Diet of <Emphasis Type="Italic">Pygathrix nigripes</Emphasis> in Southern Vietnam

Black-shanked douc langurs (Pygathrix nigripes) are Southeast Asian colobines about which primatologists know very little, but they are classed as endangered because of population decline due to habitat loss. Two preliminary studies have shown that this monkey is primarily folivorous, but there are few details of plant selection or seasonal changes in diet. We set out to observe douc langurs directly in the wild to quantify the diet during wet and dry seasons. We confirmed that the species relies on foliage throughout the year, but it includes significant proportions of fruit and flowers in its diet when those items are available. The douc langurs selected various parts from 152 species of plants in 2 national parks, but there did not appear to be favorite species that were heavily selected over others. In both the wet and dry seasons, the black-shanked douc langurs ate mostly leaves, but the diversity of plants consumed increased and the proportion of fruit eaten almost doubled in the wet season. There were diurnal shifts in food selection, but we found no evidence that the douc langurs were foraging strategically to maximize their protein or energy intake.     

喀斯特石山生境中熊猴的雨季食物组成

开展食物组成研究为人们了解灵长类对栖息地的反应提供了很好的途径,对深刻理解动物的行为可塑性及适应性具有重要意义。猕猴属(Macaca)为果食性灵长类,但是不同种类的食性差异很大;即使便同一物种,其不同地理种群也因其栖息环境不同,食物组成存在差异。一般说来,热带地区的种类比生活在较高纬度的种类采食更多的果实。于2012年7—9月,采用瞬时扫描法对广西弄岗自然保护区石山中的两群熊猴(Macaca assamensis)进行了跟踪和观察,对猴群的雨季食物组成及其日时段变化规律进行了研究。结果表明:研究期间熊猴共采食45种植物,其中乔木30种,灌木3种,藤本11种,草本1种。平均每月采食植物22.3种。嫩叶和果实是熊猴的主要食物,分别占食物组成的52.4%和46.1%(其中未成熟果实占21.3%,成熟果实占24.8%)。另外,花占食物组成的0.9%,成熟叶和其它部位分别占0.3%。石山特有植物芸香竹(Bonia saxatilis)的嫩叶提供了43.8%的食物。9种主要植物分别占食物组成的2%,共为猴群提供了85.5%的食物。分析还发现熊猴并不是严格按照环境中的植物生物量来选择食物。嫩叶在弄岗熊猴的食物中的比例高于其他地理种群,而果实低于其他地理种群,这可能与喀斯特石山中果实的丰富度和可利用性较低有关。熊猴一天中不同时间段的食物组成并不相同,主要表现在:熊猴上午时间段对成熟果实和总果实的采食比例高于下午时间段,而嫩叶的采食比例低于下午时间段。另外,不同时间段的食物组成受外界温度的影响,表现为温度与嫩叶的觅食比例成正比,与成熟果实和总果实的觅食比例成反比。这可能与猴群采取的能量平衡策略有关。对熊猴的食物组成的日时段变化规律进行首次报道,研究结果将有助于深入理解熊猴对喀斯特石山生境的适应策略。     

The diet of chimpanzees in the Budongo Forest Reserve, Uganda

Chimpanzee diets are highly variable, but in all cases dominated by ripe fruit. Previous work has suggested that in the Budongo Forest Reserve, Uganda, ripe fruit remains available throughout the year, but dietary data for chimpanzees at this site remain sparse. This paper details the diet of a recently habituated community of Budongo chimpanzees living in the Sonso region. Chimpanzees of this community spent 64.5% of their feeding time eating fruit, and 19.7% eating arboreal leaves. Terrestrial herbaceous vegetation formed only a minor component of the diet. In these general dietary characteristics, Budongo chimpanzees show greater similarity to those of the Gombe National Park than they do to those in the geographically closer and floristically more similar Kibale Forest National Park. The diet was dominated by the fruit of four species and leaves of two species, although the composition of the diet varied from month to month, remaining diverse. Figs were consumed throughout much of the year, and in consequence, should perhaps be regarded as a staple, rather than fallback, food. A period of food scarcity was not apparent in this study and future comparative nutritional investigation will be required to determine whether these chimpanzees face a time of dietary hardship.     

Nutritional geometry: gorillas prioritize non-protein energy while consuming surplus protein

It is widely assumed that terrestrial food webs are built on a nitrogen-limited base and consequently herbivores must compensate through selection of high-protein foods and efficient nitrogen retention. Like many folivorous primates, gorillas' diet selection supports this assumption, as they apparently prefer protein-rich foods. Our study of mountain gorillas (Gorilla beringei) in Uganda revealed that, in some periods, carbohydrate-rich fruits displace a large portion of protein-rich leaves in their diet. We show that non-protein energy (NPE) intake was invariant throughout the year, whereas protein intake was substantially higher when leaves were the major portion of the diet. This pattern of macronutrient intake suggests that gorillas prioritize NPE and, to achieve this when leaves are the major dietary item, they over-eat protein. The concentrations of protein consumed in relation to energy when leaves were the major portion of the diet were close to the maximum recommended for humans and similar to high-protein human weight-loss diets. By contrast, the concentrations of protein in relation to energy when gorillas ate fruit-dominated diets were similar to those recommended for humans. Our results question the generality of nitrogen limitation in terrestrial herbivores and provide a fascinating contrast with human macronutrient intake.     

Diet and feeding patterns in the kipunji (<Emphasis Type="Italic">Rungwecebus kipunji</Emphasis>) in Tanzania’s Southern Highlands: a first analysis

The diet and feeding behaviour of the kipunji (Rungwecebus kipunji) was studied over 45 months, the first dietary analysis for this species. During 9498 h of direct observation of 34 kipunji groups, a list of 122 identified foodplants was recorded. The list represents 60 families, including 64 trees, 30 herbs, 9 climbers, 7 shrubs, 6 lianas, 3 grasses and 3 ferns. Kipunji were observed eating bark, young and mature leaves, ripe and unripe fruits, flowers, pith, seed pods, rhizomes, tubers, shoots and stalks. Invertebrates, fungi, moss, lichen, and soil were also eaten. Macaranga capensis var. capensis, an early successional tree, was the most commonly consumed species, with leaves, leaf stalks, pith, flowers and bark all eaten. We demonstrate that the kipunji is an omnivorous dietary generalist, favouring mature and immature leaves, ripe and unripe fruits and bark in similar proportions, with an almost comparable fondness for leaf stalks and flowers. Kipunji appear to be adaptable foragers able to modify their diet seasonally, being more folivorous in the dry season and more frugivorous in the wet. Whereas more ripe fruit is eaten in the wet season, the proportion of unripe fruit remains similar across the year. The proportion of mature leaves and pith increases throughout the dry season at the expense of ripe fruits and bark, and this may compensate nutritionally for the lack of available dry-season ripe fruits. Relatively more pith is eaten in the dry season, more stalks at the end of the dry and beginning of the wet seasons, and bark consumption increases as the rainfall rises.     

Leaf chemistry and the biomass of folivorous primates in tropical forests

Summary Recent results have suggested that the biomass of folivorous arboreal primates per unit area in Africa and Asia is positively correlated with the average quality of leaves, expressed as the ratio of protein to fiber concentrations in a given forest. This hypothesis has been tested in different forests of Madagascar. Leaf selection of all folivorous femus species was studied in relation to leaf chemistry. Except for two populations ofLepilemur subspecies all other folivorous lemur species (including two other subspecies ofLepilemur) select leaves with high concentrations of easily extractable protein or low concentrations of fiber, or both. This confirms the prominent role of these two components in leaf selection by folivorous lemurs. The average quality of mature leaves in a given forest, expressed as the ratio of protein to fiber concentrations, is positively correlated with the biomass of folivorous lemurs. This confirms the hypothesis tested and suggests rather uniform selection processes for arboreal folivorous primates across the world.     

Feeding patterns in a small-bodied nocturnal folivore (Avahi laniger) and the influence of leaf chemistry: a preliminary study

We present data on feeding patterns in Avahi laniger and compare these data to temporal variations in leaf chemistry. Because A. laniger is one of the smallest folivorous primates and has a monogastric stomach, we hypothesized that this lemur would display behavioural adaptations to a leaf-based diet by scheduling feeding times when leaves were of highest quality. Data were collected from May to August 2004 at the Vatoharanana site in Ranomafana National Park, Madagascar. A. laniger fed during different time periods despite leaf carbohydrate and protein concentrations exhibiting little variation throughout the night. Although tannin concentrations exhibited temporal fluctuations, they did not covary with A. laniger feeding times. We suggest that A. laniger feeding times cannot be explained entirely by variations in leaf chemistry.     

Feeding Patterns and Dietary Profile of Nocturnal Southern Woolly Lemurs (Avahi meridionalis) in Southeast Madagascar

Folivory has evolved in all primate radiations but the relative importance of resource quantity and nutritional quality of food is controversial. To understand how food abundance and different nutrients affect a folivorous diet, we investigated food composition of the leaf-eating lemur Avahi meridionalis. From June to December 2004, we conducted 26 nights of focal observation (256.1 h) in Sainte Luce pluvial littoral forest (southeast Madagascar) and recorded feeding behavior of 4 radiocollared individuals. Within the subjects’ home ranges, we recorded vegetation characteristics (morphospecies, phenology, diameter at breast height) and sampled food and nonfood items for chemical analyses. A. meridionalis did not eat fruit but only leaves and flowers and did not base their choice on food abundance. Adult leaves eaten were higher in easily soluble protein than adult leaves that were not consumed. The subjects ate young leaves and flowers as soon as they became available. These young leaves contained the same concentrations of soluble protein, higher concentrations of crude protein, and lower concentrations of acid detergent fiber and sugar than mature food leaves. A. meridionalis ate leaves with condensed tannins, alkaloids, and intermediate concentrations of polyphenolics. Contrary to previous studies that considered Avahi spp. a specialist, A. meridionalis acted as leaf-eating generalists with moderate selectivity, based on nutritional quality and tolerance of a wide array of plant secondary metabolites. This illustrates the dietary flexibility within a single genus of primates that seems to be driven by environmental constraints rather than morphological or physiological adaptations.     

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。     

Diet of the Delacour's langur (Trachypithecus delacouri) in Van Long Nature Reserve,Vietnam

The six limestone langur taxa of Southeast Asia inhabit the rugged limestone karst mountains, although the reason for their current restriction to this habitat is unclear. From August 2007 to July 2008, I collected data on the diet and feeding behavior of the critically endangered Delacour's langur (Trachypithecus delacouri) in Van Long Nature Reserve, Vietnam. I used these data to evaluate the hypothesis that limestone langurs are found on karst habitat because they depend on endemic limestone plants. Feeding accounted for 29% of the langurs' activity budget. Young leaves dominated the diet monthly, seasonally, and annually. The annual diet consists of nearly 79% foliage with almost 60% young leaves. Despite a distinct wet and dry season, over the study period, seasonal variation in plant part consumption was slight. Fruit and seeds were a small contribution to the diet. Delacour's langurs ate 42 of 145 available species, and they concentrated on a subset of this number. Five plant species comprised more than 60% of the diet and 16 species comprised more than 93%. More than half of the diet came from climbers. Delacour's langurs are among the most highly folivorous of studied colobines and, along with the closely related T. leucocephalus of southern China, the most folivorous of the Asian langurs. Whether high folivory is owing to a lack of available fruits and seeds in limestone habitats is unknown. What is certain, however, is that the plant species most important in the Delacour's langurs' diet at VLNR, throughout the study, were not plants endemic to limestone. Feeding dependence alone, therefore, cannot explain the current distribution of limestone langurs on karst habitat. Am. J. Primatol. 72:317–324, 2010. © 2009 Wiley‐Liss, Inc.     

Folivory by a tropical tanager: species of plants used and the relationship between leaf consumption and fruit abundance

ABSTRACT Folivory, or eating leaves, is unusual for small passerines. Puerto Rican Spindalises (Spindalis portoricensis, Thraupidae), tanagers endemic to the island of Puerto Rico, are known to feed on leaves, but little is known about the possible importance of leaves in their diet. Our objectives were to determine the different species of leaves eaten, the percentage of their diet consisting of leaves, and the relationship between leaf consumption and fruit abundance. We used data from previous studies where the foraging activity of Puerto Rican Spindalises was systematically sampled, and from additional observations made throughout the island. We documented 160 records of folivory, with spindalises feeding on 44 plant species in 25 plant families, including monocots, dicots, gymnosperms, and pteridophytes. Spindalises fed on young leaves of 26 plant species, and mature leaves of 19 plant species. Spindalises were primarly frugivorous (83.9% of diet), but leaves were the second most frequent food item in their diets (8.9% of diet). We also found that leaf consumption was negatively correlated with the abundance of ripe fruit, suggesting that leaves were particularly important food items when less fruit was available. The frequency of folivory by spindalises in our study was less than reported for other folivorous passerines such as plantcutters (Phytotoma spp.) and saltators (Saltator spp.). Nonetheless, folivory may help spindalises cope with human‐dominated landscapes and other environmental changes on small islands.     

Diet of the Indochinese silvered langur (Trachypithecus germaini) in Kien Luong Karst area,Kien Giang Province

The Indochinese silvered langur (Trachypithecus germaini) is distributed to the west of Mekong River in Cambodia, Lao PDR, Thailand and Vietnam. During a two‐year study, from May 2014 to May 2016, we collected 320.44 hr of behavior, with 17,040 feeding bouts recorded (142 hr) for T. germaini on Chua Hang Karst Mountain, Kien Luong District, Kien Giang Province, Vietnam. Feeding accounted for 45% of the Indochinese silvered langurs’ activity budget. The plant diet of the Indochinese silvered langurs was principally composed of young leaves (58%), followed by mature leaves (9.5%), fruits (22.7%), flowers (4.7%), buds (3.3%), petioles (1.2%), and other (0.5%). A total of 58 plant species were fed on by the silvered langurs, and leaves of eight species (Phyllathus reticulatus, Ficus rumphii, Ficus tinctoria, Ficus microcarpa, Cayratia trifolia, Streblus ilicifolia, Combretum latifolium, and Streblus asper) were fed on throughout the year. P. reticulatus was most frequently eaten (13.9% feeding time, n = 1,733). Food selection differed significantly between months and seasons. The Indochinese silvered langurs ate 27 plant species in the wet season compared with 23 plant species in the dry season. Leaf chemical composition of two food categories, 16 eaten species (with 10 most frequently consumed species and six least consumed species), and four noneaten species, were analyzed. Feeding samples from eaten species in the Indochinese silvered langurs's diet contained lower amounts of condensed tannin, lignin, protein, ash, and lipids, but a higher amount of total sugar compared with samples from noneaten species. Furthermore, the most frequently consumed species contained lower amounts of lignin compared with the less frequently consumed species. Using a generalized linear model with five variables, including neutral detergent fiber (NDF), total sugar, lignin, lipid, and calcium (Ca) indicated that NDF positively correlated and lignin content negatively correlated with feeding records in the diet of these langur.     

Digestive strategies and food choice in mantled howler monkeys Alouatta palliata mexicana: bases of their dietary flexibility

Mantled howler monkeys (Alouatta palliata) occupy a wide variety of tropical habitats and are the most folivorous of New World primates. However, their diet may include fruits, buds, petioles, and flowers, as well as leaves, suggesting they must cope with variations in the nutrient composition of their food. We studied the physiological basis of the dietary flexibility of these monkeys by comparing food choice, digestive performance and patterns of digesta flow in six adults, fed diets of either leaves or a mixture of fruit and leaves. Although monkeys ate similar amounts of the two diets, they ingested more digestible protein when offered the leaf diet, on which they lost body mass, but they ingested much more soluble sugars when offered fruit and leaves on which they gained mass. Digestibilities of dry matter, fat, energy and fibre did not differ between diets, but those of crude protein, soluble sugars and minerals were higher on the fruit–leaf diet. Mean retention times in the gut of solute (Co-EDTA) and particulate markers (Cr-mordanted cell walls) did not differ between diets, but on both diets the monkeys retained the particulate marker (mean retention time ca 55 h) for longer than they did the solute marker (MRT ca 50 h). A lack of selective retention of solutes and small particles in the gastro-intestinal tract of howler monkeys probably restricts them to mixed diets but their digestive strategy is sufficiently flexible to allow them to feed on a diet of leaves when fruit is unavailable.     

Variation in total phenols and condensed tannins in Eucalyptus: leaf phenology and insect grazing

Total phenols and condensed tannins in leaves of seven species of Eucalyptus ranged from 4–40% and 0–27% respectively of the leaf dry weight. The concentrations of these compounds were variable but usually high in young and older leaves throughout the growing season, and typically increased during winter, but no other trends with season or leafage were apparent. This pattern of seasonal variation in concentrations of total phenols and condensed tannins is different to that studied for other plant species. This difference may be related to repeated production of new leaves by Eucalyptus during the growing season, and the probability that these leaves will be attacked by herbivorous insects.     

Feeding Ecology of Guerezas in the Kakamega Forest,Kenya: The Importance of Moraceae Fruit in Their Diet

Eastern black-and-white colobus (Colobus guereza), or guerezas, have long been considered to be one of the most folivorous primates. I conducted a study of the feeding ecology of two guereza groups (T and O) over an annual cycle in the Kakamega Forest of western Kenya. I found that the annual diets of both groups comprised mostly of leaves (T: 48%, O: 57%) though fruit (T: 44%, O: 33%) also accounted for a substantial portion of the diet. In the six months when fruit was most abundant, fruit consumption constituted an average of 58% of T-group's monthly diet and 42% of O-group's monthly diet. In contrast to most previous studies of colobines, in which seeds were the primary fruit item consumed, almost all of the fruit eaten by guerezas at Kakamega consisted of whole fruits. At least 72% of the whole fruits consumed by T-and O-groups were whole fruits from trees in the Moraceae family, which dominates the tree family biomass at Kakamega. Unlike at sites where guerezas consumed fruit primarily when young leaves were scarce, at Kakamega guerezas ate fruit in accordance with its availability and irrespective of the availability of young leaves. My findings demonstrate that guerezas are more dietarily flexible than was previously known, which may help to explain why the species can survive in such a wide variety of forested habitats across equatorial Africa.     

Summer feeding strategy of Spanish ibexCapra pyrenaica and domestic sheepOvis aries in south-eastern Spain

The feeding strategies of the Spanish ibexCapra pyrenaica Schinz, 1838 and sheepOvis aries Linnaeus, 1758 during summer in the alpine area of Sierra Nevada in south-eastern Spain were compared. The degree of vegetation use by the two ungulates, food availability, diet composition, species selection indices and the effect of plant availability and chemical composition on diet selection were analysed. The available biomass of herbaceous and woody resources was 87 and 13%, respectively. The degree of use was 15% for herbaceous vegetation and 10.3% for woody vegetation. The most used species were those with moderate amounts in diet composition of both ruminants. The main plant groups consumed by Spanish ibex were graminoids (57.5%), shrubs (20.5%) and chamaephytes (14.3%). Sheep consumed graminoids (41.6%), forbs (32.3%) and chamaephytes (20.9%). There was a significant correlation between the two diets, with an overlap of 56.3%. The highest indices of selection were found in items with moderate-low availability and which comprised moderate levels in the diet composition of the two species. Spanish ibex mainly selected diet according to availability, while sheep selection depended on protein content. The ibex consumed cell wall-rich resources such as grasses and shrubs, while the sheep consumed higher quality feed and includes more forbs in its diet.     

Selective herbivory by brushtail possums: Determining the age of ingested leaves using n‐alkanes

Abstract Leaves often decline in nutritional quality as they age, and selective feeding on young leaves may be nutritionally important for herbivores. Preference by mammalian herbivores for young leaves has rarely been measured in the field owing to technical difficulties. We measured preferences with respect to leaf age of an arboreal folivore, the brushtail possum (Trichosurus vulpecula Kerr), feeding on southern rata (Metrosideros umbellata Cav.; Myrtaceae) in a new application of the alkane technique. We characterized the cuticle waxes (n‐alkanes) of rata leaves that were less than 1 year old (‘1‐year’), between 1 and 2 years (‘2‐year’) and greater than 2 years old (‘>2‐year’). Simulations showed that the method accurately discriminated between 1‐year and other age groups but slightly overestimated the importance of rare components of the diet. This bias was larger when discriminating between 2‐year and >2‐year leaves apparently because they had more‐similar alkane profiles. Metrosideros umbellata leaf formed 20.8% of the diet of a population of possums from Rakiura, New Zealand, sampled in autumn 2002 (n = 33). Of the M. umbellata component, alkane analyses showed that 1‐year leaves formed 88.7 ± 3.9% of the diet despite making up only 39.5 ± 2.2% of the leaf biomass on rata trees (n = 14). The foliar concentrations of the macronutrients N, P and K all declined significantly with leaf age (P < 0.0001). Lignin content did not measurably increase with leaf age, suggesting that digestibility per se did not determine the preference of brushtail possums for young rata leaves. This study provides the first quantitative evidence that possums discriminate by leaf age and that the resulting diet is enriched in macronutrients.     

Similarities,differences, and seasonal patterns in the diets ofEulemur rubriventer andEulemur fulvus rufus in the Ranomafana National Park,Madagascar

I studied the diets of two sympatric species ofEulemur (E. fulvus rufus andE. rubriventer) in the Ranomafana National Part in southeastern Madagascar from July 1988 through August 1989. Both species were highly frugivorous throughout the year and devoted similar amounts of time to feeding daily; the composition of their diets were similar. Three aspects of both lemur species' diets were correlated with seasonal food availability: changes in interspecific dietary overlap, the length of feeding bouts, and the amount of time devoted to feeding daily. Throughout the year neither species had a more diverse or more folivorous diet than its sympatric congener. However, there were several consistent year-round differences between their diets.E. f. rufus initiated more feeding bouts daily that were shorter in duration than those ofE. rubriventer. E. f. rufus also consistently exploited more unripe fruits and mature leaves thanE. rubriventer did throughout the study.E. f. rufus migrated from the study site during a period of fruit scarcity. These dietary patterns are discussed here in regard to feeding patterns observed in other sympatric congeneric primate species, whether these dietary differences can contribute to these two species' ability to coexist, and how differences in diet may be influenced by differences in social structure. I suggest that subtle, consistent differences in diet and seasonal dietary differences are sufficient to allow these two species to coexist.     

Nutritional ecology of Preuss's red colobus monkey (Colobus badius preussi Rahm 1970) in Korup National Park, Cameroon

Studies on the nutritive ecology of red colobus monkeys (Colobus badius preussi) in Korup National Park, Cameroon, investigated food plants and the parts eaten as well as nutritive values of the various food items. Food plants were identified through direct observations and collected for laboratory determination of nutrient content. Food plants were analysed for crude protein (CP), crude fibre (CF), ether extract (EE), nitrogen‐free‐extractive (NFE) and total ash (A). A total of seventeen species belonging to nine different families were identified as food plants for red colobus. Food and feeding habits showed that the species were highly folivorous, tending to prefer the young leaves of food plants. Lecomtedoxa klaineana (27.0%) and Xylopia aethiopica (22.2%) of families Sapotaceae and Annonaceae, respectively, constituted about 50% of total food items consumed. Results of nutrient composition indicated individual nutrients varied significantly (P < 0.01) with food species.