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.     

Diet and activity pattern of howler monkeys (Alouatta palliata) in Los Tuxtlas, Mexico: effects of habitat fragmentation and implications for conservation

Accelerated deforestation is causing the rapid loss and fragmentation of primary habitat for primates. Although the genus Alouatta is one of the most studied primate taxa under these circumstances, some results are contradictory and responses of howlers to habitat fragmentation are not yet clear. In this paper, we conduct a cross-study of the available researches on mantled howlers (Alouatta palliata) in forest fragments in Los Tuxtlas, Mexico, to (1) describe the diet and activity pattern of howlers; (2) analyze the similarity in the diet across studies; and (3) relate both fragment size and howler population density with different characteristics of their diet, home range size, and activity pattern. Howlers consumed 181 plant species belonging to 54 families. Ficus was the most important taxa in the howlers' diet, followed by primary species such as Pterocarpus rohrii, Nectandra ambigens, Poulsenia armata, and Brosimum alicastrum. Secondary and non-secondary light-demanding plant species, which are representatives of disturbed habitat, contributed with a high percentage of their feeding time. Only 23% of the species consumed were the same across all the studies, suggesting that howlers adapt their diet to the food availability of their respective habitats. Population density is the best predictor of howlers' ecological and behavioral changes in response to forest fragmentation, probably owing to its relationship with food availability. Howlers respond to the increase in population densities by increasing the (1) diversity of food species in the diet; (2) consumption of non-tree growth forms; and (3) consumption of new plant items. Home range size is also predicted by population density, but fragment size is a better predictor, probably owing to the fact that howler groups can overlap their home ranges. Our results emphasize the importance of conserving the larger fragments and increasing the size of small and medium-sized ones.     

Seasonal Variations in the Diet and Feeding Effort of Two Groups of Howlers in Different Sized Forest Fragments

Little is known about how resource limitation affects the feeding ecology of primates in forest fragments. Here, we describe seasonal variation in the diet and feeding effort of 2 groups (RH and RC3) of howlers (Alouatta palliata mexicana) living in different sized forest fragments in Los Tuxtlas, Mexico. The RH group, which lived in a larger and more preserved forest fragment, with a higher availability of fruit and Ficus trees, had a higher and more constant consumption of fruit throughout the study year. Moreover, this group fed from larger food sources, i.e., trees, lianas, and shrubs, and spent more time feeding from each food source. The feeding effort, defined as the time spent feeding and traveling divided by time spent resting, of the RH group was also significantly lower and more stable than that of the RC3 group throughout the study year. As feeding effort has been positively related to stress in primates, such increases in feeding effort could have negative consequences for howlers in small or degraded forest fragments. Our study suggests that habitat characteristics interact with plant seasonality to determine the diet and feeding effort of howlers in forest fragments and that groups living in small and degraded forest fragments may be particularly vulnerable to years of low fruit production.     

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.     

Forest Fragmentation Modifies Habitat Quality for Alouatta palliata

Fragmentation reduces habitat area, increases the number of habitat patches, decreases their size, and increases patch isolation. For arboreal mammals such as howlers (Alouatta palliata), canopy modifications from fragmentation processes could also negatively affect habitat quality. We analyzed changes in the composition and plant structure of 15 fragments (1–76 ha) and compared them with vegetation from a continuous tropical rain forest reserve (700 ha) in Los Tuxtlas, Mexico. At each site, we sampled 1000 m² of all trees, shrubs, and lianas with a diameter at breast height (DBH) ≥10 cm. We obtained estimates of species richness, density, and basal area for different ecological groups, DBH ranges, and top food resources for howlers. We used a stepwise multiple regression analysis to determine relationships between fragment characteristics (size, shape index, and isolation) and plant variables. Compared to continuous forest, fragments have altered composition and plant structure, with large trees absent from the canopy. The basal area of top food resources is higher in continuous forest. Fragment size is the best explanation for the differences in composition and plant structure. The largest fragments had greater basal area of top food resources and more large primary trees in the canopy. Overall, our results suggest that fragmentation altered the habitat quality for howlers.     

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.     

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.     

Resource use by howler monkeys (Alouatta palliata) in the rain forest of Los Tuxtlas,Veracruz, Mexico

The feeding preferences of howler monkeys at their northernmost distribution in the Neotropics are reported for an annual cycle. A remarkable selectivity for 27 species representing 15 families was observed. The Moraceae and Lauraceae plant families were the most important in the diet. The howlers spent an almost equal proportion of their feeding time eating leaves and fruit, and displayed a marked preference for young leaves and mature fruit. The consumption of different plant parts was markedly seasonal and the howlers’ ranging behavior was closely associated with the availability of young leaves and mature fruit. Their home range was unusually large (ca. 60 ha) for howlers and the food species exploited occur at very low densities (93%, ≤ 4 ind/ha). They chose food items richer in protein and energy. Alkaloid compounds, present in some of the leaves, play a secondary role in their dietary selectivity.     

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.     

Primate Population Densities and Sizes in Atlantic Forest Remnants of Northern Espírito Santo,Brazil

We surveyed primates in seven fragments of Atlantic forest ranging from 210 to 24,250 ha in Espírito Santo, southeastern Brazil. The objective was to assess their population densities and fragment-specific population sizes in order to discuss the survival of primates in this highly fragmented landscape. We used line-transect sampling for censusing primates and estimated densities via DISTANCE program. Alouatta fusca, Cebus apella, Callicebus personatus, and Callithrix geoffroyi live in the study area. Although variable, primate numbers were higher in larger than in smaller fragments, the likely reasons being the lack of space and food resources (fruits) and higher impact of mesopredators (small cats) in small fragments. The estimated densities are within the range reported for other Atlantic forest sites, except for brown howlers (Alouatta fusca), which were extremely rare. Estimates of population sizes were on the order of several hundreds to thousands of individuals/species in the large-sized fragments (ca. 20,000 ha), while comparable estimates for the other areas indicate <500 individuals/species in the medium-sized reserves (ca. 2,000 ha), and <50 individuals/species in the small fragments (ca. 200 ha). These estimates suggest that only Atlantic forest fragments 20,000 ha can harbor primate populations large enough to escape extinction in the long term. Given that <20% of all protected areas remaining in the Atlantic forest are 20,000 ha, management measures are urgently needed to improve primate conservation in this biome.     

Habitat selection and diet of Western Capercaillie Tetrao urogallus in an atypical biogeographical region

Ecological features and conservation requirements of populations at the latitudinal limits of a species’ geographical range frequently differ from those in other parts of the range. Identifying such differences is key to implementing effective conservation strategies for threatened range‐edge populations especially, in the context of rapid global warming, at the lower‐latitude range edge. We studied habitat selection and diet of the endangered Cantabrian Capercaillie Tetrao urogallus cantabricus in a recently discovered population at the southernmost edge of the sub‐species’ range. This is the only Western Capercaillie population in the Mediterranean biogeographical region. We combined non‐systematic surveys based on questionnaires, reports and field sampling with data from radiotracking to assess habitat selection. Diet was surveyed by micro‐histological methods from droppings collected in the new population, which inhabits Pyrenean Oak Quercus pyrenaica forests and Scots Pine Pinus sylvestris plantations, and in two Cantabrian populations inhabiting Eurosiberian forests. Capercaillie preferred large (> 500 ha) and medium‐sized (100–500 ha) Pyrenean Oak forest fragments and large Scots Pine plantations. Forest fragments smaller than 100 ha and non‐forested habitats were always avoided. Diet differed markedly between Mediterranean and Eurosiberian populations. Bilberry Vaccinium myrtillus is common in the diet of most Capercaillie populations but was scarce in the study area and so was rare in the diet of the new population. Instead, Rockrose Halimium lasianthum was described for the first time as a major food resource for the Capercaillie and was consumed in autumn and winter. Pine needles were also heavily consumed in winter. We document for the first time the strong preference of Capercaillie for Pyrenean Oak forests and a moderately high consumption of the leaves, buds and acorns of this tree species throughout the year. Habitat selection and diet of this Mediterranean population differ from those of the core Cantabrian and other populations. Our results suggest a wider environmental tolerance (phenotypic plasticity) in the species than previously recognized. We advocate specific protection for this unique range‐edge Capercaillie population and its Pyrenean Oak forest habitat.     

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.     

Gastrointestinal parasites of Howler monkeys (Alouatta palliata) inhabiting the fragmented landscape of the Santa Marta mountain range,Veracruz, Mexico

In recent years populations of howler monkeys (Alouatta palliata) in southeastern Mexico have decreased substantially due to the transformation and loss of natural habitats. This is especially evident in the Santa Marta mountain range, Veracruz, Mexico where several studies have evaluated the impact of fragmentation on howler monkey populations in order to propose management programs for their conservation. The conditions generated by fragmentation likely change the rates of parasitic infection and could decrease howler survival. In this study, gastrointestinal parasite species richness, prevalence, and egg density of infection were determined in howler groups inhabiting five forest fragments at the Santa Marta mountain range. Two hundred and seventy‐eight fresh fecal samples were collected between October 2002 and April 2003. Three parasite species were found during the dry and the wet season in all forest fragments sampled: one unidentified species of Eimeriidae; Trypanoxyuris minutus (Oxyuridae); and Controrchis biliophilus (Dicrocoeliidae). Both the prevalence of T. minutus and infection density for all parasites differed between seasons and fragments (the largest fragment consistently differed from other fragments). Host density, distance to the nearest town, fragment size, fragment shape, and total basal area of food trees explained parasite prevalence, but each species had a different pattern. Although parasite richness was lower, prevalence and density were higher than values reported for howlers in conserved forests. These results suggest that the establishment of biological corridors and animal translocation programs must take into account the parasite ecology of each fragment to avoid higher infection rates and preclude potential consequent mortality. Am. J. Primatol. 72:539–548, 2010. © 2010 Wiley‐Liss, Inc.     

Comparison of habitat quality and diet of Colobus vellerosus in forest fragments in Ghana

The forest fragments surrounding the Boabeng-Fiema Monkey Sanctuary (BFMS) in central Ghana shelter small populations of Colobus vellerosus. Little is known about these populations or the ability of the fragments to support them, despite the fact that these fragments represent potentially important habitat for the colobus in this region. We compared the diet of three groups of C. vellerosus in the fragments to two groups in BFMS. We also examined the differences in plant species composition and food abundance among fragments. The study took place from June to November 2003. Dietary data were collected using scan sampling. Plant species composition and food abundance were evaluated using tree plots and large tree surveys. As in BFMS groups, leaves constituted the highest proportion of the diet of fragment groups, yet the colobus in fragments fed on more lianas than did those in BFMS. Over 50% of all species observed eaten by colobus in the fragments were not consumed in BFMS groups during the same season. Food abundance was similar between fragments and BFMS, although species composition differed. There was no relationship between the density of colobus and the density of food trees or percentage of food species, suggesting that other factors may be influencing the number of colobus present. This study highlights the broad dietary range of C. vellerosus, which may be a factor allowing its survival in these fragments.     

Wet and dry season diets of the Endangered Lemur catta (ring‐tailed lemur) in two mountainous rocky outcrop forest fragments in south‐central Madagascar

Lemur catta's ability to consume a wide variety of plant foods is a key to this species' survival in a time of ecological crisis across its geographic range. We examined seasonal diet variability of L. catta groups inhabiting two rocky outcrop fragments in south‐central Madagascar: Anja Reserve and Tsaranoro Valley forest. Leaves and fruit of Melia azedarach were a keystone resource for Anja lemurs in wet and dry seasons. At Tsaranoro, L. catta relied on M. azedarach and Ficus spp. in dry season, but during the wet season, neither was a dominant food resource. Top food species at both sites differed markedly from those consumed by L. catta in other habitats. At Tsaranoro, a greater proportion of lemurs engaged in feeding during the dry season compared with wet season. We attribute this to resource scarcity during dry season, when greater feeding effort is needed to maintain energy requirements. Because M. azedarach is ubiquitous throughout Anja Reserve, producing fruit and leaves year‐round, Anja lemurs can meet energy requirements with little seasonal adjustment in feeding activity. L. catta's IUCN status has been upgraded to Endangered, thus, greater insight into its diet flexibility and ability to survive on introduced plant species, can inform conservation plans in remaining wild habitats and ex situ programs.     

Are howler monkey species ecologically equivalent? Trophic niche overlap in syntopic Alouatta guariba clamitans and Alouatta caraya

According to the principle of competitive exclusion, niche differentiation allows the stable coexistence of closely related species. We analyzed dietary profile and diversity, and dietary overlap between syntopic brown howlers (BR; Alouatta guariba clamitans) and black and gold howlers (BLG; A. caraya) in the Atlantic Forest of NE Argentina, with the objective of evaluating the degree of trophic niche overlap and potential interspecific competition for food. During 12 months, we collected data on feeding behavior of two groups of each howler species using the scan sampling method, together with data on food availability. Both at the group‐ and species‐level, we analyzed feeding behavior in terms of monthly percentages of time spent feeding on each food type and specific food item, dietary diversity (Shannon index H′), and we estimated dietary overlap using the percentage index and the Morisita–Horn index (C_H). Across months, both howlers showed species‐specific preferences for certain food items, and BLG had a more diverse diet (mean±SE, H′=2.77±0.08) than BR (H′=2.39±0.09). However, diets of both species overlapped extensively (percentage index=45.64±2.97%; C_H=0.6±0.05) and diets of conspecific groups did not overlap more than diets of groups of different species. Given their high degree of trophic overlap, syntopic BR and BLG meet one of the conditions necessary for interspecific food competition to occur. Although at present we lack direct evidence for interspecific competition in these howler species, we conclude that high levels of niche overlap may have an important role in maintaining the essentially parapatric distribution of howler species throughout the Neotropics. Am. J. Primatol. 72:173–186, 2010. © 2009 Wiley‐Liss, Inc.     

Abnormal Pelage Color in an Isolated Population of <Emphasis Type="Italic">Alouatta guariba clamitans</Emphasis> Cabrera, 1940 in South Brazil

We located 4 brown howlers (1 adult male, 2 adult females, and 1 juvenile male) showing abnormally lighter pelage in 3 social groups comprising 5, 6, and 9 individuals in a 20 ha-forest fragment in the State of Rio Grande do Sul, Brazil. Two additional groups composed only of normally colored individuals also live in the fragment, which is isolated from nearby fragments by 267–1009 m. They were the only brown howlers with abnormal pelage color out of a total of 386 individuals belonging to 67 groups in 21 fragments in the 5876-ha study area. The isolation of the forest fragment, its high howler density (2.2 individuals⁄ha), and large group size (8.8 ± 2.4 individuals) may decrease the likelihood of successful immigration into the population, leading to an increased probability of inbreeding that may facilitate the expression of rare alleles.     

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.     

Effect of forest fragmentation on bird species richness in Papua New Guinea

Tropical forests worldwide are being fragmented at a rapid rate, causing a tremendous loss of biodiversity. Determining the impacts of forest disturbance and fragmentation on tropical biotas is therefore a central goal of conservation biology. We focused on bird communities in the interior (>100 m from forest edge) of forest fragments (300, 600, and 1200 ha) in the lowlands of Papua New Guinea and compared them with those in continuous forest. We surveyed bird communities using point counts, mist‐netting, and random walks, and measured habitat and microclimate characteristics at each site. We also surveyed leaf‐dwelling arthropods, butterflies, and ants, and obtained diet samples from birds to examine food availability and food preferences. We recorded significantly fewer bird species per point in the 300‐ha forest fragment than in other study sites. Overall, we recorded 80, 84, and 88 species, respectively, in forest fragments, and 102 in continuous forest. Frugivores (especially large frugivores) and insectivores had lower species richness in forest fragments than continuous forest. Our results did not support the food scarcity hypothesis, that is, the decline of insectivorous birds in forest fragments is caused by an impoverished invertebrate prey base. We also found no significant differences among forest fragments and continuous forest in microclimates of forest interiors. Rather, we found that microhabitats preferred by sensitive birds (i.e., 30% of species with the strongest preferences for continuous forest) were less common in forest fragments (19%–31% of points) than in continuous forest (86% of points). Our results suggest that changes in microhabitats may make forest fragments unsuitable for sensitive species. However, limited dispersal capabilities could also make some species of birds less likely to disperse and occupy fragments. In addition, impoverished food resources, size of the forest fragment, or hunting pressure could contribute to the absence of large frugivorous birds in forest fragments. The forest fragments in our study, preserved as village‐based protected areas, were not large enough to sustain the bird communities found in continuous forest. However, because these fragments still contained numerous bird species, preservation of such areas can be an important component of management strategies to conserve rainforests and birds in Papua New Guinea.     

Fruit diet of Alouatta guariba and Brachyteles arachnoides in Southeastern Brazil: comparison of fruit type, color, and seed size

Fruit is an important food resource for neotropical primates. In this study I compare the fruit diet of sympatric brown howlers (Alouatta guariba) and southern muriquis (Brachyteles arachnoides). Feeding behavior was studied over 12 months and fruit species consumed were identified and assigned to the categories fruit type, fruit color, and seed size. Observed-fruit feeding records were compared with expected records determined from local availability of the fruit of the tree species. I also determined dietary overlap. Fruit consumption occupied 8 and 12% of the feeding time of A. guariba and B. arachnoides, respectively. Fruit from eight tree species were consumed by the former and fruit from twenty-two species by the latter. Patterns of fruit selection of A. guariba and B. arachnoides varied widely. Although howlers and muriquis converge behaviorally by selecting fruit with common attributes (fleshy/unprotected, violet and brown/black-colored), unlike A. guariba, B. arachnoides fed on immature seeds of fleshy/protected and dry fruit. Large seeds were ingested, and defecated intact, by B. arachnoides only. There was little overlap of fruit diet even within categories that had been selected by both, suggesting that dietary divergence is occurring at the interspecific level. Different resource exploitation probably mediates the coexistence of A. guariba and B. arachnoides in low diversity, semideciduous forests, where the environment imposes narrow limits on primate food choices.