Travel Time Predicts Fecal Glucocorticoid Levels in Free-Ranging Howlers (Alouatta palliata)

Environmental stressors impact physiology in many animal species. Accordingly, the monitoring of fecal glucocorticoid metabolites (fGCM) has been increasingly used to evaluate the physiological costs of habitat disturbance on wild animal populations, providing a powerful tool for conservation and management. Several studies have suggested that primates in forest fragments have higher fGCM levels than those in continuous forests, yet the proximate causes of fGCM variation remain to be identified. In previous studies of Mexican howlers (Alouatta palliata mexicana) in Los Tuxtlas, Mexico, we found that individuals living in a smaller and more disturbed forest fragment consumed significantly less fruit and had a significantly higher feeding effort than those living in a bigger, more conserved forest fragment. Here, we aimed to examine the effects of fruit consumption and travel time on fGCM levels in the same two groups of howlers, during three sampling sessions that differed markedly in fruit availability. We found that fGCM levels (N?=?202 fecal samples) were higher in the howler group living in the smaller forest fragment and varied seasonally in both focal groups, being lowest when fruit consumption was highest. However, our results suggest that travel time is the main factor predicting fGCM levels in howlers, and that although fruit consumption may be negatively related to fGCM levels, this relationship is probably mediated by the strong effect that fruit consumption has on travel time. Our results provide important insight into the proximate causes of fGCM variation in primates in fragments and highlight the potential conservation significance of studies showing that habitat loss and transformation can lead to increases in travel time in wild primates.     

Dietary Flexibility of the Brown Howler Monkey Throughout Its Geographic Distribution

Habitat loss and fragmentation constrain the survival of most forest‐living mammals, particularly strictly arboreal primates. Because fragment size directly affects food availability, primate survival in small fragments may depend on dietary flexibility. Here, we review the literature on the diet of 29 wild groups of Alouatta guariba clamitans inhabiting forest fragments in Brazil and Argentina. We identify general feeding patterns and analyze the influence of fragment size and latitude on diet composition. Brown howlers presented a diet composed of 402 plant species belonging to 227 genera and 80 families. Rarefaction curves suggest that the richness of top food species is similar among groups living in larger (>100 ha), medium (11–100 ha) or small (1–10 ha) fragments. On average, only 12% of the plant species used as food sources by a given group was also consumed by groups from other sites. The shorter the distance between sites, the higher the diet similarity among groups. Despite their diet flexibility, brown howlers spent >80% of the total feeding records on 6–24 species belonging to genera such as Ficus, Zanthoxylum, and Eugenia. Leaves and fruits were the plant items most consumed (65% and 22% of the total feeding records, respectively). Leaf consumption was not affected by fragment size, but it was inversely related to latitude, which may be linked to an increase in the concentration of secondary metabolites in leaves at higher latitudes. We suggest that the ability of brown howlers to exploit a large number of plant food species, including native and exotic trees, shrubs, vines, and lianas, is an important trait that contributes to their survival in highly fragmented habitats along the Atlantic forest. Similar meta‐analyses of data from other howler species are necessary to test whether such dietary flexibility is a genus‐wide pattern. Am. J. Primatol. 75:16‐29, 2013. © 2012 Wiley Periodicals, Inc.     

Recruitment of Black Howler Fruit Trees in Fragmented Forests of Northern Belize

We examined recruitment of trees whose seeds are dispersed by black howler monkeys (Alouatta pigra) in forest fragments within the Community Baboon Sanctuary in north-central Belize. In the fragments, most other large frugivores are absent and howlers dominate the large frugivore community. Consequently, we expected to observe an increased representation of howler fruit trees among the sapling community. To test this prediction preliminarily we observed howler feeding behavior for one year and conducted adult tree and sapling transects in 6 locations where howlers were present. We sampled a seventh site where howlers were absent for vegetation only. We found that in 4 of 6 sites there were proportionately fewer howler fruit saplings when compared to adult tree samples. However, when recruitment of howler fruit trees in the 6 sites was compared to a site where howlers were absent, 11 of 12 species had relatively higher recruitment. The lack of recruitment among howler fruit trees in general likely reflects differences in responses of individual species, as well as disruptions from human activities to natural processes within the forests. The observed lower recruitment patterns of howler trees suggest that over time, the abundance of the trees will likely decline and thus affect the foraging behavior and possibly survival of howlers in the fragmented forests.     

Foraging habits of Alouatta palliata mexicana in three forest fragments

The activity patterns and diet of howler monkeys (Alouatta palliata mexicana) were studied in 3 forest fragments in south-eastern Mexico: Playa Escondida (PLA), Agaltepec Island (AGA) and Arroyo Liza (LIZ). Intersite differences offered the opportunity to investigate the foraging adaptations of howler monkeys in response to population and habitat size. In the largest fragment (PLA), the howlers' diet was based on high-quality items (fruit and young leaves). In AGA, where the density of howlers was the highest, their diet was mostly folivorous with a marked exploitation of uncommon food items such as vines, lianas, shrubs and herbs. The dietary differences in AGA were accompanied by more time spent travelling and less time spent resting. Although LIZ was the smallest fragment and had a high howler density, the small group size and the use of energy-minimizing strategies (less time spent travelling and more time spent resting) probably allowed howlers to maintain a frugivorous diet.     

Ecological strategies of woolly monkeys (Lagothrix lagotricha) at Tinigua National Park,Colombia

Information on the use of space, activity patterns, diet, and social interactions were recorded for a group of woolly monkeys (Lagothrix lagotricha) during 13 months at Tinigua National Park, Macarena, Colombia. In this region, fruit abundance changes throughout the year with a peak during March–April (beginning of the rainy season) and less fruit during September-November (end of rainy season). Woolly monkeys spent most of their time in mature forest where fruit abundance is higher than in opendegraded or flooded forests. Changes in habitat used by monkeys were coupled with changes in fruit supply across vegetation types. On an annual basis, woolly monkeys spent 24% of point samples locomoting, 36% resting, 36% feeding, and 4% on other activities. However, these proportions varied across the year depending on fruit availability. Based on instantaneous samples, the diet consisted mostly of fruits (60%), arthropods (23%), vegetative parts and flowers (17%), and other items (1%). Non-lactating females and juveniles spent more time eating insects than adult males and lactating females; however, significant differences between classes were detected only during the period of fruit scarcity. These differences are probably due to the high extent to which non-lactating females and juveniles were excluded from fruiting trees by males. The high proportion of arthropods in their diet is unusual for primates with large body size and is a possible factor influencing group cohesiveness in woolly monkeys. © 1994 Wiley-Liss, Inc.     

Seasonal variation in the positional behavior of red howling monkeys (Alouatta seniculus)

Recent studies on the positional behavior of primates reveal that significant seasonal variation occurs in both locomotion and postures that is related to changes in diet and foraging techniques. Howling monkeys (genusAlouatta), which also have a seasonally varied diet, are predicted to have correspondingly varied positional behaviors. Two groups of red howling monkeys were studied in a primary rain forest in French Guinana during the dry and wet seasons. During the dry season, when howler diet is based mainly on leaves, howlers traveled more frequently by quadrupedal walking on large supports, a mode of progression that is probably inexpensive energetically and relatively stable. During feeding, quadrupedal and tripedal stand contributed considerably, a posture probably associated with the equal distribution of leaves within a tree crown. In contrast, during the wet season, when fruit was abundant, howlers fed very frequently by sitting on large supports, probably because fruit consumption required more time for special manipulation. However, most seasonal changes in feeding postures, and in travel and feeding locomotion, were difficult to associate directly with dietary shifts. These behavioral changes may be more highly correlated with slight modifications in microhabitat use (horizontal and vertical daily ranges, similar and alternative arboreal pathways) that are not considered in this paper.     

The Influence of Seasonal Variation on Chimpanzee (Pan troglodytes schweinfurthii) Fallback Food Consumption, Nest Group Size, and Habitat Use in Gishwati, a Montane Rain Forest Fragment in Rwanda

The increased number of primates living in fragmented habitats necessitates greater knowledge of how they cope with large-scale changes to their environment. Chimpanzees (Pan troglodytes) are exceptionally vulnerable to forest fragmentation; however, little is known about chimpanzee feeding ecology in fragments. Although chimpanzees have been shown to prefer fruit when it is available and fall back on more abundant lower quality foods during periods of fruit scarcity, our understanding of how chimpanzees use fallback foods in forest fragments is poor. We examined how chimpanzees cope with periods of fruit scarcity in Gishwati Forest Reserve, a disturbed montane rain forest fragment in Rwanda. We assessed seasonal changes in chimpanzee diet and the use of preferred and fallback foods through fecal and food site analysis. We also examined seasonal variation in nest group size and habitat use through marked nest censuses. We found that chimpanzees experienced a seasonal reduction in preferred fruit availability, which led to a seasonal diet shift to more fibrous foods, including several that functioned as fallback foods. Our results suggest that during periods of fruit scarcity the chimpanzees also reduced nest group size. However, we found that the chimpanzees did not alter their habitat use between high- and low-fruit seasons, which suggests that the small size of the forest limits their ability to change their seasonal habitat use. Consequently, fallback foods appear to be particularly important in small food-impoverished habitats with limited ranging options.     

Differences in diet and activity pattern between two groups of Alouatta palliata associated with the availability of big trees and fruit of top food taxa

The threat that forest fragmentation and habitat loss presents for several Alouatta taxa requires us to determine the key elements that may promote the persistence of howler monkeys in forest fragments and to evaluate how changes in the availability of these elements may affect their future conservation prospects. In this study we analyzed the relationship between the availability of both big trees of top food taxa (BTTFT) (diameter at breast height>60) and fruit of top food taxa (FrTFT) in the home ranges of two groups of Alouatta palliata mexicana occupying different forest fragments in Los Tuxtlas, Mexico, and their diet and activity pattern. Both study groups preferred big trees for feeding and the group with lower availability of BTTFT in their home range fed from more, smaller food sources. Furthermore, both study groups also increased the number of food sources when their consumption of fruit decreased, and the group with lower availability of FrTFT in their home range fed from more food sources. The increase in the number of food sources used under such conditions, in turn, set up a process of higher foraging effort and lower rest. In summary, our results support other studies that suggest that the availability of big trees and fruit may be two important elements influencing the persistence of howler monkeys in forest fragments. Am. J. Primatol. 71:654–662, 2009. © 2009 Wiley‐Liss, Inc.     

The Ranging Costs of a Fallback Food: Liana Consumption Supplements Diet but Increases Foraging Effort in Howler Monkeys

Lianas are important components in the dynamics of tropical forests and represent fallback foods for some primates, yet little is known about their impact on primate ecology, behavior or fitness. Using 2 yr of field data, we investigated liana consumption and foraging effort in four groups of howler monkeys (two in bigger, more conserved forest fragments and two in smaller, less conserved fragments) to assess whether howler monkeys use lianas when and where food availability is scarce, and how liana consumption is related to foraging effort. Howler monkeys in smaller fragments spent more time consuming lianas and liana consumption was negatively related to the consumption of preferred food resources (fruit and Ficus spp.). Further, travel time was positively related to liana feeding time, but not to tree feeding time, and howler monkeys visited a greater number of food patches when feeding from liana leaves than when feeding from tree leaves. Our results suggest that these increases in foraging effort were related to the fact that lianas are mainly a source of leaves, and that liana patch size was probably smaller than tree patch size. While these results were clear when analyzing all four groups combined, however, they were not always significant in each of the groups individually. We suggest that this may be related to the differences in group size, patch size and the availability of resources among groups. Further studies are necessary to assess whether these dietary and behavioral adjustments negatively impact on the fitness and conservation of primates in fragments.     

Behavioral modifications in northern bearded saki monkeys (<Emphasis Type="Italic">Chiropotes satanas chiropotes</Emphasis>) in forest fragments of central Amazonia

We investigated behavioral differences among seven groups of northern bearded saki monkeys (Chiropotes satanas chiropotes) living in five forest fragments and two areas of continuous forest at the Biological Dynamics of Forest Fragments Project study area, located approximately 80 km north of Manaus, Brazil. We collected data in six research cycles from July–August 2003 to January 2005–April 2006. When bearded saki monkeys were present in a study area, we followed the group from dawn until dusk for three consecutive days. Every 5 min, we conducted behavioral scans of all visible individuals. There was a positive relationship between forest size and group size, but animals in the small forest fragments lived at greater densities. Bearded saki monkeys in the smaller fragments spent more time resting, less time traveling, and less time vocalizing, but there was no relationship between forest size and the amount of time spent feeding. Our results indicate that the main behavioral differences among the groups are related to the amount of forest resources (e.g., fruit trees, space) available to the monkeys in the smaller fragments, as well as the resulting smaller group sizes. We stress the need to preserve large tracts of forest and provide connectivity between forest patches.     

Resource Use in a Landscape Matrix by an Arboreal Primate: Evidence of Supplementation in Black howlers (Alouatta pigra)

Across the tropics, landscapes of continuous rain forest are being replaced by forest fragments embedded in a matrix of pastures and farmlands. This conversion has endangered many species, including arboreal primates. Species vary, however, in how they are able to supplement their diets from the matrix, although this is rarely studied in primates. We studied two groups of black howlers (Alouatta pigra) for a total of 1156 h, one inhabiting a smaller fragment (0.4 ha) and the other a larger fragment (20 ha). monkeys inhabiting the smaller fragment spent more time in the matrix than in the habitat fragment, spending 50 % of their time (335 of 667 h) in an abandoned mango (Mangifera indica) plantation, 8.8 % in scattered trees, and 0.2 % in pastures. In contrast, monkeys in the larger fragment spent 75 % of their time (368 of 489 h) in the forest fragment and only 25 % of their time in the matrix. Feeding in the matrix accounted for 53 % and 12 % of the foraging time for groups in the smaller and larger fragments respectively. We suggest that Alouatta pigra can use resources in the matrix to supplement their diet by means of crop raiding or taking other resources in many fragmented landscapes and that this may be true also for many fragment-dwelling primates. It is important to include a consideration of the matrix in conservation planning, considering both the total resources available to primates and the consequences of crop raiding for farmers.     

Feeding and general activity patterns of a howler monkey (Alouatta palliata) troop living in a forest fragment at Los Tuxtlas,Mexico

The feeding behavior and general activity patterns of a howler monkey troop living in a 3.6 ha forest fragment were studied at Los Tuxtlas, Mexico, for an annual cycle. Monthly samples of their feeding behavior indicated that they used 52 species of 24 plant families as sources of food. Of these, 67% were trees, which accounted for 96% of total feeding time recorded. Ten species of Moraceae, Cecropiaceae, Anacrdiaceae. and Sapotaceae contributed to 70% of the trees used and to almost 90% of feeding time. The number of plant species used per monthly record varied from 7 to 31 with an average of 19.9 species. Young leaves and ripe fruit were the principal items in the monthly diet of howlers and average percent of time spent consuming these plant parts was 46.7% and 34.8%, respectively. The use of tree species was found to be associated to their importance value and to their pattern of spatial dispersal in the study site. Availability of young leaves was fairly constant from month to month, but it presented a seasonal pattern, and there was a significantly lower number of tree species bearing ripe fruit through the year with brief pulses of production. The monthly activity pattern was found to be related to variations in the availability of young leaves and ripe fruit as well as to the values of the intermonthly overlap in plant species used. Resting and feeding presented a bimodal pattern of occurrence throughout the day that seemed to be related to variations in maximum ambient temperatures. Results are discussed in light of the small size and shape of the forest fragment inhabited by the howler troop. Am. J. Primatol. 48:167–183, 1999. © 1999 Wiley‐Liss, Inc.     

Foraging ecology of howler monkeys in a cacao (Theobroma cacao) plantation in Comalcalco, Mexico

Recent evidence indicates that primate populations may persist in neotropical fragmented landscapes by using arboreal agroecosystems, which may provide temporary habitats, increased areas of vegetation, and connectivity, among other benefits. However, limited data are available on how primates are able to sustain themselves in such manmade habitats. We report the results of a 9-month-long investigation of the feeding ecology of a troop of howler monkeys (n = 24) that have lived for the past 25 years in a 12-ha cacao plantation in the lowlands of Tabasco, Mexico. A vegetation census indicated the presence of 630 trees (> or =20 cm diameter at breast height (DBH)) of 32 shade species in the plantation. The howlers used 16 plant species (13 of which were trees) as sources of leaves, fruits, and flowers. Five shade tree species (Ficus cotinifolia, Pithecellobium saman, Gliricidia sepium, F. obtusifolia, and Ficus sp.) accounted for slightly over 80% of the total feeding time and 78% of the total number trees (n = 139) used by the howlers, and were consistently used by the howlers from month to month. The howlers spent an average of 51% of their monthly feeding time exploiting young leaves, 29% exploiting mature fruit, and 20% exploiting flowers and other plant items. Monthly consumption of young leaves varied from 23% to 67%, and monthly consumption of ripe fruit varied from 12% to 64%. Differences in the protein-to-fiber ratio of young vs. mature leaves influenced diet selection by the monkeys. The howlers used 8.3 ha of the plantation area, and on average traveled 388 m per day in each month. The howlers preferred tree species whose contribution to the total tree biomass and density was above average for the shade-tree population in the plantation. Given the right conditions of management and protection, shaded arboreal plantations in fragmented landscapes can sustain segments of howler monkey populations for many decades.     

Primate Group Size and Abundance in the Caatinga Dry Forest, Northeastern Brazil

The Caatinga dry forest poses a series of ecological challenges for mammals in general and primates in particular. The erratic rainfall pattern impacts on plant diversity and phenological patterns; from year to year there is marked variability in fruit production and failure to fruit is common. The harshness apparently accounts for the impoverished mammalian fauna. However, data on primate abundance, distribution, and possible environmental effects on primate density are lacking in this type of dry forest. I censused the primate community in 3 habitats of the Serra da Capivara National Park, Piaui, NE Brazil, over a total distance of 318 km. Overall, the abundance of primates in the Caatinga dry forest is very low as a consequence of low abundance of food resources both in space and time. Alouatta caraya (predominantly folivorous) occurs at extremely low density, and during the dry season are apparently confined to canyon areas, where trees retain their leaves. Callithrix jacchus has morphological feeding specializations for gum-eating, and gum is an important resource during food bottleneck periods. Nonetheless, Callithrix jacchus occurs at comparatively low densities. Group sizes for howlers and marmosets in the Caatinga are significantly smaller than in other forest types. Contrarily, Cebus apella libidinosus had an average group size within the range reported for Amazonian and Atlantic forests. Researchers consider the generalized diet of capuchins as the explanation for their similar abundance in different habitats, indicating relative independence from ecological constraints. However, I suggest that capuchin foraging style and cognitive abilities are important factors accounting for their unreduced group size and density even under extreme conditions.     

Use of space, activity patterns, and foraging behavior of red howler monkeys (Alouatta seniculus) in an Andean forest fragment in Colombia

Howler monkeys are among the most studied primates in the Neotropics, however, behavioral studies including estimation of food availability in Andean forests are scarce. During 12 months we studied habitat use, behavior, and feeding ecology of two groups of red howler monkeys (Alouatta seniculus) in an isolated fragment in the Colombian Andes. We used a combination of focal animal and instantaneous sampling. We estimated fruit production (FP) using phenology transects, and calculated young leaf abundance by observing marked trees. The home range area used by each group was 10.5 and 16.7 ha and daily distances traveled were 431 ± 228 and 458 ± 259 m, respectively. We found that both groups spent most of their time resting (62–64%). Resting time did not increase with leaf consumption as expected using a strategy of energy minimization. We did not find a relationship between daily distances traveled and leaf consumption. However, howlers consumed fruits according to their availability, and the production of young leaves did not predict feeding time on this resource. Overall, our results are similar to those found on other forest types. We found that despite limited FP in Andean forests, this did not lead to a higher intake of leaves, longer resting periods, or shorter traveling distances for red howlers. Am. J. Primatol. 73:1062–1071, 2011. © 2011 Wiley‐Liss, Inc.     

Diets of some French guianan primates: Food composition and food choices

We studied food choices of black spider monkeys (Ateles paniscus) and red howlers (Alouatta seniculus) in an undisturbed tropical forest of French Guiana for 6 months in the rainy season. We made additional observations on tufted capuchins (Cebus apella) and examined the differences and similarities in feeding behavior with respect to the plant specific composition of the habitat and the biochemical characteristics of their food. Capuchins and spider monkeys mainly fed on ripe fruit pulp, to which they added invertebrates (capuchins) or young leaves (spider monkeys); their plant diet was more varied than that of howlers, which contained approximately equal proportions of ripe fruits and young leaves of many species, with large monthly variations in these food categories. BothAteles andAlouatta tended to feed preferentially on abundant plant species, with a large overlap in their fruit choices. The former species included a high proportion of soluble sugars and a low proportion of protein in its diet compared, to howlers. Leaves of several species selected byAlouatta reacted positively when screened for alkaloids and phenolic compounds. Besides specific metabolic requirements and adaptations to deriving nutrients from distinct food types, we hypothesize that the ability to taste sugars, which varies among primates, affects the range of foods appearing to be palatable and, consequently, contributes to the differentiation of feeding niches.     

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.     

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.     

Diet of the Critically Endangered Brown Spider Monkey (Ateles hybridus) in an Inter‐Andean Lowland Rainforest in Colombia

Brown spider monkeys (Ateles hybridus) are one of the least known and more threatened primates in the Neotropics. Recognized as a species about a decade ago, field studies on these endangered primates have mainly focused on estimating local population densities. Since 2006, we habituated a group of wild brown spider monkeys at Serranía de Las Quinchas, Colombia, and studied their feeding ecology during 2.5 years using focal “subgroup” sampling, and conducted phenological surveys in order to estimate habitat‐wide fruit availability. Based on 847 hr of behavioral follows, brown spider monkeys spent approximately 25% of their time in feeding activities, and fed from fruits and leaves on at least 123 plant species. Ripe fruits were the most important item in the diet of A. hybridus at Las Quinchas comprising 92% of their feeding time. Probably due to the minor variation in the monthly proportion of fruits in brown spider monkey's diet throughout this study, there was no relation between habitat‐wide fruit availability and the proportion of fruit included in their monthly diet. The diet of brown spider monkeys at Las Quinchas is toward the high end of fruit intake, even within other wild spider monkeys’ populations, suggesting that these endangered primates might also be facing the challenges of being a large bodied fruit specialist under a regional scenario of habitat loss and fragmentation. Am. J. Primatol. 74:1097‐1105, 2012. © 2012 Wiley Periodicals, Inc.     

Feeding Strategies of Brown Howler Monkeys in Response to Variations in Food Availability

Primates display varying degrees of behavioral flexibility that allow them to adjust their diet to temporal changes in food availability. This trait might be critical for the survival of folivorous-frugivorous species inhabiting small forest fragments, where the availability of food resources tends to be lower than in large fragments and continuous forests. However, the scarcity of studies addressing this issue hampers our understanding of the adaptive behaviors that favor the survival of these primates in low-quality habitats. We conducted a 36-mo study testing the hypothesis that brown howler monkeys (Alouatta guariba clamitans) are able to adjust their diet in response to local and seasonal changes in resource availability. We compared the diet of six free-ranging groups inhabiting three small (<10 ha) and three large (>90 ha) Atlantic forest fragments in southern Brazil and estimated the temporal availability of their top food species (i.e., those species that together contribute ≥80% of total feeding records). We found that brown howlers exploited similarly rich diets in small (45, 54, and 57 plant species) and large (48, 51, and 56 species) fragments. However, intermonth diet similarity was higher for groups in small fragments, where howlers also fed on plant items from nine alien species. Fruits and leaves were the most consumed plant items in both small (42% and 49% of feeding records, respectively) and large (51% and 41%, respectively) fragments. The consumption of young leaves was higher in small than in large fragments, whereas the consumption of other plant items did not show a pattern related to fragment size. Regarding the contribution of growth forms as food sources, only the exploitation of palms showed a pattern related to fragment size. Palms contributed more to the diet of groups inhabiting large fragments. The availability of seasonal food items–ripe fruits and young leaves–influenced their consumption in both habitat types. Therefore, brown howlers cope with local and seasonal fluctuations in food availability by opportunistically exploiting resources. We believe that this feeding flexibility is a key component of the phenotypic plasticity that enables howlers to thrive in disturbed habitat patches, where periods of scarcity of preferred foods shall be more common.