶猴对不同喀斯特石山生境的行为适应对策。结果表明:黑叶猴3个地理种群共采食77科、182属、259种植物,木本植物(包括乔木和灌木)是其最重要的食物来源。3个地理种群均表现出较高程度的叶食性,树叶分别占其食物组成的52.7%、63.9%和79.7%。然而,3个地理种群的食物组成却存在显著差异,种群间总的食物重叠指数仅为1.33%-7.43%,主要食物的重叠指数仅为0-13.51%。这些差异可能与栖息地中植被组成的差异有关。与弄岗和麻阳河黑叶猴相比,扶绥黑叶猴采食更多的嫩叶和灌木。人为干扰对栖息地植被结构的影响可能是导致上述差异的主要原因。我们认为黑叶猴不同地理种群食物组成的显著差异反映其觅食的灵活性以应对不同喀斯特石山生境。     

海南热带雨林国家公园不同植被类型的大型真菌多样性

为揭示海南热带雨林国家公园大型真菌多样性及不同植被类型对真菌群落的影响, 本研究于2020年和2021年湿季对海南热带雨林国家公园内7个管理局辖区开展了大型真菌多样性调查, 比较了不同植被类型(山地雨林、低地雨林、低地雨林次生林、人工林)的大型真菌生活型(共生型、腐生型)组成差异。从设置的58条1 km长的样带内采集到1,869份子实体标本, 根据子实体形态与ITS rDNA序列分析, 从中鉴定出562种真菌, 涉及17目64科174属, 其中80%以上的物种由伞菌目、牛肝菌目、红菇目、多孔菌目、鸡油菌目、锈革孔菌目和炭角菌目构成。大型真菌的营养型以腐生型(占48.2%物种)和共生型(44.8%)为主。每条样带的平均物种丰富度和多度以中海拔的山地雨林最高, 分别为28 ± 5种和33 ± 6个, 而人工林最低, 分别为11 ± 1种和11 ± 2个。植被类型主要影响共生型大型真菌物种丰富度(P = 0.026)和子实体多度(P = 0.019)及Shannon-Wiener多样性(P = 0.028), 但对腐生型大型真菌的影响并不显著。多响应置换过程(multiple response permutation procedure, MRPP)检验结果表明, 不同植被类型对共生型与腐生型大型真菌群落物种组成均有显著影响(腐生型: P = 0.004, 共生型: P = 0.041)。冗余分析(redundancy analysis, RDA)的结果表明, 植被类型对腐生型和共生型真菌群落物种组成差异的解释度均较低(共生型: R² = 0.068, P = 0.004; 腐生型: R² = 0.067, P = 0.004)。海拔仅对腐生型真菌群落物种组成产生微弱影响(R² = 0.029, P = 0.001), 而对共生型真菌影响不显著(R² = 0.024, P = 0.072)。在不同保护地之间, 共生型(R² = 0.148, P = 0.001)与腐生型(R² = 0.123, P = 0.002)真菌物种组成均具显著差异; 基于样带‒真菌矩阵的网络图显示, 海南热带雨林国家公园内尖峰岭、霸王岭、五指山等国家级自然保护区的山地雨林是共生型大型真菌多样性较高区域, 应作为共生型真菌与宿主的优先保护区域。     

Feeding strategy of François' langur and white-headed langur at Fusui, China

We investigated the diet of a group of François' langur inhabiting a forest fragment between August 2002 and July 2003 to add to our knowledge of behavior and ecology of this langur. Our objective was to study whether the François' langur would adopt a feeding strategy similar to the white‐headed langur in the same nature reserve. Data indicated that a total of 37 plant species were eaten by François' langur; however, only ten species accounted for 90% of the total feeding time. Four species belong to the ten most dominant tree species within the vegetation quadrats. Ninety‐four percent of feeding time of François' langur was spent on leaves, whereas the remaining 6% of time was spent on fruits, flowers and twigs. Results also demonstrated that François' langur are similar to white‐headed langur in being predominantly folivorous and that Francois' langur adopted a similar strategy to white‐headed langur in diet, dietary variation and time spent on feeding. François' langur exhibited a habitat use preference for the middle zone of the forest, whereas the white‐headed langur utilized the bottom zone of the forest. Further analysis indicates that human interference and habitat fragmentation caused by agriculture also impact habitat use. Am. J. Primatol. 70:320–326, 2008. © 2008 Wiley‐Liss, Inc.     

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.     

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.     

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.     

Dietary breadth and resource use of François' langur in a seasonal and disturbed habitat

Previous studies of Trachypithecus species indicated that they were selective feeders that concentrated on relatively few food species/items. From January to December 2005, I quantified potential food availability and the food species/items eaten by five groups of François' langur (Trachypithecus francoisi) in the Mayanghe Nature Reserve (MNR), People's Republic of China. These langurs fed on 164 species, of which the top ten accounted for 51% of all feeding records. Langurs consumed more species (91) in the spring than in other seasons (73 summer, 75 autumn, and 67 winter), and only 38 species were consumed in all seasons. Nontree food species, such as bushes and lianas, accounted for 47% of the total feeding records and for a majority (68%) of the feeding records in winter. The annual diet consisted of leaves (64% of feeding records), fruit and seed (32%), and other nonfoliage items (4%); the langurs switched from being more folivorous in spring (93%) and summer (79%) to being more frugivorous in autumn (53%) and winter (56%). There was no correlation between the proportion of feeding records and the food availability in the most frequently consumed species, indicating that these langurs were selective feeders; there were significant correlations between consumption and abundance in both the entire set of 112 food species and the set of 86 infrequently consumed species, indicating that foods that are more available are eaten more frequently. It appeared that in the seasonal and disturbed habitat, feeding decisions and diet composition of the langur may be driven more by food availability, and less by animal's selectivity, than at other sites. The results indicate that François' langur copes with habitat alterations by broaden its dietary breadth; this has implications for the adaptive significance of dietary breadth, and has implications for future conservation strategies for species which exist in degraded habitats. Am. J. Primatol. 73:1176–1187, 2011. © 2011 Wiley Periodicals, Inc.     

Population census of the white-headed langur (Trachypithecus leucocephalus) at Longrui Karst Hills, Guangxi, China

A population census of white-headed langurs (Trachypithecus leucocephalus) was conducted using sleeping-site counts and the line-transect method from July to August 2002 at the Longrui Karst Hills, Guangxi, China. This area had been the largest habitat of the langurs across their range before the 1990s. While our survey revealed 146 old sleeping-sites, no white-headed langurs or fresh sleeping-sites were found in this area. Our study indicated that there had previously been a large population of langurs at Longrui Karst Hills, but now the langurs are possibly locally extinct in this area. If langurs still exist within the area, the population density must be very low. Interviews with local people confirmed that the number of white-headed langurs has been decreasing since the 1980s. Poaching was very common in the past and continues to be a problem at present, and it is likely that this has caused the decrease in the white-headed langur population at Longrui Karst Hills.     

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.     

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.