Diet of the red howler monkey (Alouatta seniculus) in French Guiana

We report the feeding behavior and food preferences of a troop of red howler monkeys (Alouatta seniculus) over two annual cycles in primary tropical rain forest in French Guiana. The monkeys used 195 plant species from 47 families as food. Major food categories were young leaves (54%), mature fruits (21.5%), and flowers (12.6%). Other food categories included old leaves, immature fruits, termitarium soil, bark, and moss. The monkeys were less selective than other howler groups, since 19 plant species contributed 1% to their diet and accounted for only 35.7% of their total diet. The Sapotaceae was the most frequently eaten plant family and represented >10% of the total diet.     

Fallback Foods of Red Leaf Monkeys (Presbytis rubicunda) in Danum Valley, Borneo

Animals in Southeast Asia must cope with long periods of fruit scarcity of unpredictable duration between irregular mast fruiting events. Long-term data are necessary to examine the effect of mast fruiting on diet, and particularly on the selection of fallback foods during periods of fruit scarcity. No such data is available for colobine monkeys, which may consume substantial amounts of fruits and seeds when available. We studied the diet of red leaf monkeys (Presbytis rubicunda, Colobinae) in Danum Valley, Sabah, northern Borneo, using 25 mo of behavioral observation, phenology and vegetation surveys, and chemical analysis to compare leaves eaten with nonfood leaves. The monkeys spent 46% of their feeding time on young leaves, 38% on seeds, 12% on whole fruits, 2.0% on flowers, 1.0% on bark, and 1.2% on pith. They spent more time feeding on seeds and whole fruit when fruit availability was high and fed on young leaves of Spatholobus macropterus (liana, Leguminosae) as fallback foods. This species was by far the most important food, constituting 27.9% of the total feeding time, and the feeding time on this species negatively correlated with fruit availability. Consumed leaves contained more protein than nonconsumed leaves, and variation in time spent feeding on different leaves was explained by their abundance. These results suggest that red leaf monkeys show essentially the same response to the supra-annual increase in fruit availability as sympatric monogastric primates, increasing their seed and whole-fruit consumption. However, they depended more on young leaves, in particular Spatholobus macropterus, as fallback foods during fruit-scarce periods than did gibbons or orangutans. Their selection of fallback food appeared to be due to both nutrition and abundance.     

Diet of the brown howler monkeyAlouatta fusca in a semi-deciduous forest fragment of southeastern Brazil

The diet of a group of six brown howlers was studied weekly during 12 months in a reserve of 250 ha of secondary, mesophytic, semi-deciduous forest. The phenology of 186 trees of 72 species and 29 families was monitored simultaneously. Scan sampling was used to record the diet from dawn until dusk on a total of 60 days of observation, yielding 718 hr of animal-observer contact and 2,943 feeding scans. The diet was composed of leaves (73%), flowers (12%), and fruits (5%), from 68 identified plant species.Celtis iguanae, Cassia ferruginea, andInga spp. were the main food sources, accounting for approximately 50% of the diet. Young leaves (59%) were preferred to mature leaves (31%), trees contributing 56% and lianas 41% of the leaf diet. The ingestion of young leaves was correlated to the availability of these items, however, the correlations were not significant for flowers and fruits. The diet was poorer in fruits and richer in young leaves of lianas in comparison to other howler monkey species studies, probably as a consequence of the liana abundance in this forest fragment.     

Selectivity and Variability in the Diet of Milne-Edwards' Sifakas (Propithecus diadema edwardsi): Implications for Folivory and Seed-Eating

Propithecus diadema edwardsi combines a strategy of high selectivity for many food items with the use of more readily available and abundant foods. Selectivity is seen in relation to taxonomic affiliations of plants eaten, forest composition, and phenological patterns. Feeding time is concentrated on relatively uncommon species as indicated by high selection ratios. Annual and monthly dietary diversity is low, with food choices restricted to a relatively narrow set of species: a total of 82 and 73 plant species were eaten by the two groups of which, only 22 and 16% contributed >1% feeding time, however, these values fall with commonly reported ranges. Plant parts eaten varied widely from month to month, with overall variance in seed-eating higher than variance in young or mature leaf feeding. No correlation between percentage feeding time and food availability is significant; instead, monthly feeding patterns are better explained by the availability of particular plant species, which also supports a view of high selectivity in food choice. In contrast to selectivity for many seed, whole fruit, and new leaf sources, reliably abundant food sources appear to be leaves from lianas, epiphytes, and parasites, which were eaten in all focal months. While dramatic interannual differences in dietary composition in response to the absence of a preferred seed source highlight the interpretation of folivores as surviving food crunches by turning to readily available, poor quality resources, this strategy is not unique to folivores.     

The role of phytochemistry in dietary choices of Tana River red colobus monkeys(Procolobus badius rufomitratus)

We conducted a phytochemical survey of tree species growing within the riverine forests of the Tana River National Primate Reserve in Kenya to understand better the feeding ecology of an endangered resident primate, the Tana River red colobus monkey (Procolobus badius rufomitratus).Young leaves, which make up a large percentage of this monkey's diet, are significantly higher in nitrogen and lower in acid detergent fiber than more abundant mature leaves are. Phenolic chemistry had little inhibitory effect on feeding by P. b. rufomitratus.Choice among tree species by P. b. rufomitratusappears to be influenced largely by leaf availability,once an acceptable threshold of nitrogen and fiber is reached When mature leaves are eaten, they selected species that are high in nitrogen and low in fiber. A significantly higher nitrogen content was found for the mature leaves of all leguminous versus nonleguminous tree species. Consequently, the availability of certain types of mature leaf species during periods of preferred food scarcity may prove critical to groups of Tana River red colobus monkeys.     

The diet of the olive colobus monkey,Procolobus verus,in Sierra Leone

Observations of the feeding behavior of wild olive colobus monkeys (Procolobus verus)were made in southern Sierra Leone, West Africa. Most data came from systematically sampling one habituated social group, which inhabited old secondary rain forest on Tiwai Island. Olive colobus at Tiwai were highly selective feeders, obtaining most of their food from a small number of uncommon middle-canopy trees, from the forest understory, and from climbers. Mature leaf blades were largely ignored; young foliage (particularly of climbers) was a year-round dietary staple, while seeds, flowers, and mature leaf petioles were seasonally important. It is argued that this diet results largely from selection for two different attributes of food: high digestibility and physical location. Preferred foods had low fiber and tannin contents, while preferred feeding sites were in thick low growth and climber tangles. P. verusis the smallest extant primate species using a foregut fermentation system. Observations of its diet accord with inferences drawn from dental anatomy, digestive physiology, and considerations of body size.     

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.     

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.     

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.     

High flexibility in diet and ranging patterns in two golden monkey (Cercopithecus mitis kandti) populations in Rwanda

Many primates exhibit behavioral flexibility which allows them to adapt to environmental change and different habitat types. The golden monkey (Cercopithecus mitis kandti) is a little-studied endangered primate subspecies endemic to the Virunga massif and the Gishwati forest in central Africa. In the Virunga massif, golden monkeys are mainly found in the bamboo forest, while in the Gishwati forest they live in mixed tropical montane forest. Here we describe and compare the diet of golden monkeys in both fragments. Over 24 consecutive months from January 2017 we used scan sampling to record feeding and ranging behavior of two Virunga groups and one Gishwati group totaling ca. 240 individuals. We also examined the phenology of bamboo and fruit trees, key seasonal food plant species for the monkeys. Golden monkeys fed on more than 100 plant species. The Virunga groups were mostly folivorous (between 72.8% and 87.16% of the diet) and fed mostly on young bamboo leaves and bamboo shoots, while 48.69% of the diet of the Gishwati group consisted of fruit from 22 different tree and shrub species. Bamboo shoots and fruit are seasonally available foods and were consumed regularly throughout the period when they were available. Despite being the smallest of the three study groups, the Gishwati group had a larger home range area (150.07 ha) compared to both Virunga groups (25.24 and 91.3 ha), likely driven by the differences in availability and distribution of fruit and bamboo in the habitats. Like other blue monkey subspecies, golden monkeys appear to have a flexible dietary strategy enabling them to adjust diet and ranging behavior to local habitats and available food resources. Additional studies and continuing conservation efforts are needed to better understand how variation in feeding and ranging ecology affects reproduction, population growth, and carrying capacity.     

Dietary generalists and nutritional specialists: Feeding strategies of adult female blue monkeys (Cercopithecus mitis) in the Kakamega Forest,Kenya

Generalist primates eat many food types and shift their diet with changes in food availability. Variation in foods eaten may not, however, match variation in nutrient intake. We examined dietary variation in a generalist‐feeder, the blue monkey (Cercopithecus mitis), to see how dietary food intake related to variation in available food and nutrient intake. We used 371 all‐day focal follows from 24 adult females (three groups) in a wild rainforest population to quantify daily diet over 9 months. We measured food availability using vegetation surveys and phenology monitoring. We analyzed >700 food and fecal samples for macronutrient content. Subjects included 445 food items (species‐specific plant parts and insect morphotypes) in their diet. Variation in fruit consumption (percentage of diet and total kcal) tracked variation in availability, suggesting fruit was a preferred food type. Fruits also constituted the majority of the diet (by calories) and some fruit species were eaten more than expected based on relative availability. In contrast, few species of young leaves were eaten more than expected. Also, subjects ate fewer young leaves (based on calories consumed) when fruit or young leaves were more available, suggesting that young leaves served as fallback foods. Despite the broad range of foods in the diet, group differences in fiber digestibility, and variation that reflected food availability, subjects and groups converged on similar nutrient intakes (grand mean ± SD: 637.1 ± 104.7 kcal overall energy intake, 293.3 ± 46.9 kcal nonstructural carbohydrate, 147.8 ± 72.4 kcal lipid, 107.8 ± 12.9 kcal available protein, and 88.1 ± 17.5 kcal structural carbohydrate; N = 24 subjects). Thus, blue monkeys appear to be food composition generalists and nutrient intake specialists, using flexible feeding strategies to regulate nutrient intake. Findings highlight the importance of simultaneously examining dietary composition at both levels of foods and nutrients to understand primate feeding ecology.     

Diet in one group of woolly spider monkeys,or muriquis (Brachyteles arachnoides)

Data on feeding behavior in one group of muriquis (Brachyteles arachnoides) were collected during a 14 month study at Fazenda Montes Claros, Minas Gerais, Brazil. Muriquis devoted an average of 51% of their annual feeding time to leaves, 32% to fruits, and 11% to flowers. The high proportion of leaves in their diet is consistent with predictions from the energetics of body size. However, they devoted a greater proportion of their feeding time to patchy fruit and flower resources than was expected from comparisons with smaller, sympatric howler monkeys. Muriqui diet varied with the availability of preferred food types across sample months. Fruit and flower consumption corresponded to the availability and abundance of these food resources in the forest. Leaves contributed substantially to muriqui diet throughout the year but appeared to be eaten primarily to provide necessary protein and/or necessary bulk. Male and female diets differed only in the greater proportion of feeding time females devoted to flowers.     

Diet and Feeding Ecology of <Emphasis Type="Italic">Ateles chamek</Emphasis> in a Bolivian Semihumid Forest: The Importance of <Emphasis Type="Italic">Ficus</Emphasis> as a Staple Food Resource

We describe temporal patterns of food consumption by Peruvian spider monkeys (Ateles chamek) in a semihumid forest in lowland Bolivia. We assessed dietary composition in relation to temporal variation in abundance, duration, and synchrony of different food items in their home range. We collected data from September 2003 to September 2004, in the forestry concession La Chonta, Department of Santa Cruz. Throughout the period of detailed feeding data collection (February-September 2004), Ateles chamek used Ficus as a staple food resource. Figs constituted almost 50% of their diet in terms of total time spent feeding, and subjects consumed them to a great extent even during times of high overall food availability. This is contrary to the general expectation that for Neotropical frugivores, Ficus is a fallback food in times of fruit scarcity, rather than a staple food resource. Surprisingly, despite being considered ripe fruit specialists, Ateles chamek spent 18% of their feeding times eating unripe figs. Ateles chamek consumed unripe figs all through the year, including periods when ripe figs and other ripe fruit were abundant. We identify other important fallback foods for Ateles chamek in the forest, in particular the ripe fruit of Myrciaria sp.     

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 and feeding behaviour of samango monkeys (Cercopithecus mitis labiatus) in Ngoye Forest, South Africa

The samango monkey occurs at the southern limit of the range of Cercopithecus mitis. Greater climatic seasonality at this latitude results in more predictable fruiting patterns. In addition, there are no diurnal sympatric primate frugivores. Under these conditions, the diet and feeding strategies of samango monkeys would be expected to differ notably from those of central or east African C. mitis subspecies. Contrary to these expectations, the preliminary observations reported here indicate that diets of samango and blue monkeys differ only superficially in the proportions of items eaten. Similarities in feeding behaviour are especially marked during the dry season period when fruit is not abundant. Both samango and blue monkeys tend to be less selective in their choice of food species and to eat available food species regardless of their energy content; a shift toward less nutritious items such as leaves is also noted. Feeding behaviour during the summer wet season is characterized by the selection of fruits with high-energy values. A high proportion of visits by the monkeys to areas of greater food availability suggests a concentration of feeding effort in food patches and the selection of higher energy food species within patches.     

Diet ofColobus polykomos on Tiwai Island: Selection of food in relation to its seasonal abundance and nutritional quality

A group of Colobus polykomosat Tiwai, Sierra Leone, demonstrated seasonal flexibility in its diet, with seeds, young leaves, and mature leaves each dominating the diet at different times. Comparison of food consumption with phenological data indicates that seeds are eaten whenever available and are preferred to other foods, while young leaves are preferred to mature leaves. Colobus polykomosalso prefer liane to tree leaves, despite the relatively high quality of mature tree foliage at the Tiwai site. Analysis of protein, fiber, and energy values of foods selected and items available, but not eaten, suggests that preference is related to protein and energy maximization. Leguminous plants, especially Papilionaceae and Mimosaceae, are highlighted as important food sources for C. polykomos;seeds and leaves from these families have a high nitrogen content, and the protein content of leguminous seeds often equals or exceeds that found in leaves. It is predicted, therefore, that colobines living in habitats with a high density of legumes will feed heavily on seeds, subject to constraints such as seasonal availability. Where suitable leguminous species are less common, a mixture of fruits, seeds, and young or mature leaves or both is likely to be selected. The results of this and other recent studies of colobines do not support the notion that colobines are specialist folivores.     

The feeding ecology and activity budget of proboscis monkeys

A group of proboscis monkeys (Nasalis larvatus) consisting of an alpha‐male, six adult females, and several immatures was observed from May 2005–2006. We collected over 1,968 hr of focal data on the adult male and 1,539 hr of focal data on the six females in a forest along the Menanggul River, Sabah, Malaysia. Availability and seasonal changes in plant species consumed by the focal monkeys were determined by vegetation surveys carried out across an area of 2.15 ha along 200–500 m trails in riverine forest. A total of 188 plant species were consumed by the focal monkeys. The activity budget of members of our study group was 76.5% resting, 19.5% feeding, and 3.5% moving. Young leaves (65.9%) and fruits (25.9%) accounted for the majority of feeding time. Over 90% of fruit feeding involved the consumption of unripe fruits and in the majority of case both the fruit flesh and seeds were eaten. Although fruit eating was rare in some months, during other times of the year time fruit feeding exceeded the time devoted to young leaves. We found that monthly fruit availability was positively related to monthly fruit eating and feeding activity, and seasonal fluctuations in dietary diversity were significantly affected by fruit eating. These results suggest that fruit availability and fruit‐eating behaviors are key factors that influence the activity budget of proboscis monkeys. Earlier assumptions that colobine monkeys are obligate folivores do not apply well to proboscis monkeys and certain other colobines. Our findings may help contribute to a better understanding of the dietary adaptations and feeding ecology of Asian colobines. Am. J. Primatol. 71:478–492, 2009. © 2009 Wiley‐Liss, Inc.     

Influence of Forest Seasonality on Gibbon Food Choice in the Rain Forests of Barito Ulu,Central Kalimantan

We describe the diet of two hybrid gibbon groups (Hylobates mulleri x H. agilis) in relation to forest seasonality. We collected data over 12 mo in lowland dipterocarp forest in the Barito Ulu research area, Central Kalimantan, Indonesia. Although non-fig fruit was the main dietary item (52–64% of diet), gibbon diet was most strongly influenced by the availability of flowers. During periods when flowers were most abundant and the gibbons increased consumption of them, they also ate figs or young leaves more often. We suggest that although flowers are nutritionally rich sources of food, providing relatively high levels of protein compared to fruit, they are unlikely to satiate gibbon hunger and they seek dietary bulk from figs or young leaves, because they are easily obtained. Rainfall also influenced food choice, and non-fig fruit availability had a weak influence on fruit selection for one group. The group concentrated feeding on the fruit of a few species when fruit was most abundant and ate a greater diversity of species when fruit was scarce. Gibbon diet appeared not to be influenced by changes in availability of figs, young leaves and diversity of fruiting species.     

Differences in Diet Between Spider Monkey Groups Living in Forest Fragments and Continuous Forest in Mexico

Forest fragmentation can lead to reductions in food availability, especially for some large‐bodied tropical mammals such as spider monkeys. During a 15\xE2\x80\x90mo period, we assessed the diet of Geoffroyi's spider monkey (Ateles geoffroyi) in continuous forest and fragments in the Lacandona region, southern Mexico, and related differences in diet to differences in vegetation structure and composition. We found that both forest types presented top food species for monkeys (e.g., Spondias spp., Brosimum alicastrum), but the sum of the importance value index of these species and the density of large trees were lower in fragments than in continuous forest. We also found that, compared with continuous forest, monkeys in fragments diversified their overall diet, increased consumption of leaves, and reduced the time they spent feeding on trees in favor of more time feeding on hemiepiphytes (particularly Ficus spp.) and palms, both of which were common in fragments. We attribute these changes to the relative food scarcity of the most favored feeding plants in forest fragments. Overall, our findings suggest that monkeys are able to adjust their diet to food availability in fragments, and thus persist in small‐ and medium‐sized fragments. Although it is unlikely that the small size of two of the three study fragments (14 and 31 ha) can maintain viable populations of monkeys in the long term, they may function as stepping stones, facilitating inter‐fragment movements and ultimately enhancing seed dispersal in fragmented landscapes.     

Monthly and diurnal variations in food choice byMacaca fuscata yakui during the major fruiting season at Yakushima Island, Japan

An ecological survey was conducted on two groups of wild Japanese macaques (Macaca fuscata yakui) inhabiting the warm-temperate, broad-leaved forest of Yakushima Island. The survey was designed to determine and explain monthly and diurnal variations in both duration of feeding and food choice, and to explain these variations from a nutritional perspective. Feeding activity increased during the hour before sunset, while leaf eating in particular tended to be observed later in the day throughout the study period. A diverse and unstable diet promoted monthly differences in fat and protein consumption. The mean lipid and calorific content of the food and the rate of protein intake were high in early autumn.Ficus fruits were not selected when unripeArdisia sieboldii fruits were available, but were an important food resource in a general context. Differences in nutritional intake at different times of the day were determined by the combination of species eaten although no single species was preferred at a particular time. Similarities with experimental results in laboratory animals suggest that a physiological, rather than a behavioural, response was regulating nutritional intake. A high consumption of proteins before rest might produce satiety in the macaques and give them the opportunity to digest and absorb complicated or toxic metabolites slowly during the night.