Influence of food dispersion on feeding activity and social interactions in captive Lophocebus albigena and Cercocebus torquatus torquatus

We analysed the impact of the distribution of food items on feeding activity and social interactions in captive red-capped and grey-cheeked mangabeys. Three different feeding situations were presented: food items were either placed in a single heap or dispersed in several smaller heaps on the ground or in space. Social interactions were estimated by the frequency of positive, as well as of negative social interactions and by the structure of visual social attention. Feeding activity was estimated by proximity to food and feeding frequency. When food items were presented in single heaps, the adult males monopolized the food and monitoring of conspecifics increased. Social interactions and social gazes decreased in numbers when food items were dispersed in 3D. Gazes were directed more frequently towards the adult males when food items were presented in a single heap. Juveniles and some nonreproductive adult monkeys were the most affected by food competition situations. Inter-group variations, within a given species, of the behavioural responses observed in relation to the distribution of food items were evidenced. This suggests that social context, i.e., individual histories and relationship between group members, plays an important part in the expression of the activities of each member in a group.     

Aimed Throwing as a Means of Food Transfer Between Tufted Capuchins (Cebus apella)

We examined aimed throwing as a means of food transfer in tufted capuchins (Cebus apella). We conducted this research in three phases. In Phase 1 we provided food to monkeys in one of two groups housed 1 m apart. We did not provide food to subjects in the second group. An observer recorded each instance in which a subject in the first group threw food toward one in the second group. In Phase 2 we provided a group of capuchins with food and noted each instance in which a subject threw food toward an empty cage. In Phase 3 we provided food simultaneously to two groups of capuchins and noted each instance of food-throwing between them. In Phase 1 subjects in one group threw food toward subjects in a second group, which, when provided the opportunity, did not throw food toward capuchins in the first group. Thrown food was either caught, retrieved, or lost on the test room floor. The rate of throwing decreased significantly when subjects were presented with an empty cage and when both groups of subjects were given food. We propose that psychological processes which underlie aimed throwing and food sharing came into existence through convergent evolution in large-brained, extractive foraging primates. We further speculate that although a well-developed system of exchange, based on contingent reciprocity, may occur among primates only inHomo,simpler transfer systems involving voluntary unidirectional passing of food from one individual to another appear to be more widespread among primates than previously thought and can be expressed in rather unusual circumstances such as those in this experiment.     

Social Transmission of Feeding Techniques in Tonkean Macaques?

We studied the influence of social partners on food processing and drinking techniques in a semifree-ranging group of Tonkean macaques. In a first study, immature subjects observed their mothers repeateadly eating unfamiliar fruits. They were subsequently allowed to consume the same fruits. We found no evidence for social learning when comparing the technical variants used by the mother and her offspring. In a second study, we recorded the techniques displayed by the monkeys while drinking and eating familiar food. We compared the technical variants used by the members of two different maternal lineages. Whereas a majority of techniques did not differ between lineages, some variants were peculiar to each of them, this effect was independent from age. The results indicate that the possibility of technique transmission should not be excluded among macaques.     

Response to novel food and the role of social influences in common marmosets (Callithrix jacchus) and Goeldi's monkeys (Callimico goeldii)

Neophobia, defined as showing caution toward novel features of the environment, is widespread in birds and mammals; it can be affected by ecology, early experience, and social context. In this study, we aimed to (i) investigate the response to novel food in adult common marmosets and Goeldi's monkeys and (ii) assess the role of social influences. We used an experimental paradigm employed previously with capuchin monkeys and children, in which a subject (observer) was presented with a novel food under three conditions: (i) Presence: group members did not have food; (ii) Different color: group members received familiar food whose color differed from that of the observer's novel food; (iii) Same color: group members received familiar food of the same color as the observer's novel food. Although most common marmosets tasted and/or ate the novel food, none of the Goeldi's monkeys ate it and only two sampled it. Differences in home range size and early social experience might explain the divergent behavior of the two species. Observers of both species similarly attended to group members and their visual attention increased with the number of group members eating, especially when the observer's and group members' foods were perceptually similar. However, we observed social influences on explorative behavior in Goeldi's monkeys but not on explorative or eating behavior in common marmosets. This result might be explained by the different pattern of response to novel food observed in the two species. Moreover, social influences on Goeldi's monkeys' behavior were nonspecific, i.e. they were not based on an appreciation that the food is safe because eaten by group members.     

The diet and feeding behavior of the silver leaf monkey (Trachypithecus auratus sondaicus) in Indonesia

I describe the diet and feeding behavior of silver leaf monkeys (Trachypithecus auratus sondaicus) in the Pangandaran Nature Reserve, West Java, Indonesia, and compare a group living in old secondary rain forest with a group living in mixed plantation/secondary forest to determine intraspecific variation in feeding behavior and the importance of the plantation species in the diet of the monkeys. Young leaves and leaf buds made up slightly less than half of their diets, with both groups showing a preference for a few species when seasonally available. Fruits and flowers of a few species were also preferentially selected when available. These included sweet, fleshy fruits, which most other colobines tend to avoid. Young leaf intake was greatest in months when fruit intake was low. Mature leaves were rarely eaten. Both groups spent approximately 20% of feeding time foraging on Moraceae species. Differences in the diet of the two groups were related largely to differences in vegetational composition and the availability and abundance of food items for the species common to both sites. Teak (Tectona grandis) was the top food species of the group living in mixed plantation/secondary forest, with the midribs of young leaves preferentially selected. Young leaves ofT. grandis, available throughout the study, provided a staple food and were eaten when preferred foods were scarce. More favored food items were available to the group living in old secondary forest, though none was a staple food.     

Food Transfer and Development of Feeding Behavior and Food-Associated Vocalizations in Cotton-Top Tamarins

Food transfer between adults and infants is common in many marmoset and tamarin monkeys, and is often accompanied by vocalizations. We hypothesized that vocalizations by adults in a food transfer context creates an opportunity for infants to learn not only what foods are appropriate but what vocalizations are appropriate in feeding contexts. We studied the development of feeding behavior and food-associated vocalizations in 10 infant cotton-top tamarins through the first 20 wks of life. Infants obtained solid food through transfers from older group members, primarily the adult male, beginning at weeks 5–6. Both adults and infants vocalized during food transfers with adults, producing rapid sequences of the call types adults normally give when feeding. Infants were usually successful in obtaining food primarily when the adult was vocalizing. The sooner infants were active participants in food transfers, the sooner they began to feed independently. In the early weeks, infants produced a large number of vocal types during food transfers, but with increasing age there was a steady increase in the number of adult-form food calls and a reduction in other, non-food-associated calls. Infants that fed independently at an early age produced fewer non-food-associated calls by the last month of observation. Infants called at higher rates to their most preferred food. Food transfers accompanied by vocalizations may provide an opportunity for infants to learn about appropriate foods as well as the vocalizations that accompany feeding in adults, and may represent a form of 'coaching' or information donation by adults.     

Utilization of fruiting trees by monkeys as analyzed from feeding traces under fruiting trees in the tropical rain forest of Cameroon

Individual trees of the food species of monkeys were identified by placing plastic tapes with an identification number on them in the tropical rain forest of Cameroon, West Africa. In order to determine the use of the feeding trees by monkeys, the ground under each of the trees was checked at least once a week to see if there were any fallen fruits or traces of feeding on fruits. Some fruit species were not fed on by either monkeys or large arboreal squirrels. Among the food species common to both the monkeys and large squirrels, a larger proportion in terms of quantity in each species was mainly eaten by the monkeys except in the case of super-abundantly fruit producing species. The monkeys and large arboreal squirrels were well segregated in their diets. Larger proportions (more than 85% for most of the monkeys' major foods) of fruits of larger sizes were made to fall on the ground by the monkeys and squirrels. The monkeys displayed a tendency to visit fruiting trees rather evenly (even rate of visit = even frequency of visit/duration of fruiting) not ignoring any area of the home range, although a small difference in this tendency was observed between the two study periods, one an abundant season and the other a poor fruiting season. On average, one associated polyspecific group of monkeys encountered only 14 fruiting trees per day. On the other hand, fruits were available all around the year, as the fruiting periods of different tree species were widely distributed around the year, or the fruiting periods of some species were very long. Although the monkeys are able to depend heavily on fruits, the quantity of fruits is not so great. The population size of monkeys is well balanced with the available food supply in the tropical rain forest of West Africa.     

Decisions on time allocation to different food patches by Japanese monkeys (Macaca fuscata)

From the standpoint of the possible development of an optimal foraging strategy, the time allocation to different food patches and the decision as to when to leave any given patch were examined in wild Japanese monkeys (Macaca fuscata). The investigations yielded the following results: (1) There was no tendency for feeding bouts to be of longer duration in higher-quality patches, i.e. the monkeys did not employ an energy-maximizing strategy during their feeding bouts. (2) The speed of feeding barely decreased with passage of time during a bout. In most cases, therefore, a decrease in feeding speed was not considered relevant to the decision to leave a patch. These findings are evaluated in terms of social factors, and it appears that following a monkey's own family group or the troop usually contributes to the timing of leaving a food patch.     

Feeding strategies of Japanese monkeys against deterioration of habitat quality

Field observations of the feeding behaviour of Japanese monkeys were carried out from autumn to winter on Kinkazan Island which is covered with cool temperate forest. As a result, the following two points became clear: (1) the available food items were fixed for a long time; and (2) the habitat quality deteriorated monotonously because the monkeys themselves or their competitors, such as wild mice, utilized the food resources. Against the decrease in food intake caused by this deterioration of the habitat quality, the monkeys controlled the decrease in food intake by employing the following strategies: (1) they recovered their feeding speed by exploiting new food patches (patch-increase strategy); (2) they extended the time spent on feeding (time-extension strategy); and (3) they changed their food (food-change strategy). The former two strategies operated earlier than the third one.     

Food-exchange with humans in brown capuchin monkeys

To assess how brown capuchin monkeys (Cebus apella) delay gratification and maximize payoff, we carried out four experiments in which six subjects could exchange food pieces with a human experimenter. The pieces differed either in quality or quantity. In qualitative exchanges, all subjects gave a piece of food to receive another of higher value. When the difference of value between the rewards to be returned and those expected was higher, subjects performed better. Only two subjects refrained from nibbling the piece of food before returning it. All subjects performed two or three qualitative exchanges in succession to obtain a given reward. In quantitative exchanges, three subjects returned a food item to obtain a bigger one, but two of them nibbled the item before returning it. Individual differences were marked. Subjects had some difficulties when the food to be returned was similar or equal in quality to that expected.     

Diet and Feeding Ecology of Woolly Monkeys in a Western Amazonian Rain Forest

I investigated the diet and feeding ecology of two social groups of woolly monkeys (Lagothrix lagotricha poeppigii) in Yasuní National Park, Ecuador between April 1995 and March 1996. Woolly monkeys in Yasuní were predominantly frugivorous, with fruits comprising ca. 77% of the yearly diet; the next most common food type in the diet was insect and other animal prey. The fruit diet of woolly monkeys in Yasuní is the most diverse yet recorded for any ateline primate, including spider monkeys (Ateles), which are often regarded as ripe fruit specialists: 208 distinct morphospecies of fruits were consumed by woolly monkeys either during the study or during several preceding months of pilot work. Nonetheless, close to one-third of the yearly diet came from just 3 plant genera—Inga, Ficus, and Spondias—and only 20 genera each contributed to 1% of the diet. For one study group, the proportion of ripe fruit in the diet each month was correlated with the habitat-wide availability of this resource, a pattern evidenced by several other ateline species. However, the relationship was not apparent in the second study group. The modal party size for feeding bouts on all food types was a single monkey, and, contrary to reports for other atelines, neither feeding party size nor the total number of feeding minutes that groups spent in food patches was well predicted by patch size. Both results highlight the independent nature of woolly monkey foraging. Given that woolly monkeys and closely-related spider monkeys focus so heavily on ripe fruits, their very different patterns of social organization are intriguing and raise the question of just how their ecological strategies differ. Two important differences appear to be in the use of animal prey and in the phytochemical composition of the ripe fruits that they consume: spider monkeys rarely forage for animal prey, and woolly monkeys seldom consume the lipid-rich fruits that are an important part of spider monkey diets.     

Effects of feeding cycles on circadian rhythms in squirrel monkeys

Squirrel monkeys (Saimiri sciureus) were housed singly in cages equipped with a tree for climbing to measure locomotor activity, and with a movable food cup that could be arrested automatically. The animals were kept in continuous dim illumination (LL), twice interrupted by several weeks of entrainment by a light-dark (LD) 12:12 cycle. Apart from three control sections in which the food cups were unlocked continuously (ad libitum feeding), food was accessible for 3 hr per day only, with interfeeding intervals varying from 23 to 26 hr (periodic restricted feeding, or RF). During LD entrainment, the imposition of an RF schedule resulted in anticipatory behaviors, represented by increased tugs at the food cup and a pause in locomotor activity preceding the feeding time. In LL, the animals showed free-running circadian rhythms of locomotor and "feeding" activity that nearly always persisted when ad libitum feeding was replaced by RF. The period (tau) of the free-running rhythm was slightly modulated in relation to the varying interfeeding intervals (T), but entrainment was never achieved except in one test with an animal whose tau was very close to T. It is concluded that periodic availability of food represents an extremely weak zeitgeber, if any, for the circadian pacemaker of squirrel monkeys.     

Feeding Competition in <Emphasis Type="Italic">Macaca fascicularis</Emphasis>: An Assessment of the Early Arrival Tactic

In primate species with unidirectional dominance relationships, rank order restricts the access of nondominant females to clumped resources. However, females might attempt to bypass the rank order by reaching feeding sites before the highest ranking individuals (early arrival tactic) when there are net benefits. We therefore analyzed the order of arrival to the feeding site of the adult members of a captive group of long-tailed macaques. We used 2 experimental conditions that differed in the spatial distribution of a fixed amount of food (large vs. small patch). Though each condition induced contest competition, it was stronger in the small-patch condition. Arrival order does not correlate with dominance rank in either experimental condition. The α-male and α-female reached the feeding site 10–30 s after the beginning of the test. Some females seized on opportunities to reach the feeding site before them, especially in the large-patch condition. They used the early arrival tactic when the risks of aggression were relatively low, which subjects accomplished either by being dominant or by being nondominant but tolerated by the α-male. Social tolerance may provide individuals with an alternative means to obtain resources. In sum, variation in food abundance and distribution may affect the extent to which rank order determines order of arrival to feeding sites. A higher rank may confer priority in the choice of tactics, but not necessarily priority of access to the resources themselves.     

Grooming,kinship, and co‐feeding in captive spider monkeys (Ateles geoffroyi)

Allo‐grooming is perhaps the most powerful affiliative behavior observed in non‐human primates. However, the functional significance of grooming in New World monkeys has not yet been fully understood, perhaps because grooming is less frequently observed in platyrrhines. To differentiate the potential role of affiliative investment and/or kinship on sharing access to food (co‐feeding) in spider monkeys, behavioral data on grooming, embracing, and feeding were collected from two different groups of captive study subjects: a familiar/kin group and a non‐familiar/non‐kin group. The results of this study suggest that family‐related spider monkeys that engage in grooming tend to share access to food resources more than unfamiliar conspecifics that do not groom. One explanation for this difference is that the unfamiliar study subjects had not yet invested in the affiliative social network, were not reciprocating their affiliative investments and hence, had a higher tendency toward single animal monopolizing resources. Degree of relatedness alone was not found to be a determinant for sharing the access to food, suggesting that familiarity in spider monkeys is based on the extent to which animals invest in affiliative relationships. In this study, only animals that had engaged in long‐term grooming and recognized each other as familiar shared the access to food. Therefore, it might be likely that in spider monkeys, long‐term grooming of high intensity has to be developed for co‐feeding to occur. Zoo Biol 20:293–303, 2001. © 2001 Wiley‐Liss, Inc.     

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

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

Factors affecting feeding decisions in a group of black lemurs confronted with novel food

The behavior of black lemurs presented with novel and familiar food in the presence or absence of the dominant female was analyzed in order to examine the influence of the latter on feeding decisions at a group level. Results showed that individual learning occurred with high quality food, while the dominant female influenced the eating behavior of other group members when food was of low attractiveness or unfamiliar.     

Gastrointestinal responses to feeding in a frequently feeding colubrid snake (Natrix maura)

Ectotherm vertebrates show physiological mechanisms that reduce metabolic costs during prolonged fasting. Once feeding, these animals adopt a wide variety of metabolic responses such as changes in gastrointestinal organ masses. Up-regulatory responses after feeding have been widely explored in infrequently feeding snakes like pythons, whereas few studies have been devoted to frequently feeding snakes. In this study, we have considered the gastrointestinal responses after feeding in a frequent feeder, the viperine snake Natrix maura, in the Ebro Delta rice fields. In this habitat, viperine snakes are exposed to long periods of food deprivation due to the lack of available prey as a consequence of the man-induced rice cycle. We weighed prey items and full gut masses, and measured length of combined esophagus and stomach, and intestine of viperine snakes belonging to a wide range of sizes. Snakes concentrate foraging activity when rice fields were flooded. In this period, gut masses increased. Likewise, intestines increased in length during the feeding period, which suggests that viperine snakes probably experience a postfeeding hypertrophy of their small intestines that contributes to their larger length. Once the intestine length was corrected for the snake size, it was shown that adults present longer intestines than immature snakes, reflecting an increase in the posterior part of the body linked to the gonads development. This study contributes to explore the physiological responses to feeding in frequently feeding snakes modelled by abrupt shifts of food availability.     

Diet and activity budget of <Emphasis Type="Italic">Rhinopithecus roxellana</Emphasis> in the Qinling Mountains,China

We collected data on diet and daytime activity budget, and investigated the phenology of food trees and food abundance for a group of Rhinopithecus roxellana on the East Ridge of Yuhuangmiao in the Qinling Mountains from November 2001 to December 2003. We calculated the seasonal activity budget using data collected by scan sampling from 84 full-day observations (winter 16, spring 18, summer 28, autumn 22 days). During scan sampling we recorded behavioral states, and the food items and species consumed. The subjects consumed 84 plant species, including trees and shrubs of 29 families, and lichens. Food species varied seasonally. The overall diet of R. roxellana consisted of 29.4% fruit/seeds, 29.0% lichens, 24.0% leaves, 11.1% bark, 4.2% buds, 1.3% twigs and 1.0% unidentified items. Because the abundance of different food items varied seasonally, the monkeys had to shift their major food items seasonally. The annual activity budget of R. roxellana was 36.2% time spent resting, 35.8% feeding, 22.9% moving, and 5.1% other behavior. Seasonal changes in activity budget were observed. R. roxellana spent more time moving in autumn, when the quality of the food might be highest, and least time moving in winter when the food quality might be lowest. Thus, this type of monkey has a passive foraging strategy.     

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

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

Time budget of the green monkey,Cercopithecus sabaeus: Some optimal strategies

Members of a single group of green monkeys spent, on average, 44.8% of their waking time foraging, 46.7% resting, and 8.5% in social activities, over 1 year. There was significant variation in activity budgets over months (ranging from 35 to 55% of the time spent feeding). Diurnal rhythms of feeding and ranging were influenced by the daily cycle of temperature in predictable ways in different seasons: in the dry season, activity was reduced if it was too hot or too cold, while the temperature in the wet season did not affect activities. Feeding was also synchronized among individuals on a finer time scale, irrespective of the time of day. There was closer synchrony when feeding on less common foods. An ecological model of foraging time and energetics was tested, using estimates of the costs and benefits of foraging and predicting how these are optimally balanced in relation to the food density. Both feeding time and distance traveled increased as food availability increased. Costs and benefits were balanced over several days. Comparisons between populations of Cercopithecus aethiopswere made; differences in time budgets were compared with differences in the availability and quality of food. Insufficient comparative data are available for firm conclusions about the role of different energetic and nutritive strategies in population differences.