Geographic variation in tool use on Neesia fruits in orangutans

Geographic variation in the presence of skilled behavior may reflect geographic variation in genetic predispositions or ecological conditions (accompanied by reliable expression during development), or it may reflect the vagaries of invention and the appropriate social conditions for persistence. In this study, we compare the feeding techniques and tool-using skills used by orangutans to extract the nutritious seeds from Neesia fruits between Suaq Balimbing on Sumatra and Gunung Palung on Borneo, and map the distribution of Neesia tool use in Sumatran swamps. We show that neither genetics nor ecology is sufficient to explain the distribution of this tool use, confirming earlier findings on chimpanzees. We conclude that the ability to learn to use tools determines the geographic distribution. It is impossible to distinguish between the history of invention and the conditions for social transmission as the causal factors, but the high density and the social tolerance at Suaq Balimbing create propitious conditions for the maintenance of the skill as a tradition once it has been invented. High orangutan densities in the other Sumatran coastal swamps with Neesia tool use support the conclusion that suitable transmission conditions are the critical factor to explain the geographic distribution of skills such as feeding tool use.     

The conditions for tool use in primates: implications for the evolution of material culture

In order to identify the conditions that favored the flourishing of primate tool use into hominid technology, we examine inter- and intraspecific variation in manufacture and use of tools in extant nonhuman primates, and develop a model to account for their distribution. We focus on tools used in acquiring food, usually by extraction. Any model for the evolution of the use of feeding tools must explain why tool use is found in only a small subset of primate species, why many of these species use tools much more readily in captivity, why routine reliance on feeding tools is found in only two species of ape, and why there is strong geographic variation within these two species. Because ecological factors alone cannot explain the distribution of tool use in the wild, we develop a model that focuses on social and cognitive factors affecting the invention and transmission of tool-using skills. The model posits that tool use in the wild depends on suitable ecological niches (especially extractive foraging) and the manipulative skills that go with them, a measure of intelligence that enables rapid acquisition of complex skills (through both invention and, more importantly, observational learning), and social tolerance in a gregarious setting (which facilitates both invention and transmission). The manipulative skills component explains the distribution across species of the use of feeding tools, intelligence explains why in the wild only apes are known to make and use feeding tools routinely, and social tolerance explains variation across populations of chimpanzees and orangutans. We conclude that strong mutual tolerance was a key factor in the explosive increase in technology among hominids, probably intricately tied to a lifestyle involving food sharing and tool-based processing or the acquisition of large, shareable food packages.     

Chimpanzee insectivory in the northern half of Uganda’s Rift Valley: do Bulindi chimpanzees conform to a regional pattern?

Insects are a nutritious food source for many primates. In chimpanzees, insectivory is most prevalent among communities that manufacture tools to harvest social insects, particularly ants and termites. In contrast to other long-term study sites, chimpanzees (Pan troglodytes schweinfurthii) in Budongo Forest and Kibale National Park, Uganda, rarely eat insects and have small foraging tool kits, supporting speculation that infrequent insectivory—technically aided or otherwise—characterises chimpanzees in this part of Uganda’s Rift Valley. To expand the dataset for this region, insect foraging was investigated at Bulindi (25 km from Budongo) over 19 months during two studies in 2007–2008 and 2012–2013. Systematic faecal analysis demonstrated that insectivory is a habitual foraging activity at this site. Overall levels of insect consumption varied considerably across months but were not predicted by monthly changes in rainfall or fruit intake. Unlike their Budongo and Kibale counterparts, Bulindi chimpanzees often consume ants (principally weaver ants, Oecophylla longinoda) and use sticks to dig out stingless bee (Meliponini) ground nests. In other respects, however, insectivory at Bulindi conforms to the pattern observed elsewhere in this region: they do not manufacture ‘fishing’ or ‘dipping’ tools to harvest termites and aggressive or hard-to-access ants (e.g., army ants, Dorylus spp.), despite availability of suitable prey. The Bulindi data lend support to the supposition that chimpanzees in this part of the Rift Valley rarely exploit termites and Dorylus ants, apparently lacking the ‘cultural knowledge’ that would enable them to do so most efficiently (i.e., tool use). The study’s findings contribute to current debates about the relative influence of genetics, environment and culture in shaping regional and local variability in Pan foraging ecology.     

The socioecology of fission-fusion sociality in Orangutans

Fission-fusion systems can have the group or the individual as their primary unit. In group-based fission-fusion systems, predation risk reduction is the major benefit to grouping, in the individual-based ones the benefits are likely to be primarily social. Orangutans, like chimpanzees, are examples of an individual-based fission-fusion species. The orangutans inhabiting a Sumatran swamp forest (Suaq Balimbing) are more likely than elsewhere to form travel parties. As elsewhere, the main benefits of grouping are social: mating opportunities, protection from harassment and socialization of infants. Most animals also incur costs, but these are relatively low at Suaq Balimbing due to the high productivity of the swamp. Costs seem to be disproportionately high for females with mid-sized infants, who avoid parties.     

Forest Structure and Support Availability Influence Orangutan Locomotion in Sumatra and Borneo

The influence of habitat structure and support availability on support use is an important aspect of understanding locomotor behavior in arboreal primates. We compared habitat structure and support availability in three orangutan study sites—two on Sumatra (Pongo abelii) in the dry‐lowland forest of Ketambe and peat swamp forest of Suaq Balimbing, and one on Borneo (Pongo pygmaeus wurmbii) in the disturbed peat swamp forest of Sabangau—to better understand orangutan habitat use. Our analysis revealed vast differences in tree and liana density between the three sites. Sabangau had a much higher overall tree density, although both Sumatran sites had a higher density of larger trees. The two peat swamp forests were more similar to each other than to Ketambe, particularly with regard to support availability. Ketambe had a wider variety of supports of different sizes and types, and a higher density of larger lianas than the two peat swamps. Orangutans in all three sites did not differ substantially in terms of their preferred supports, although Sumatran orangutans had a strong tendency to use lianas, not observed in Sabangau. Differences in observed frequencies of locomotor behavior suggest the homogeneous structure of Sabangau limits the locomotor repertoire of orangutans, with high frequencies of fewer behaviors, whereas the wider range of supports in Ketambe appears to have facilitated a more varied locomotor repertoire. There were no differences among age‐sex classes in the use of arboreal pathways in Suaq Balimbing, where orangutans selected larger trees than were typically available. This was less apparent in Sabangau, where orangutans generally used trees in relation to their environmental abundance, reflecting the homogeneous nature of disturbed peat swamp forest. These results demonstrate that forest architecture has an important influence on orangutan locomotion, which may become increasingly important as the structure of orangutan habitat continues to be altered through human disturbance. Am. J. Primatol. 74:1128‐1142, 2012. © 2012 Wiley Periodicals, Inc.     

Individual variation in the rate of use of tree-hole tools among wild orang-utans: implications for hominin evolution

Primate tool use varies among species, populations, and individuals. Individual variation is especially poorly understood. Orang-utans in the Sumatran swamp forest of Suaq Balimbing varied widely in rates of tool use to extract honey, ants or termites from tree holes and in the degree to which they specialized on this tree-hole tool use. We tested whether individual variation was best explained by effects of social dominance, habitat differences, or by opportunities for socially learning the skills during ontogeny. There was no evidence for the first two hypotheses. However, we found a strong relationship between tool use specialization and mean female party size, which was used as a proxy for the opportunities for socially mediated learning in a foraging context during their development. This use was justified because females are rather philopatric and their mean party size remained stable over time, thus reflecting long-term tendencies. The correlation was not an artifact of a direct effect of party size on tool use tendencies, and did not hold for males, the dispersing sex. Thus, variation in the number of opportunities for social learning explains tool use variation within populations, corroborating hypotheses for between-population variation. The emergence of human culture was accompanied by vastly improved mechanisms of social learning. In order for these improvements to be favored by natural selection, the cultural potential must have actually been expressed. Thus, a combination of strong sociability and a reliance on tool-using or other technical skills acquired through social learning must have characterized early hominins.     

The development of wild immature Sumatran orangutans (Pongo abelii) at Ketambe

Orangutans have the longest immature period and inter-birth interval of all ape species. This may be explained by a slow life history, the need to develop skills or by their relatively solitary lifestyle, which prevents a mother from associating with two offspring. This study of wild immature orangutans at the Ketambe Research Station, Indonesia, describes, with partly cross-sectional, partly longitudinal data, their development to independence. The study subjects ranged from 1 to 11 years of age. Data on their activity budget, diet, mother–offspring proximity and maintenance of proximity, association with conspecifics and play behavior were collected. The results indicate that immature orangutans can provide for their own food and transport, and therefore were independent of direct maternal care, at an age of possibly 3 but more clearly 6 years. This is similar to chimpanzees, and refutes the slow life history hypothesis. Immature orangutans remain within their mother’s vicinity until the age of 8 years, indicating a dependence on indirect maternal care, and this coincides with the period during which the mother does not produce another offspring. A female orangutan seems unable to associate with an older immature while caring for a new infant. This is consistent with the solitary-lifestyle hypothesis and corroborates the results obtained with the Sumatran orangutan population at Suaq Balimbing. However, why an immature depends indirectly on its mother for such a long period remains unclear. It is possible that it needs to develop ecological or social skills or needs the protection of its mother. Unfortunately, no data were available to distinguish between these possibilities.     

Insect prey foraging strategies in Callicebus oenanthe in northern Peru

Titi monkeys (genus Callicebus) are small-bodied platyrrhines that supplement their predominantly frugivorous diet with variable amounts of leaves, seeds, and/or arthropod prey. Notable interspecific variation in the amount of insect prey in the diet has been observed in Callicebus, ranging from 0% to 20%. In this study, I investigate the degree and type of prey foraging in a little-known species, Callicebus oenanthe inhabiting a fragmented, secondary forest on the foothills of the Andes in northern Peru. I present data on prey type, prey search and capture techniques, substrate/vegetation use, foraging height, prey capture efficiency, and seasonal variation of insect prey foraging in one group of C. oenanthe observed from January to August 2005. Insect prey accounted for 22% of the diet, the highest amount reported for any Callicebus species to date, and insects from at least six different orders were included. C. oenanthe was mainly an investigative forager of hidden prey, manipulating easy-to-open substrates such as rolled up leaves, and hunted ant swarms and larger insects opportunistically. Insect foraging was predominant during the dry season (26%) and decreased during the wet season (13%). The study group foraged mostly in the understory (2-6 m) within vine-laden shrubs and trees, which may conform to an anti-predator strategy of crypticity. Overall the group had an 83% insect capture success rate. These data suggest that insect prey is an important part of the diet of C. oenanthe and may be especially notable during periods of resource scarcity. This study adds to the knowledge concerning insect prey foraging in Callicebus, which can have an important role in defining ecological strategies in the selection of secondary protein food resources within a given ecosystem.     

Orangutan diet,range, and activity at Tanjung Puting,Central Borneo

Previously, wild orangutan feeding and ranging behaviors have been described only from populations in hilly or mountainous regions. The Tanjung Puting study focuses on an orangutan population in a swampy lowland area near sea level. Tanjung Puting also differs from other areas in the virtual absence of large figs, which are significant orangutan food sources elsewhere. During a 4-year period and 6804 hr of observation, focal orangutans were recorded in 11,338 foraging bouts accounting for 3805 hr. Composition and phenology of the forest habitat were documented. The orangutans were predominantly frugivorous, with fruit-eating accounting for 61% of the foraging time. However, the overall variety in their diet was remarkable; 317 different food types have been identified, including fungus, insects, and honey. Orang-utans were strongly opportunistic foragers, with the composition of their diet varying markedly from month to month. During most months orangutans fed on a complex mix of fruit, leaves, bark, insects, and small vines. During some months fruit was not the major component of the diet. All orangutans foraged in both the dry-ground mixed dipterocarp forest and the peatswamp forest habitats found in their ranges. Adult males and females utilized different proportions of certain resources in their diets. Prime adult males also ranged further per day and spent more time on the ground than prime adult females. At Tanjung Puting contact with other orangutans usually increased a focal orangutan’s day length, day range, and amount of time spent moving. This suggests that foraging alone maximized each orangutan’s foraging returns by minimizing the day range traveled. Orangutan solitariness is the result of a large body size and of a predominantly frugivorous and opportunistic diet.     

Insectivory by Gorilla gorilla gorilla in Southeast Cameroon

Our study extends quantitative analyses of insect-eating by gorillas (Gorilla gorilla gorilla) to Cameroon. During a 2-mo period (May–June 2001), we recorded and analyzed feeding traces on plants and insects and in gorilla feces. We found 180 feeding traces, 17% of which involved insectivory. Seventy-eight percent of the fecal samples (n = 36) contained insects. Ants were found in 61% of the samples, termites in 39%, while 56% of the samples contained remains of other insects. We added 14 new species to the known insect diet of western lowland gorillas. Overall, social insects are predominant. The choice of prey by the Ntonga gorillas gives new clues for the existence of cross-cultural differences among gorilla populations. A comparison of the overall frequency of insectivory with those at other sites in Central Africa indicate a possible effect of forest disturbance on the insectivorous behavior of gorillas. The study suggests the existence of temporal variation in ant- and termite-eating by gorillas.     

Seasonal patterns of diet and ranging in two species of tamarin monkeys: Stability versus variability

From June through December, data were collected on the diet and ranging patterns of moustached (Saguinus mystax) and saddle-back (Saguinus fuscicollis) tamarin monkeys in the Amazon Basin of northeastern Peru. During this 7-month period, insects and nonleguminous fruits accounted for 83% of tamarin feeding and foraging time. Despite marked seasonal variation in rainfall and forest productivity, patterns of habitat utilization, day range, dietary diversity, resource exploitation, and activity budget remained relatively stable throughout the year. Moustached and saddle-back tamarins appear to solve problems of food acquisition and exploit patchily distributed feeding sites using a relatively limited set of foraging patterns. In general, these primates concentrate their daily feeding efforts on several trees from a small number of target plant species. These feeding sites are uncommon, produce only a small amount of ripe fruit each day, and are characterized by a high degree of intraspecific fruiting and flowering synchrony. Trees of the same species are frequently visited in succession, and individual feeding sites are revisited several times over the course of 1–2 weeks. This type of foraging pattern occurred during both dry and wet seasons and when exploiting fruit, nectar, legume, and exudate resources. Seasonal variation in the percentage of feeding and foraging time devoted to insectivory was also limited. In this investigation, there was no consistent evidence that temporal changes in overall forest fruit production had a major impact on the feeding, foraging, or ranging behavior of either tamarin species.     

Functional Invertebrate Prey Groups Reflect Dietary Responses to Phenology and Farming Activity and Pest Control Services in Three Sympatric Species of Aerially Foraging Insectivorous Birds

Farming activity severely impacts the invertebrate food resources of farmland birds, with direct mortality to populations of above-ground arthropods thorough mechanical damage during crop harvests. In this study we assessed the effects of phenological periods, including the timing of harvest, on the composition and biomass of prey consumed by three species of aerial insectivorous birds. Common Swifts Apus apus, Barn Swallows Hirundo rustica and House Martins Delichon urbica breed sympatrically and most of their diet is obtained from agricultural sources of invertebrate prey, especially from oil-seed rape crops. We categorized invertebrate prey into six functional groups, including oil-seed rape pests; pests of other arable crops; other crop-provisioned taxa; coprophilous taxa; and taxa living in non-crop and mixed crop/non-crop habitats. Seasonality impacted functional groups differently, but the general direction of change (increase/decrease) of all groups was consistent as indexed by prey composition of the three aerial insectivores studied here. After the oil-seed rape crop harvest (mid July), all three species exhibited a dietary shift from oil-seed rape insect pests to other aerial invertebrate prey groups. However, Common Switfts also consumed a relative large quantity of oil-seed rape insect pests in the late summer (August), suggesting that they could reduce pest insect emigration beyond the host plant/crop. Since these aerially foraging insectivorous birds operate in specific conditions and feed on specific pest resources unavailable to foliage/ground foraging avian predators, our results suggest that in some crops like oil-seed rape cultivations, the potential integration of the insectivory of aerial foraging birds into pest management schemes might provide economic benefits. We advise further research into the origin of airborne insects and the role of aerial insectivores as agents of the biological control of crop insect pests, especially the determination of depredation rates and the cascading effects of insectivory on crop damage and yield.     

Nutritional composition of actual and potential insect prey for the Kasekela chimpanzees of Gombe National Park,Tanzania

Humans, all great ape species, and some lesser apes consume insects. Insects can provide comparable nutritional yields to meat on a gram‐for‐gram basis and may serve as an important source of energy, fat, protein, minerals, and vitamins for hominoids. Although potential insect prey are abundant in ape habitats, patterns of insectivory are not consistent across species or populations. Efforts to understand these patterns are complicated by a lack of nutritional data. We collected samples of insects consumed by the Kasekela chimpanzee community of Gombe National Park, Tanzania, as well as of some insects found within the community range and ignored by these chimpanzees but known to be preyed upon by Pan elsewhere. We determined the gross energy (GE), estimated metabolizable energy (ME), fat, protein, fiber, and ash content of these samples following standard methodologies. We use these data to test the hypothesis that Kasekela chimpanzees choose insect prey (at least in part) based on energy and/or macronutrient content. On a fresh‐weight, per‐gram basis, the insect prey consumed by Kasekela chimpanzees had significantly higher fat and lower ash content than other assayed insects, and on a fresh‐weight, per‐foraging‐unit (“per‐insect,” “per‐dip,” or “per‐nest”) basis were significantly higher in GE, fat, and protein. On a per‐gram basis, the assayed insects were generally comparable in energy and macronutrients to wild vertebrate meat. We conclude that Kasekela chimpanzees do favor insects that are high in energy, fat, and protein, and that the potential macronutrient yields from some forms of insectivory are not trivial. © 2012 Wiley Periodicals, Inc.     

Adaptation to Forest Life During the Reintroduction Process of Immature <Emphasis Type="Italic">Pongo abelii</Emphasis>

Reintroduction of Sumatran orangutans can be an important tool to conserve the species in the face of the current decline of wild populations. Monitoring ex-captive orangutans during the reintroduction process provides insight into their adaptation to forest life and is important for evaluating the success of such programs. We investigated activity patterns, food choice, height use, and nest-building abilities of 8 immature orangutans at a reintroduction station in Sumatra. All focal individuals spent most of their time in the forest feeding, but at a lower proportion than wild conspecifics. Among the focal individuals, the behavior of orangutans that avoided human contact differed in several aspects from those that were human bonded. Their diet was composed mainly of fruits, similar to the food choice of wild orangutans, and their ground-avoidance and superior nest-building abilities indicated more effective predation-avoidance behavior vs. that of human-bonded animals. Finally, these immatures showed regular social contact with more experienced conspecifics, which ultimately may facilitate the social learning of local feral expertise. Human-bonded individuals, in contrast, preferentially stayed on the ground and at low heights and rarely built nests, thereby increasing predation risk. Their choice of leaves over fruits as foods indicated a less effective foraging strategy. Nevertheless, this group exhibited individual learning trends toward an adult-like foraging pattern and height use. We conclude that social interactions with conspecifics and humans, especially during an early developmental period, can affect adaptation to forest life and probably also influence the success of orangutan reintroduction processes.     

Diurnal foraging‐mode shifts and food availability in nectarivore assemblages during winter

Abstract Temporal fluctuations of food resources and foraging activities have been studied extensively, especially at longer timescales (monthly, seasonally, among years). However, short‐term variation (e.g. within days) is less well understood. Here we systematically quantified diurnal patterns of foraging by nectarivorous birds (meliphagid honeyeaters) that numerically dominated stands of a winter‐flowering eucalypt, the red ironbark, Eucalyptus tricarpa, in central Victoria, Australia. Diurnal variation in food resources also was measured. Data were collected in winter. Anecdotal observations that honeyeaters change from almost exclusive nectarivory early in the day to a higher fraction of insectivory – especially aerial hawking – later in the day were confirmed, although in areas of high flowering intensity, nectar‐feeding remained the dominant foraging activity throughout the day. Local climatic factors (ambient temperature, windiness and cloud cover) all varied systematically through the day. Together, results were consistent with a change in foraging emphasis to greater insectivory as a function of elevated activities of insects (especially aerial ones), which was probably fostered by higher ambient temperatures. Contrary to energetic expectations, the nectarivores were very active early in the morning when ambient temperatures averaged approximately 3°C, well below thermoneutral temperatures. We deduced that the potential benefits of gathering as much energy‐rich nectar as possible before it was depleted outweighed the high costs of activities at low temperatures.     

Context-dependent specialisation drives temporal dynamics in intra- and inter-individual variation in foraging behaviour within a generalist bird population

Individual niche variation is common within animal populations, and has significant implications for a wide range of ecological and evolutionary processes. However, individual niche differences may also temporally vary as a result of behavioural plasticity. While it is well understood how niche variation is affected by changes in resource availability, comparatively little is known about the extent to which individual niche differences may vary within the annual cycle due to internal drivers. Here, we assess how time- and energy-constraints imposed by incubating and brood rearing affect inter- and intra-individual variation in the foraging behaviour of lesser black-backed gulls, a generalist seabird with strong individual niche variation. To this end, we compared daily foraging trips of 22 breeding and 23 non-breeding GPS-tracked adult gulls from two colonies in the Southern Bight of the North Sea over the course of the breeding season. We find that breeding birds, unlike non-breeding ones, did indeed alter their foraging behaviour during the breeding season. Both sexes reduced their searching effort by increasingly revisiting earlier foraging locations, allowing for shorter and more frequent foraging trips. Breeding females also showed pronounced shifts in their habitat use and strongly specialised on urbanised foraging habitats throughout the breeding season. Hence, while individual variation in habitat use remained largely consistent within non-breeders and in breeding males, individual variation among breeding females almost completely disappeared. Female lesser black-backed gulls are on average smaller, and therefore often outcompeted by males for the most profitable food sources. The temporal specialisation on spatially reliable anthropogenic food sources during breeding hence suggests a complex interplay between intrinsic competitive constraints, resource reliability and shifting time- and energy budges in shaping temporal dynamics in individual niche variation within our study population.     

Strengthened insectivory in a temperate fragmented forest

Habitat fragmentation modifies ecological patterns and processes through changes in species richness and abundance. In the coastal Maulino forest, central Chile, both species richness and abundance of insectivorous birds increases in forest fragments compared to continuous forest. Through a field experiment, we examined larvae predation in fragmented forests. Higher richness and abundance of birds foraging at forest fragments translated into more insect larvae preyed upon in forest fragments than in continuous forest. The assessed level of insectivory in forest fragments agrees with lower herbivory levels in forest fragments. This pattern strongly suggests the strengthening of food interactions web in forest fragments of coastal Maulino forest.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Begging for information: mother-offspring food sharing among wild Bornean orangutans

Transfer of solid food from mothers or other adults to dependent offspring is commonly observed in various primate species and both nutritional and informational benefits have been proposed to explain the function of such food sharing. Predictions from these hypotheses are tested using observational data on wild orangutans (Pongo pygmaeus wurmbii) at Tuanan, Central Borneo, Indonesia. In 1,145 hr of focal observation and 458 recorded food interactions between four pairs of females with offspring it was found that virtually all transfers were initiated by the offspring and that younger infants solicited food more often and did so for a greater variety of items than older offspring. All offspring primarily solicited food that was difficult to process, i.e., inaccessible to them. Furthermore, the amount of food solicitation was negatively correlated with ecological competence. Hence food sharing seemed to be related to an offspring's skill level, as suggested by the informational hypothesis. In contrast, offspring did not solicit high-quality items more than low-quality items and food sharing did not peak around the age of weaning, as predicted by the nutritional hypothesis. Mothers were usually passively tolerant, allowing offspring to take food but hardly ever provisioned. Parent-offspring conflict concerning food sharing was only observed well after weaning. Thus, by taking food directly from the mother, young orangutans were able to obtain information about the affordances and nutritional value of food items that were otherwise out of their reach and could familiarize themselves with the mother's diet. In species such as orangutans or other apes, characterized by a broad diet that requires extractive foraging, informational food transfer may be vital for an immature to acquire complex feeding skills and adult diet.     

Cross-site differences in foraging behavior of white-faced capuchins (Cebus capucinus)

Researchers have identified a variety of cross-site differences in the foraging behavior of free-ranging great apes, most notably among chimpanzees (Pan troglodytes) and more recently orangutans (Pongo pygmaeus), that are not due to obvious genetic or ecological differences. These differences are often referred to as "traditions." What is not known is whether this high level of interpopulation variation in behavior is limited to hominoids. In this study, we use long-term data from three Costa Rican field sites that are geographically close and similar ecologically to identify potential foraging traditions in white-faced capuchins (Cebus capucinus). Foraging traditions are predicted in Cebus because of many behavioral and morphological convergences between this genus and the great apes. The processing techniques used for the same food species were compared across sites, and all differences found were classified as present, habitual, or customary. Proximity data were also analyzed to determine if social learning mechanisms could explain variation in foraging behavior. Of the 61 foods compared, we found that 20 of them are processed differently by capuchins across sites. The differences involve pound, rub, tap, "fulcrum," "leaf-wrap," and "army ant following." For most of the differences with enough data to analyze, the average proximity score of the "matched" dyads (two individuals within a group who shared a "different" processing technique) was statistically higher than the average proximity score of the remaining "unmatched" dyads.     

The insectivorous-frugivorous diet of the golden-handed tamarin (Saguinus midas midas) in French Guiana

The diet of Saguinus midas is characterised by analysis of stomach contents samples on an annual and seasonal basis. The diet comprises significant proportions of both fruit and insects all year round, although fruit use shows important seasonal variations. The tamarins reduce their use of fruit resources during the peak fruiting season and instead exploit increased opportunities for insectivory. This is suggested to be a response to differences in gross fruit availability as a function of both fruiting patterns and the presence of competitors for the available fruit resources.