Ecological Correlates of Ranging Behavior in Bearded Sakis (Chiropotes sagulatus) in a Continuous Forest in Guyana

Group size and the distribution and quality of food resources are among the most important determinants of primate ranging behavior. In this study, I use the framework of the ecological constraints model to assess correlates of range size of a free-ranging group of bearded sakis (Chiropotes sagulatus). Bearded sakis are among the widest ranging neotropical primates, yet the lack of data from continuous forest populations has made understanding the factors influencing such large ranges difficult. I collected data on ranging behavior and diet during 44 full-day follows over 15 mo. The focal group used a home range of ca. 1000 ha and had daily path lengths of 2.8–6.5 km (mean?=?4.0 km). Daily path length did not significantly correlate with group size, patch quality, food availability, or the spatial distribution of feeding trees. Monthly home range size significantly positively correlated with group size and patch quality. The focal group had significantly shorter paths when ripe fruit consumption was higher and had more diverse diets, visited more food patches, and used larger monthly home ranges when they consumed a higher percentage of seeds. The results of this study, combined with other recent studies of Chiropotes in continuous forest, suggest that large home ranges (approaching 1000 ha) are characteristic of the genus. Although range size may be related to group size and food patch size, I suggest nutrient mixing and the need to balance the effects of seed secondary compounds as additional explanations for the large ranges of bearded sakis.     

Pithecia pithecia's behavioral response to decreasing fruit abundance

In this article we describe the behavioral responses of a group of white-faced sakis' (Pithecia pithecia) to fruit and water scarcity. Six sakis were observed on Round Island in Guri Lake, Venezuela, between March and May 1996. These months are considered the dry season and the beginning of the wet season. Sakis specialize in eating seeds. During the present study only one tree species, Licania discolor (Chrysobalanaceae), fruited in substantial numbers. Licania seeds accounted for 88% of the time the sakis spent eating fruit in March, 87% in April, and 80% in May. We estimate that the sakis' intake of Licania seeds dropped from 2,573 seeds in the 15-day observation period in March to 956 seeds in the 16-day observation period in May. The sakis not only spent less time eating Licania, they ate the seeds at a much slower rate. The drop in the sakis' feeding rate was probably due to increased local search and inspection times. In response to the scarcity of fruit, the sakis ate more young leaves, insects, and flowers. Feeding bouts became more frequent but shorter. Mean distances between feeding bouts fell significantly and the sakis revisited trees less often.     

Fruit Finding by Mangabeys (Lophocebus albigena): Are Monitoring of Fig Trees and Use of Sympatric Frugivore Calls Possible Strategies?

Frugivorous forest primates face a continual challenge to locate ripe fruit due to the poor visibility characterizing a heavily vegetated habitat and the spatial and temporal unpredictability of their fruit sources. We present two hypotheses regarding fruit finding in gray-cheeked mangabeys (Lophocebus albigena). The first hypothesis is that mangabeys monitor nonfruiting fig trees by visiting and checking them for fruit at a higher rate than control trees that do not produce preferred fruit. We test this hypothesis by comparing rates of visitation to focal fig trees and control trees. The second hypothesis is that mangabeys use sympatric frugivore loud calls to locate fruit sources. We test this hypothesis (1) observationally, by comparing the rates at which mangabeys visit calling sites of sympatric frugivores and matched control areas; and (2) experimentally, by following mangabey responses to playbacks of tape-recorded calls: the black-and-white-casqued hornbill (Bycanistes subcylindricus) long call, the great blue turaco (Corythaeola cristata) rattling kok, the adult male mangabey whoopgobble, and the chimpanzee (Pan troglodytes) pant hoot. We tested the hypotheses via data from a single group of mangabeys in the Kibale National Park, Uganda. There is no evidence that mangabeys monitor fig trees for the presence of fruit, but they may use the calls of hornbills to locate fruit. Statistical evidence that mangabeys use conspecific whoopgobbles and chimpanzee pant hoots in fruit finding is lacking, though anecdotal observations suggest this possibility. There is no evidence for use of turaco calls in fruit finding.     

Diet and Food Choice in Peruvian Red Uakaris (<Emphasis Type="Italic">Cacajao calvus ucayalii</Emphasis>): Selective or Opportunistic Seed Predation?

Even primates considered dietary specialists tend to eat a combination of fruit pulp, seeds, other plant parts, or animals. Specialist seed predators could either feed on seeds preferentially, or to avoid competition when ripe pulps are scarce. Pitheciin monkeys have specialized dentition that allows them to feed on seeds protected by hard shells, and the upper limit on the hardness of these is likely to be a function of jaw size. We recorded the diet of Peruvian red uakaris (Cacajao calvus ucayalii) on the Yavari River, Peru, to test the prediction that this seed predator would feed on the seeds of hard-shelled fruits preferentially over softer ones in relation to their availability in the forest. We also tested predictions that adult male, adult female, and juvenile diets would differ, with larger individuals eating more hard fruits. Uakaris ate 55.4% seeds, 38.9% pulps and arils, and 5.6% other items, but proportions varied through the year. More pulps, especially from the palm Mauritia flexuosa, were eaten when fruit availability was low, and more hard fruits were positively selected for than softer ones. Juveniles did not open the hardest fruit species opened by adults, and adult males ate harder fruits than females. These results provide evidence that seed eating in some primates has evolved beyond a means of avoiding competition for the ripe pulps typically preferred by many primates. Specialist seeding-eating primates therefore occupy divergent niches that require separate consideration from those of similar-sized primates.     

Seed predation by monkeys and macaws in eastern Venezuela: Preliminary findings

Feeding data collected concurrently on bearded sakis (Chiropotes satanas) and red and green macaws (Ara chloroptera) on a large island in Guri, Venezuela provides preliminary evidence that these two seed predators have similar diets. Individuals of both species were equally capable of opening very hard, protected fruits. Of the seven fruit species used by macaws during the study period, four species were also ingested by sakis at the same stage of ripeness, two species were ingested at different stages of ripeness (macaws earlier than sakis), and one species was never observed to have been eaten by sakis. The second finding, that macaws ingest young seeds of the Anacardiaceae and Burseraceae families and the bearded sakis ingest only the ripe mesocarp of these species suggests that the most distinguishable difference in their diets might be a tolerance of toxins by the macaws that act as feeding deterrents for the monkeys. Although we did not document the location of local clay licks in eastern Venezuela, the use of clay licks by macaws in Peru (Munn, 1992) suggests that this activity (that is not practiced by the sakis) may be helpful in detoxifying or ameliorating the effects of ingesting chemically protected fruit. deceased.     

Dispersal of a Human-Cultivated Crop by Wild Chimpanzees (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) in a Forest–Farm Matrix

With the conversion of natural habitats to farmland, nonhuman primates (hereafter primates) are increasingly exposed to agricultural crops. Although frugivorous primates are important seed dispersers that sometimes feed on agricultural fruits, evidence for dispersal of crops by primates is lacking. Here, we examine flexible feeding on cacao (Theobroma cacao) fruit and seed dispersal patterns by chimpanzees (Pan troglodytes verus) at Bossou in Guinea, and consequent cacao germination and survival. From direct observations, we confirm that cacao fruit is not an important food to chimpanzees, representing 0.23 % of focal animal feeding time. Chimpanzees ingest cacao pulp and either spit out the large seeds intact from unripe cacao fruit or swallow the seeds from ripe cacao fruits, which are consequently deposited in feces. From ecological surveys we show that chimpanzees distributed cacao extensively throughout their home range, at a mean distance of 407 m?±?SE 0.6 (N?=?90 clusters, range: 4–1130 m) from cacao plantations. As distance from the cacao plantation increased, cacao plants were more likely to survive. Other factors, including number of cacao plants in a cluster, plant height, and openness of the understory did not predict short-term cacao survival. Cacao plants within the forest did not produce fruit. By contrast, when chimpanzees deposited seeds in a plantation, cacao plants produced fruits as a result of farmers’ maintenance of the area. Our local-scale findings emphasize the complex behavioral and ecological interconnections between coexisting humans and primates in agricultural landscapes and generate interesting questions regarding primate niche construction and crop “ownership” related to who “plants” the crop.     

Notes on the ecology of buffy saki monkeys (Pithecia albicans,Gray 1860): A canopy seed-predator

Data on the ecology and diet of buffy sakis (Pithecia albicans) were obtained during a 20-month study in an entirely undisturbed terra firme forest in the upper Urucu river, Amazonas, Brazil. Groups of 3–7 sakis found in a 900-ha study plot used large home ranges (147–204 ha), which overlapped extensively with those of neighboring groups. Similar to other pitheciines, buffy sakis were primarily seed predators, relying heavily on young seeds of certain key plant families, such as the Sapotaceae and Leguminosae. Ripe fruits, ripe seeds, young leaves, flowers, and nectar were eaten to a lesser extent. Whether or not feeding, sakis spent most of their time in the canopy and subcanopy, a pattern similar to that of other southwestern Amazonian saki species, but sharply different from that of Guianan sakis (Pithecia pithecia), which use considerably lower levels of the forest. Comparisons are made between different Pithecia species to show whether and how P. albicans diverges ecologically from its congeners. © 1993 Wiley-Liss, Inc.     

Western Purple-faced Langurs (Semnopithecus vetulus nestor) Feed on Ripe and Ripening Fruits in Human-modified Environments in Sri Lanka

Although most colobines feed mainly on leaves and a few feed heavily on seeds, colobine digestive adaptations for folivory are thought to preclude the high use of ripe fleshy fruits. In this long-term study of Semnopithecus vetulus nestor, the endemic western purple-faced langur of Sri Lanka, I investigated the feeding ecology and dietary flexibility for fruit feeding in 2 free-ranging groups (PT1 and R1) living in human-modified environments with abundant cultivated fruit, at Panadura and Piliyandala, for 19 mo and 13 mo respectively, using scan-sampling, vegetation enumeration, and phenological studies. In contrast to folivorous forest-living colobines, including other subspecies of Semnopithecus vetulus, my focal groups used more fruit (>50%) than foliage (PT1: 36%; R1: 34%). Both groups used many plant species (PT1 115; R1 59), but selected their food species, fruits over leaves, and young leaves over mature leaves. Fruit use was independent of young leaf availability. Notably, 78.4% and 83.4% of fruits consumed by PT1 and R1 were fleshy and human-edible, most of which were ripening or ripe (PT1: 72.4%; R1: 94.8%). The main fruit for both groups was Artocarpus heterophyllus (Moraceae; jakfruit), a cultivar with fleshy fruit. These findings differ from previous understanding of colobine diets. I suggest that environmental factors, such as the abundance and nature of available fruits, and the absence of arboreal-primate fruit competitors, could influence the use of ripe fleshy fruits by colobines strongly, highlighting the need to review the dietary and digestive flexibility of this group in changed and changing natural environments to formulate effective conservation action.     

Factors Affecting Nesting Site Choice in Chimpanzees at Tshibati,Kahuzi-Biega National Park: Influence of Sympatric Gorillas

We recorded 310 fresh chimpanzee night nests at 72 nest sites to determine their choice of tree and site for nesting vis-à-vis the effects of sympatric gorillas. Chimpanzees did not use trees for nesting according to their abundance, but instead tended to nest in fruit trees that they used as food sources. Nesting patterns of chimpanzees may vary with nesting group size, the type of vegetation, and fruit species eaten or not eaten by gorillas. When chimpanzees lodged as a small group in the secondary forest, they nested more frequently in trees bearing ripe fruits eaten only by themselves than in those with fruit eaten also by gorillas. When they lodged as a large group in the primary forest, they nested more frequently in trees bearing ripe fruits eaten by both apes. Nest group size is positively correlated with the availability of preferred ripe fruits in secondary forest. These findings not only reflect the larger foraging groups at the larger fruiting trees but also suggest that chimpanzees may have tended to occupy fruiting trees effectively by nesting in them and by forming large nest groups when the fruits attracted gorillas. Competition over fruits between gorillas and chimpanzees, due to their low productivity in the montane forest of Kahuzi, may have promoted the chimpanzee tactics.     

Primates take weather into account when searching for fruits

Temperature and solar radiation are known to influence maturation of fruits and insect larvae inside them . We investigated whether gray-cheeked mangabeys (Lophocebus albigena johnstonii) of Kibale Forest, Uganda, take these weather variables into account when searching for ripe figs or unripe figs containing insect larvae. We predicted that monkeys would be more likely to revisit a tree with fruit after several days of warm and sunny weather compared to a cooler and more cloudy period. We preselected 80 target fig trees and monitored whether they contained ripe, unripe, or no fruit. We followed one habituated monkey group from dawn to dusk for three continuous observation periods totalling 210 days. Whenever the group came within a 100 m circle of a previously visited target tree for a second time, we noted whether or not individuals proceeded to the trunk, i.e., whether they "revisited" or simply "bypassed" the tree. We found that average daily maximum temperature was significantly higher for days preceding revisits than bypasses. The probability of a revisit was additionally influenced by solar radiation experienced on the day of reapproach. These effects were found only for trees that carried fruit at the previous visit but not for trees that had carried none. We concluded that these nonhuman primates were capable of taking into account past weather conditions when searching for food. We discuss the implication of these findings for theories of primate cognitive evolution.     

Tracking Bat‐Dispersed Seeds Using Fluorescent Pigment1

Tracking the dispersal of seeds by fruit‐eating animals in tropical rain forests is crucial to further our understanding of plant–frugivore interactions and their impacts upon forest regeneration and plant population dynamics. We tested the feasibility of tracking bat‐dispersed seeds in a Philippine lowland rain forest with the help of fluorescent pigment. The powder was mixed with acetone and sprayed to ripe fruits of fig trees, i.e., Ficus septica and F. variegata. During nightly monitoring using a hand‐held ultraviolet lamp bat deposits (seed‐containing spat outs and feces) could successfully be detected. Distances and directions of deposit sites to the focal trees were recorded and seed shadow areas were analyzed. Bats dispersed most of the seeds less than 50 m away from the parent plants resulting in seed shadow areas < 0.30 ha in size. An in situ fruit preference experiment showed that fluorescent powder is unlikely to deter bats from feeding on ripe figs. In conclusion, the technique is simple, inexpensive, noninvasive, applicable to different fields of research and allows one to follow the fate of seeds from known sources.     

Dietary Response of Chimpanzees and Cercopithecines to Seasonal Variation in Fruit Abundance. I. Antifeedants

In order to understand dietary differentiation among frugivorous primates with simple stomachs, we present the first comparison of plant diets between chimpanzees and cercopithecine monkeys that controls for food abundance. Our aim was to test the hypothesis that monkeys have a more diverse diet as a result of their dietary tolerance for chemical antifeedants. Our study species are chimpanzees, blue monkeys, redtail monkeys, and gray-cheeked mangabeys living in overlapping ranges in Kibale National Park, Uganda. We indexed food abundance by the percentage of trees having ripe fruit within the range of each group; it varied widely during the year. Chimpanzees spent almost 3 times as much of their feeding time eating ripe fruits as the monkeys did and confined their diets almost exclusively to ripe fruits when they were abundant. Monkeys maintained a diverse diet at all times. When ripe fruit was scarce chimpanzee and monkey diets diverged. Chimpanzees relied on piths as their main fallback food, whereas monkeys turned to unripe fruits and seeds. For each primate group we calculated the total weighted mean intake of 5 antifeedants; condensed tannins (CT), total tannins assayed by radial diffusion (RD), monoterpenoids (MT), triterpenoids (TT), and neutral-detergent fiber (NDF). Monkeys had absolutely higher intakes of CT, RD, MT, and TT than those of chimpanzees, and their intake of NDF did not differ from that of chimpanzees, appearing relatively high given their lower body weights. However contrary to expectation, dietary divergence during fruit scarcity was not associated with any change in absolute or relative intake of antifeedants. For example, fruit scarcity did not affect the relative intake of antifeedants by cercopithecines compared to chimpanzees. Our results establish chimpanzees as ripe-fruit specialists, whereas cercopithecines are generalists with a higher intake of antifeedants. The low representation of ripe fruits in the diets of cercopithecines has not been explained. An important next step is to test the hypothesis that the difference between Kibale chimpanzees and cercopithecines represents a more general difference between apes and monkeys.     

Responses of dispersal agents to tree and fruit traits in Virola calophylla (Myristicaceae): implications for selection

Variation in traits affecting seed dispersal in plants has been attributed to selection exerted by dispersal agents. The potential for such selection was investigated in Virola calophylla (Myristicaceae) in Manú National Park, Peru, through identification of seed dispersal agents and of tree and fruit traits significantly affecting the quantity of seeds dispersed. Seventeen bird and one primate species (the spider monkey, Ateles paniscus) dispersed its seeds. Spider monkeys dispersed the majority of seeds (a minimum of 83% of all dispersed seeds). Visitation by dispersal agents depended only on the quantity of ripe fruit available during a tree observation. In contrast, seed removal increased with both greater quantity of ripe fruit and aril: seed ratio. When analyzed separately, seed removal by birds increased only with greater aril: seed ratio, whereas seed removal by spider monkeys was affected by the quantity of ripe fruit and phenological stage. The finding that dispersal agents responded differently to some tree and fruit traits indicates not only that dispersal agents can exert selection on traits affecting seed dispersal, but also that the resulting selection pressures are likely to be inconsistent. This conclusion is supported by the result that the proportion of the seed crop that was dispersed from individual trees, which accounted for cumulative dispersal by all agents, was not influenced by any tree or fruit trait evaluated. Comparing these results with those from studies of V. sebifera and V. nobilis in Panama revealed that the disperser assemblages of these three Virola species were congruent in their similar taxonomic representation. In Panama the proportion of V. nobilis seed crop dispersed was related positively to aril: seed ratio and negatively to seed mass, a result not found for V. calophylla in Peru. The greater importance of dispersal by primates versus birds in V. calophylla, relative to V. nobilis, may explain this difference. Thus, variation in disperser assemblages at regional scales can be another factor contributing to inconsistency in disperser-mediated selection on plant traits.     

Feeding habits of chimpanzees (Pan troglodytes), red-tail monkeys (Cercopithecus ascanius schmidti) and blue monkeys (Cercopithecus mitis stuhlmanii) on figs in Budongo Forest Reserve, Uganda

Feeding habits of chimpanzees, red‐tail and blue monkeys on figs (Ficus) were studied in compartment N3 of Budongo Forest Reserve, western Uganda, from September 1997 to March 1998. The aim was to examine the spatial and temporal foraging habits of chimpanzees, red‐tail monkeys and blue monkeys on figs in the forest reserve. Both scan and focal sampling methods were used to assess the foraging habits of the primates. It was found that the primates fed on emerging leaves of Ficus mucuso Ficalho, F. varifolia Warb. and F. exasperata Vahl. They also preferred ripe fruits to emerging, young and unripe fruits. The primates spent 78% of the morning eating fruits and leaves and inhabited fig trees with fruits for about 4 h. Fig trees with ripe fruits attracted larger numbers of primate groups. It is concluded that information on the feeding habits of chimpanzees and monkeys is required in order to have a clear understanding of the social behaviour and pattern of movement of the primates and to assist in predicting the likely impacts of poor forest management, forest degradation and loss of food resources on their populations.     

Fruit and seed ecology of wild robusta coffee (Coffea canephora Froehner) in Kibale National Park, Uganda

Coffee fruit production, frugivorous activities and seed longevity were investigated during two seasons. The average densities of fruiting coffee trees during the first sampling period ranged from four to seven per 25 m² and during the second sampling period, four fruiting trees per 25 m² were estimated. Fruiting coffee trees were significantly less than the non-fruiting trees ( P < 0.05, t -test). Periods of maximum coffee fruit fall coincided with those of lowest rainfall. Coffee fruit yield per tree from ground collections was 7–40 m⁻² in 1992 and 16–38 m⁻² in 1993/1994. Split coffee berries ranged between 71.7% and 83.6% of all fruits collected from the forest floor in the 1993/94 sampling period. Coffee seeds collected from the forest floor were mostly undamaged. Black and white colobus and redtail monkeys were observed to feed on ripe coffee fruits but did not crush the seeds. Coffee seed viability declined rapidly during storage. Seeds left on the forest floor survived longest, those stored under laboratory conditions lost viability fastest, and those in cold storage showed intermediate longevity.     

Seed handling in chimpanzees (Pan troglodytes) and redtail monkeys (Cercopithecus ascanius): Implications for understanding hominoid and cercopithecine fruit-processing strategies and seed dispersal

Primates are confronted with an array of constraints in feeding on fruit, including the removal of adhesive, energy-rich pulp from seeds. In this paper, I discuss how primates meet this challenge and present data on the fruit-processing and seed-handling behavior of chimpanzees and redtail monkeys in Kibale National Park, Uganda. These data are then related to these species' services as seed dispersers. Particular attention was paid to the methods by which primates removed pulp from seeds, the density of seed clumps that they deposited (by spitting, dropping, or defecating) to the forest floor, and the distance seeds were moved from parent trees. Distance and density differences in chimpanzee and redtail seed dispersal resulted from distinct fruit-processing and seed-handling methods. It was observed, in general, that redtail monkeys engaged in fine oral processing and were seed spitters: most seeds were dispersed in close proximity to parent trees (84% of spat seeds <10 m of parent tree), and deposited singly (100% seeds spat singly). In contrast, chimpanzees were coarse fruit processors and seed swallowers: seeds were defecated in denser clumps (e.g., a mean of 149 large seeds/dung sample and hundreds of small seeds/dung sample), far from parent trees. I evaluate the factors that shape patterns of fruit processing in hominoids and cercopithecines, and argue that the observed seed handling differences can be attributed to differences in digestive retention times, oral anatomy, and alternative mechanisms by which to avoid the cost of seed ballast. Am J Phys Anthropol 109:365–386, 1999. © 1999 Wiley-Liss, Inc.     

Beans with bugs: Covert carnivory and infested seed selection by the red-nosed cuxiú monkey

Members of the Neotropical primate genus Chiropotes eat large volumes of immature seeds. However, such items are often low in available proteins, and digestion of seeds is further inhibited by tannins. This suggests that overall plant-derived protein intake is relatively low. We examined the presence of insect larvae in partially eaten fruits, compared with intact fruit on trees, and examined fecal pellets for the presence of larvae. We found that red-nosed cuxiú (Chiropotes albinasus) individuals may supplement their limited seed-derived protein intake by ingesting seed-inhabiting insects. Comparison of fruits partially eaten for their seeds with those sampled directly from trees showed that fruits with insect-containing seeds were positively selected in 20 of the 41 C. albinasus diet items tested, suggesting that fruits with infested seeds are actively selected by foraging animals. We found no differences in accessibility to seeds, that is, no differences in husk penetrability between fruits with infested and uninfested seeds excluding the likelihood that insect-infestation results in easier access to the seeds in such fruits. Additionally, none of the C. albinasus fecal samples showed any evidence of living pupae or larvae, indicating that infesting larvae are digested. Our findings raise the possibility that these seed-predating primates might provide net benefits to the plant species they feed on, since they feed from many species of plants and their actions may reduce the populations of seed-infesting insects.     

Ranging behavior and foraging ecology of lowland woolly monkeys (Lagothrix lagotricha poeppigii) in Yasuní National Park,Ecuador

In a year-long study, I investigated the ranging behavior of lowland woolly monkeys (Lagothrix lagotricha poeppigii) in a terra firma rainforest in Yasuní National Park, Ecuador, and examined the relationship between ranging, diet, food availability, and food patch use for this population. In Yasuní the total home range sizes for two social groups were 124 and 108 ha, which are much smaller than has been reported previously for Lagothrix elsewhere in its geographic distribution. The mean yearly day range estimates for these same groups were 1,792 m and 1,878 m, which are well within the range of variation previously reported. Ranging behavior was not correlated with the current habitat-wide abundance of ripe fruit, which comprises 76.3% of the yearly diet for this population, but was associated with one measure of likely insect prey abundance and with the availability of immature fruits, a minimal part of the diet. Specifically, one study group moved significantly greater distances during months of high likely insect prey abundance and when immature fruits were abundant. The second study group also traveled farther when likely insect prey abundance was high and when immature fruits were abundant, although the latter relationship only approached significance. This group also devoted significantly more of its daily activity budget to travel during these times. These results indicate that variation in ripe fruit abundance makes no meaningful contribution to explaining variation in ranging behavior for this population of woolly monkeys. Instead, the results raise the possibility that some aspects of the ranging behavior of frugivorous primates may be related to the availability of alternative food sources, such as animal prey, or to monitoring the phenological status of important fruit trees, rather than simply reflecting the degree of intragroup feeding competition.     

Alternative seed-handling strategies in primates: seed-spitting by long-tailed macaques (Macaca fascicularis)

Summary The seeds in fruits consumed by primates may be chewed and digested, swallowed and defecated intact, or separated from the flesh and spat out. We show by a combination of close field observations and experiments with caged animals, that long-tailed macaques (Macaca fascicularis) have a remarkably low threshold of 3–4 mm for swallowing seeds and also that wild macaques rarely break them. The seeds of 69% of the ripe fruit species eaten are spat out intact or cleaned outside the mouth and dropped. Seed-spitting significantly reduces the swallowed food bulk and may lessen the risk of releasing seed toxins during mastication. However, it requires that even small fruits are processed in the mouth one or a few at a time. We suggest that fruit storage in the cheek pouches of cercopithecine monkeys allows them to spit seeds individually without excessively slowing fruit intake while feeding on patchily distributed fruit. In contrast, Apes and New World monkeys apparently swallow and defecate most ripe seeds in their diet and colobine monkeys break and digest them, detoxifying seed defenses by bacterial fermentation.     

Intratree Variation in Fruit Production and Implications for Primate Foraging

We tested the hypothesis that fruit quantity and quality vary vertically within trees. We quantified intratree fruit production before exploitation by frugivores at different heights in 89 trees from 17 species fed on by primates in Kibale National Park, Uganda. We also conducted a pilot study to determine if the nutritional value of fruit varied within tree crowns. Depending on the species and crown size, we divided tree canopies into 2 or 3 vertical layers. In 2-layered trees, upper crowns produced fruits that were 9.6–30.1% bigger and 0.52–140 times the densities of those from lower crowns, with one exception. Among 2-layered trees, upper crowns produced a mean of 46.9 fruits/m³ (median 12.1), while lower crowns produced a mean of 14.1 fruits/m³ (median 2.5). Among 3-layered trees, upper crowns produced a mean density of 49.9 fruits/m³ (median 12.5), middle crowns a mean of 16.8 fruits/m³ (median 6.6), and lower crowns a mean of 12.8 fruits/m³ (median 1.8). Dry pulp and moisture were systematically greater per fruit in the highest compared to the lowest canopy layers (22.4% and 16.4% respectively in 2-layered trees, 49.7% and 21.8% respectively in 3-layered trees). In 1 tree of Diospyros abyssinica, a pilot nutritional study showed that upper crown ripe fruit contained 41.9% more sugar, 8.4% more crude proteins, and 1.8 times less of the potentially toxic saponin than lower crown ripe fruit, but the result needs to be verified with more individuals and species of trees. We discuss the consequences of intratree variations in fruit production with respect to competition among frugivorous primates.