黑叶猴食物组成及其季节性变化

对灵长类来说,喀斯特石山生境是一种比较独特的生活环境。喀斯特森林具有生物量低,物种多样性高的特点,生活于其中的灵长类的食物资源受环境的影响强烈,有明显的季节性变化。黑叶猴(Trachypithecus francoisi)是生活在喀斯特石山的特有灵长类动物。在食物资源波动较大的石山生境中,黑叶猴如何在食物匮乏时期获得食物?是否采取了大多数灵长类的应对策略,还是采用适应喀斯特环境的特殊对策?为了回答这些问题,2005年9月至2006年8月,在广西弄岗国家级自然保护区选择一群黑叶猴作为观察对象,采用瞬时扫描取样法收集相关的食性数据。通过比较不同季节食物组成的差异来探讨黑叶猴如何应对喜食食物(嫩叶、果实)的季节性缺乏的问题。结果表明:黑叶猴共采食92种植物,其中乔木和灌木38种,藤本植物52种,草本植物1种,寄生植物1种。猴群平均每月采食食物22.8种(6-34种),月平均食物多样性指数为2.5(1.6-3)。在弄岗黑叶猴的食物组成中,树叶占总觅食记录的71%,其中嫩叶46.9%,成熟叶24.1%的;果实、花和种子分别占13.2%、6.3%和4.3%;其他食物类型占5.4%,分别包括茎1.8%,叶柄1.1%,未知部位2.5%。黑叶猴的食物组成具有明显的季节性差异。在旱季,平均月食物种类和食物多样性指数分别为29.3和2.8,明显高于雨季的16.2和2.1。在雨季,嫩叶在食物中的比例明显高于旱季(61.5%相对32.3%);在旱季,猴群对成熟叶的采食比例明显高于雨季(39.4%相对8.9%),且在该季节才利用种子作为食物,占食物组成的8.6%。黑叶猴的食物组成受食物可获得性的影响。主要表现为嫩叶在食物中所占的比例与其可获得性有显著的正相关关系(r=0.865,n=12,P=0.000)。另外,当成熟叶和果实的可利用性降低时,黑叶猴的食物多样性指数提高(成熟叶:r=-0.602,n=12,P=0.039;果实:r=-0.716,n=12,P=0.009)。黑叶猴在嫩叶丰富的雨季,它们采食更多的嫩叶;而在喜食性食物短缺的旱季,它们采取选择更多种类的成熟叶和种子为食,同时它们采食的种类和食物多样性也相应地增加。这与大多数叶食性灵长类的适应策略相似。     

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。     

同域分布黑叶猴和熊猴的活动时间分配比较

2005年9月至2006年8月,在广西弄岗国家级自然保护区选择一群黑叶猴和两群熊猴作为观察对象,采用瞬时扫描取样法收集相关的行为数据.通过比较两种灵长类活动时间分配的数据来探讨食物资源在时间和空间分布上的差异对两种灵长类动物活动时间分配的影响.结果表明:休息在黑叶猴的活动时间分配中所占比例明显高于熊猴; 熊猴用于移动的时间比例明显高于黑叶猴.两种灵长类采取不同的策略以应对喜食食物的季节性短缺.当旱季嫩叶和果实的可获得性降低时,黑叶猴明显增加用于移动和觅食的时间比例,相应减少用于休息的时间.与此相反,熊猴明显减少用于移动的时间比例,相应地增加用于社会性理毛的时间比例.分析表明,食物的可获得性以及食物组成的差异可能是影响同域黑叶猴和熊猴活动时间分配差异的重要因素.     

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。     

川金丝猴食谱的地域性差异比较

对不同地区川金丝猴食物组成的比较有助于了解其对不同生境食物供应的适应性。本研究通过对陕西川金丝猴猴群食谱的长期记录,并汇集了国内对四川-甘肃和湖北的川金丝猴食性研究结果,集成了这三个地理种群川金丝猴各自的地域性食谱,共计有136 种植物(隶属35 科)被该物种作为采食对象。对来自这三个不同区域(陕西、四川-甘肃、湖北)的川金丝猴食谱组成的比较,发现有近半数的植物是三个地理种群共同的采食对象,但其食谱组成差异明显。这可能源于各地植物本身的多样性差异,及不同地理种群对各种食物采食偏好的不同所致。对其相关聚类分析结果显示,陕西和湖北的猴群在食谱组成上相近,但四川-甘肃的猴群与前两个地区猴群的食谱组成差异极大。然而湖北和四川-甘肃种群在食物的选择上采用了近乎相同的偏好倾向,而陕西的猴群与它们明显不同。我们初步分析认为造成食谱组成和采食偏好差异的原因可能是各地理种群活动地海拔带重叠度不同、森林类型不同、它们在不同林型中活动的时间分配不一。很明显,就我们目前所掌握的有关川金丝猴食谱组成来看,该物种不应该仅仅被认为是一个叶食性灵长类动物,而应该是一个泛化采食者。     

广西弄岗黑叶猴栖息地选择与利用

2005年9月至2006年8月对广西弄岗国家自然保护区内一群黑叶猴进行行为观察,采用瞬时扫描法观察并收集猴群的栖息地利用数据,分析黑叶猴对喀斯特石山栖息地的选择与利用规律。结果表明,黑叶猴对不同山体部位的利用有显著性差异(χ~2=43.063,df=4,P0.001)。黑叶猴对崖壁的利用频率最高(占总记录的36.67%±9.44%),其次是山坡(32.30%±9.57%),对山脚(14.15%±5.01%)、山顶(11.24%±8.42%)和平地(5.63%±2.92%)的利用较少。分析发现,黑叶猴对山体部位的利用没有显著的季节性差异(山顶:Z=-0.160,P=0.837;崖壁:Z=-0.320,P=0.749;山坡:Z=-0.480,P=0.631;山脚:Z=-1.601,P=0.109;平地:Z=0,P=1)。黑叶猴将崖壁作为主要的休息场所,山坡和山脚为主要的移动和觅食场所。黑叶猴对栖息地的利用受食物可获得性的影响。当食物中花的可获得性降低时,猴群增加对山脚的利用;当嫩叶可获得性降低时,猴群增加在崖壁移动的频率;当果实的可获得性升高时,猴群增加在平地觅食的频率。食物组成与黑叶猴栖息地利用也有关系。总体来看,黑叶猴对山顶的利用频率与花的觅食比例呈显著正相关;对山脚的利用频率与果实+种子的觅食比例呈显著正相关。猴群在山顶休息的频率随花和成熟叶的觅食比例的上升而上升;在山脚休息的频率随果实+种子的觅食比例的上升而上升。猴群在山坡觅食的频率随果实+种子的觅食比例的下降而上升;在山脚觅食的频率随嫩叶的觅食比例的下降而上升。另外,平均最低温度与猴群在平地觅食的频率呈负相关关系。分析表明食物的分布和数量对黑叶猴栖息地利用有重要影响,黑叶猴对栖息地的选择是在觅食利益与风险之间进行权衡的结果。     

秦岭南坡川金丝猴的食物组成及季节性变化

2013年3月至2014年2月,在秦岭南坡观音山自然保护区大坪峪颜家沟内选取一群半野生川金丝猴的成年个体作为研究对象,采用瞬时扫描取样法收集其觅食的食物类型数据,目的是为该猴群建立食谱,并通过对比不同季节内觅取的食物组成差异探讨秦岭南坡川金丝猴如何应对喜食食物的季节性缺乏。结果表明：秦岭南坡川金丝猴共采食53种植物（包括34种乔木、13种灌木、6种藤本植物）和4种大型真菌,分别占取食植物组成的64.3%、25.3%、8.0%和2.2%,春、夏、秋、冬季节取食种类和多样性指数分别为20种(3.93)、19种(3.73)、21种(3.87)和25种(4.12)。在秦岭南坡川金丝猴的食物组成中,地衣占总觅食记录的22%;树叶占20%,其中嫩叶9%和成熟叶11%;种子、树皮、芽苞、果实和叶柄分别占16%、15%、11%、 9%和7%。该猴群觅取的植物部位具有明显的季节性差异。春季,对树皮和芽苞的觅食量较高,分别为28%和25%;夏季,增加了对成熟叶的采食量(29%),而减少了芽苞的觅取量(5%);秋季,以取食种子和果实为主,分别占总觅食的48% 和16%;冬季,地衣的采食量达到最大值（41%）。觅食的食物组成与食物的可获得性呈正相关性(R = 0.984, P < 0.01),这与大多数叶猴的适应策略类似,在喜食食物短缺的冬季,它们选择更多的地衣和树皮为食,同时它们采食的种类和食物多样性也有相应增加。     

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。     

黑叶猴在喀斯特石山生境的觅食活动

2002 年 8 月至2003 年 7 月,对一群生活在广西扶绥珍稀动物保护区内孤立石山中的黑叶猴进行观察,通过收集有关栖息地利用的数据,从中探讨栖息地的破碎化对其生态习性的影响.研究结果表明：猴群的觅食活动在石山各个部位的分布存在着显著差异,它们更偏好在山腰觅食（66.1%）,而较少利用山脚（18.6%）和山顶（15.3%）.从横向水平来看,猴群的觅食活动主要集中在6个觅食斑块,总面积为7.94 hm^2,约占栖息地面积的18.9%.觅食斑快中黑叶猴喜食食物的植株密度明显高于非觅食斑块.虽然树叶在黑叶猴食物组成中占绝大部分,但其食物组成具有明显的季节性变化.与之相对应,猴群对觅食斑块的利用也表现出明显的季节性差异.     

蒙古野驴的秋季食性分析

2006 年9 月在新疆卡拉麦里山有蹄类野生动物保护区内采集了25 堆蒙古野驴粪便和10 科共29 种植物标本,采用粪便显微组织学分析方法分析了蒙古野驴的秋季食性,并初步探讨了其食性与周围环境的关系。结果表明,蒙古野驴秋季食性广, 共采食8 科26 种植物。这些植物按照其在食物组成中的比例可分为3 大类:主要食物为梭梭、针茅、驼绒藜, 共占食物组成的61.3% ;常采食植物为柽柳、角果藜、蒿、琵琶柴、芨芨草等11种,共占食物组成的33.3%; 少见采食的植物为盐爪爪、獐毛、顶羽菊、黑果枸杞、里海盐爪爪等12 种,共占食物组成的6.7% 。按科别而论,蒙古野驴采食最多的是藜科植物,其次是禾本科植物。从被采食植物的分布区域分析,其采食区域广阔,包括了保护区内所有的生境类型。      

Nutrient contents predict the bamboo‐leaf‐based diet of Assamese macaques living in limestone forests of southwest Guangxi,China

Determining the nutrient factors influencing food choice provides important insight into the feeding strategy of animals, which is crucial for understanding their behavioral response to environmental changes. A bamboo‐leaf‐based diet is rare among mammals. Animals’ food choice and nutritional goals have been explained by several frameworks; however, the influence of nutrients on food choice in bamboo‐leaf‐based macaques is not yet available. Assamese macaques (Macaca assamensis) inhabiting limestone forests are characterized by such a bamboo‐leaf‐based diet, predominantly consuming young leaves of Bonia saxatilis, a shrubby, karst‐endemic bamboo. We studied the feeding behavior of one group of Assamese macaques using instantaneous scan sampling in limestone forests of the Guangxi Nonggang National Nature Reserve in southwest Guangxi, China. We compared the nutrient content of staple food and nonfood items and examine the role of key nutrients in the food selection of macaques. Our results showed that young leaves of bamboo B. saxatilis contained more water, crude protein, phosphorus, and less tannin than nonfood items. Furthermore, staple foods contained a higher content of water and less content of calcium than nonfood items. More specifically, quantities of water, crude protein, calcium, and phosphorus in food items were critical factors affecting feeding time on a specific plant item. Our results suggest that young bamboo leaves could meet macaques’ required protein and water intake, while enabling them to maintain their mineral balance, consequently facilitating to maintain the primates’ bamboo‐leaf‐diet in the limestone forest. Our findings confirm the effects of nutrient contents in food choice of Assamese macaques, highlighting the importance of the nutrient contents in maintaining their bamboo‐based diet and the need to increase the knowledge on their nutritional strategy adapted to the bamboo‐dominated diet inhabiting the unique limestone habitat.     

Food Habits of Formosan Rock Macaques (Macaca cyclopis) in Jentse,Northeastern Taiwan,Assessed by Fecal Analysis and Behavioral Observation

We studied the food habits of the Formosan rock macaques (Macaca cyclopis) in Jentse via fecal analysis and direct field observation from October 1991 to September 1992, and recorded macaques eating 51 plant species and insects of 5 orders. Macaques in Jentse spent more time feeding on fruits than on other plant parts or insects. However, there was seasonal variation in their food habits, i.e., they spent a higher proportion of time feeding on fruits and insects in summer, and on leaves and stems in winter. The major plant species consumed and the patterns of seasonal variation in the macaque diets identified by fecal analysis and field observation are similar. However, there are discrepancies between results from the two methods in some of the finer details of estimated dietary composition.     

Spatial patterns in the diet of the Japanese macaque Macaca fuscata and their environmental determinants

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Variations in diet composition of sympatric Trachypithecus francoisi and Macaca assamensis in the limestone habitats of Nonggang,China

Comparative studies of sympatric species are essential for understanding behavioral and ecological adaptation as well as the mechanisms that can reduce resource competition to allow coexistence.Fran?ois' langurs(Trachypithecus francoisi) and Assamese macaques(Macaca assamensis) are sympatric primate species found in the limestone seasonal rainforests of Nonggang Nature Reserve,southwestern Guangxi, China. To explore their different adaptation strategies, we collected data on diet using scan sampling at 15-min intervals. Our results revealed that Fran?ois' langurs showed a more flexible diet composition than Assamese macaques.Fran?ois' langurs increased dietary diversity and mature leaf consumption in response to seasonal scarcity of preferred young leaves and fruits, whereas Assamese macaques relied heavily on young bamboo leaves(Indocalamus calcicolus) in most months.These variations reflect the differences in digestive physiology, morphology, and the temporal and spatial distribution of food resources.     

Diet of Grauer's Gorillas in the Montane Forest of Kahuzi,Democratic Republic of Congo

We describe the diet of a semihabituated group of Grauer's gorillas (Gorilla beringei graueri) inhabiting the montane forest of Kahuzi-Biega National Park, Democratic Republic of Congo, based on direct observations, feeding remains in their fresh trails, and fecal samples collected over 9 yr. We examined fruit availability in their habitat; consumption of fruit, vegetative, and animal food; and daily intake of vegetative plant food using a transect, fruit monitoring trails, fecal analysis, and tracing of the animal's daily trails between consecutive nest sites. The fruit food repertoire of Kahuzi gorillas resembles that of western and eastern lowland gorillas inhabiting lowland tropical forests, while their vegetative food repertoire resembles that of mountain gorillas inhabiting montane forests. Among 236 plant foods (116 species), leaves, pith, and barks constitute the major parts (70.2%), with fruit making up the minor part (19.7%). About half (53.2%) of the total fecal samples included fruit remains. The gorillas used leaves, stems and other vegetative plant parts as staples. Their fruit intake was similar to that reported for mountain gorillas in Bwindi. They ate animal foods, including earthworms, on rare occasions. Variation in fruit consumption was positively associated with variation in fruit production. The gorillas ate fig fruits frequently; fig intake is positively correlated with that of other fruits, and figs were not fallback foods. They relied heavily on bamboo shoots on a seasonal basis; however, no bamboo shoots were available for several years after a major flowering event. Our results support the argument that variation in gorilla diets mostly reflects variation in vegetational composition of their habitats.     

Niche separation of sympatric macaques, Macaca assamensis and M. mulatta, in limestone habitats of Nonggang, China

Comparative studies of sympatric species are essential in understanding those species’ behavioral and ecological adaptations as well as the mechanisms that can reduce resource competition enough to allow coexistence. We collected data on diet, activity budget and habitat use from two sympatric macaque species, the Assamese macaque (Macaca assamensis) and the rhesus macaque (M. mulatta), in a limestone seasonal rainforest of Nonggang Nature Reserve, southwestern Guangxi, China. Our results show that the two sympatric macaques differ in diet, activity budget, and habitat use: (1) out of the 131 plant species that were used by both macaque species as food over the year, only 15 plant species (11 %) were shared. Rhesus macaques used more plant species as major foods, and had higher dietary diversity and evenness indexes than Assamese macaques. (2) Assamese macaques fed predominantly on leaves, whereas rhesus macaques fed more selectively on fruits. The rhesus macaques’ diet varied according to season, and was significantly correlated to season fluctuation in fruit availability. (3) Assamese macaques devoted more time to resting, and less time to feeding than rhesus macaques (4) Assamese macaques were present mostly on the cliff, and tended to stay on the ground, whereas rhesus macaques were present mostly on the hillside, and showed preference to lower and middle canopy. The observed differences in diet and habitat use between the two macaque species represent behavioral patterns enabling their coexistence.     

The diet and feeding behavior of the silver leaf monkey (Trachypithecus auratus sondaicus) in Indonesia

I describe the diet and feeding behavior of silver leaf monkeys (Trachypithecus auratus sondaicus) in the Pangandaran Nature Reserve, West Java, Indonesia, and compare a group living in old secondary rain forest with a group living in mixed plantation/secondary forest to determine intraspecific variation in feeding behavior and the importance of the plantation species in the diet of the monkeys. Young leaves and leaf buds made up slightly less than half of their diets, with both groups showing a preference for a few species when seasonally available. Fruits and flowers of a few species were also preferentially selected when available. These included sweet, fleshy fruits, which most other colobines tend to avoid. Young leaf intake was greatest in months when fruit intake was low. Mature leaves were rarely eaten. Both groups spent approximately 20% of feeding time foraging on Moraceae species. Differences in the diet of the two groups were related largely to differences in vegetational composition and the availability and abundance of food items for the species common to both sites. Teak (Tectona grandis) was the top food species of the group living in mixed plantation/secondary forest, with the midribs of young leaves preferentially selected. Young leaves ofT. grandis, available throughout the study, provided a staple food and were eaten when preferred foods were scarce. More favored food items were available to the group living in old secondary forest, though none was a staple food.     

Altitudinal and seasonal variations in the diet of Japanese macaques in Yakushima

Altitudinal and seasonal variations in the diet of Japanese macaques in Yakushima, southwestern Japan, were studied for 2 years by means of fecal analysis. The altitudinal range of fecal samples collected was 30 m to 1,203 m above sea level, and it was divided into three zones: low-zone forest (0–399 m), middle-zone forest (400–799 m), and high-zone forest (800 m–1,230 m). There was a considerable altitudinal and seasonal variation in the macaques' diet. Seed/fruit and animal matter were eaten more in the lower zones, whereas more fiber and fungi were consumed in the higher zones. In all of the zones, they ate seed/fruits the most in autumn (September–November) and the least in spring (March–April). They ate fibrous food the most in spring and the least in autumn. Macaques relied on seed/fruits heavily in the lower zone for a longer period than in the higher zones. Macaques in the high-zone forest ate almost no seed/fruit foods from March to May. Altitudinal variations in availability of seed/fruit foods seem to have influenced the altitudinal variations in diet. Total basal area of seed/fruit-food trees, species richness of seed/fruit-foods, main seed/fruit-food types available, and annual fleshy-fruit production all decreased with increasing altitude. Both interannual variation and annual cyclicity of diet were found in all zones. Electronic Publication     

Northern pigtailed macaques rely on old growth plantations to offset low fruit availability in a degraded forest fragment

Space-use and foraging strategies are important facets to consider in regard to the ecology and conservation of primates. For this study, we documented movement, ranging, and foraging patterns of northern pigtailed macaques (Macaca leonina) for 14 months in a degraded habitat with old growth Acacia and Eucalyptus plantations at the Sakaerat Biosphere Reserve in northeastern Thailand. We used hidden Markov models and characteristic hull polygons to analyze these patterns in regard to fruit availability. Macaques' home range (HR) was 599 ha and spanned through a natural dry-evergreen forest (DEF), and plantation forest. Our results showed that active foraging increased with higher fruit availability in DEF. Macaques changed to a less continuous behavioral state during periods of lower fruit availability in DEF, repeatedly moving from foraging to transiting behavior, while extending their HR further into plantation forest and surrounding edge areas. Concomitantly, macaques shifted their diet from fleshy to dry fruit such as the introduced Acacia species. Our results showed that the diet and movement ecology adaptations of northern pigtailed macaques were largely dependent on availability of native fruits, and reflected a “high-cost, high-yield” foraging strategy when fresh food was scarce and dry fruit was available in plantation forest. Conversely, wild-feeding northern pigtailed macaque populations inhabiting pristine habitat approached a “low-cost, low-yield” foraging strategy. Our results outline the effects of habitat degradation on foraging strategies and show how a flexible species can cope with its nutritional requirements.     

广西西南部熊猴对石山森林的利用

栖息地利用直接反映灵长类适应环境或应对环境变化所采取的行为策略,是了解和评估物种适应能力与进化潜力的重要途径.石山森林是一类特殊的生境,栖息在石山森林中的灵长类动物可能展现出独特的适应策略.为了解熊猴(Macaca assamensis)对石山森林的利用策略,2012年9月至2013年8月采用瞬时扫描法对广西弄岗国家级...