Sex and group differences in feeding on animals by wild chimpanzees in the Mahale Mountains National Park,Tanzania

Sex differences in animal prey intake were revealed by fecal analysis among wild chimpanzees of the large-sized M-group (ca. 100 members) in the Mahale Mountains National Park, Tanzania: prime adult or old males feed more on vertebrates, while adult females more onCamponotus ants. By contrast, such differences were not obvious in the neighboring, small-sized K-group (ca. 20–30 members), despite the similar environment in which the two unit-groups lived. Such sex and group differences may be explained in terms of various factors, either ecological or social, or both, but social factors seem most responsible in particular for the group differences. It seems likely that increased capture rate of vertebrates per unit-group in the larger-sized M-group results in increased per capita intake of meat among prime adult or old males. Also, the more frequent interactions among prime adult or old males of M-group appear to reduce the frequency of theirCamponotus ant-fishing behavior.     

Social Relationships Among Male <Emphasis Type="Italic">Alouatta pigra</Emphasis>

We investigated male social relationships in 2 groups of black howlers (Alouatta pigra) in Palenque National Park, Mexico, over a 14-mo study characterized by frequent changes in male group membership. Both single males and pairs of males entered our focal groups. Single males tended to join groups, whereas pairs of males entering groups together successfully evicted resident males. The 19 male dyads across the 11 periods defined by changes in group membership were rarely in close proximity or interacting with one another. Nevertheless, 1 male formed significantly closer associations with resident females in 6 periods and achieved higher mating success during 4 of the periods when ≥1 female was sexually active. In the other 5 periods, no particular male maintained significantly closer associations with resident females, which may be a result of the rapid sequence of changes in male membership and corresponding social instability. Resident males participated frequently and consistently in mutual howling, but the resident male with the strongest female associations and highest mating success initiated howling bouts more frequently in all but 1 of the 6 socially stable periods. Though still preliminary, our findings suggest that variation in social relationships among male black howlers may be more related to their ability to establish strong relationships with females than to their relationships with one another.     

Rates of predation on mammals by gombe chimpanzees, 1972–1975

Rates of chimpanzee predation on mammals are calculated using data on 75 kills recorded during focal observation in Gombe National Park, Tanzania, from January 1972 to April 1975. The chimpanzees were members of two study communities (Kanyawara, or Northern, and Kahama, or Southern, community), and were observed as focal individuals for 14,583 hr by more than 30 researchers and field assistants working in pairs. The rate of predation by females was too low to allow reasonable estimates. For males, the mean rate of killing during the study period was 0.31 kills per male per 100 hr (N=17 males), or 4.65 kills per 100 hr in the two communities. In contrast to results from Mahale Mountains, there was no difference in predation rate between wet and dry seasons. However, predation rates varied over time, increasing by four times between the first three and last four seasons of the sample period. In an average year the 15 adult and subadult male chimpanzees are calculated to have killed 204 prey per year in an area of 16 km², varying between 99 and 420 prey per year in periods of low and high predation rate. Red colobus were the most frequent prey, followed by bushpig and bushbuck. Predation rates varied greatly on different prey species, and were not related to either the proportion of time spent within 200 m of male chimpanzees, or to their population densities. In relation to encounter rates and population density, baboons, blue monkeys, and redtail monkeys were killed at a fraction of the rate of red colobus monkeys, which suffered severe mortality from chimpanzee predation. Predation on bushpig and bushbuck also appears to have been high in relation to population density. The amount of food provided by predation is estimated to have averaged 600 kg per year for chimpanzees in the two communities (totalling 14–17 adult or subadult males, 18–20 adult of subadult females, and about 19 infants or juveniles). This suggests that adult males consumed around 25 kg of meat per year, although any average figure undoubtedly masks considerable individual variation. Present data suggest that chimpanzees in Gombe and Tai National Park, Ivory Coast, prey on mammals at rates higher than other populations.     

Sex differences in diet and foraging behavior in white-faced capuchins (Cebus capucinus)

I examined sex differences in diet and foraging behavior in two groups of white-faced capuchin monkeys (Cebus capucinus)in a tropical dry forest at Santa Rosa National Park, Costa Rica. I tested three hypotheses: sex differences in diet and foraging behavior are best explained by (1) sexual dimorphism, (2) the energy demands of pregnancy and lactation for females;and (3) avoidance of competition between the sexes. Sexual dimorphism offered the best explanation of sex differences in the diet and foraging behavior of C. capucinus,accurately predicting that males do more strenuous foraging activity, make less use of small foraging supports, and spend more time on or near the ground. Females spent more time foraging than males did but probably obtained a lower protein yield per unit foraging time. Females exploited more small and embedded invertebrates, while males ate more large invertebrates and vertebrate prey. Pregnant and lactating females spent more time resting and less time foraging than other females did, increasing their foraging return by focusing on foods requiring little handling. There was little evidence of competition avoidance between the sexes.     

Male-female interactions of sika deer (cervus nippon) in nara park through allogrooming during breeding and rutting seasons

The frequency and pattern of interactions between males and females of sika deerCervus nippon were surveyed in 3 sites of Nara Park, central Japan, mainly via observation of allogrooming frequency and aggressive behavior throughout the breeding and rutting seasons. In the breeding season, the Daibutsuden group contained several adult males while Ukimido and Hakuba groups contained only a few. Most allogroomers were adult and subadult females. Females groomed the same sex more than the opposite sex at Daibutsuden, but at the other 2 sites, there was no such difference. In general, male-female interactions through allogrooming frequencies in the breeding season were not significantly fewer than intra-sex interactions. Males that attacked females tended to be groomed by females in the breeding season. In the rutting season, all adult males identified in the breeding season were absent at the usual observation sites, and newly arrived males showed defensive and/or mating behaviors. Females groomed adult males in the rutting season regardless of group status or display of sexual interactions. This suggests that male-female allogrooming in the breeding season does not relate to mating in the following rutting season, but may reduce tension in the group during the current breeding season.     

Dispersal Patterns Among Olive Colobus in Taï National Park

In Primates, females are more likely to be philopatric than males. However, in some species like Procolobus verus, females or individuals of both sexes disperse. In Taï National Park, Ivory Coast, olive colobus groups are small, with one or two adult males and 6 females. Dispersal is common for juveniles and adults of both sexes. Adult male dispersal is less common than adult female dispersal. Adult females immigrated especially into small, one-male groups indicating that food competition played a role. Furthermore, unknown sexually receptive females visited resident groups and mated with the resident males for a few days before disappearing again. Adult males dispersed when this improved their mating opportunities. All juveniles left their natal groups. The dispersal of juveniles may be a strategy to prevent inbreeding with their parents. Dispersal by juvenile males furthermore seemed to be the result of mate competition. The high dispersal rates, visits by receptive females, and dispersal of all individuals in the population suggest that moving between groups is a strategy that can be used ad hoc in several situations more easily in the olive colobus than in most other primates. The predation risks related to moving between groups were reduced by dispersing in conspecific or allospecific groups and by dispersing to neighboring groups.     

General Gregariousness and Specific Social Preferences among Wild Chimpanzees

Wild chimpanzees form temporary parties that vary in size and composition. Previous studies have revealed considerable intraspecific variation in party compositions. We examined patterns of association among age, sex, and reproductive classes of chimpanzees at Ngogo in the Kibale National Park, Uganda. We employed a class-based association index and a randomization procedure to control for confounding factors and to test for differences between classes. Results indicate that males associated with other males significantly more than expected if all classes behaved equivalently, while females generally associated with individuals of the same sex less than expected. To interpret these patterns we used two additional indices that separate associations into two components: general gregariousness and preference for particular classes of associates. Males and estrous females were more gregarious than other classes, while anestrous females were less so. After controlling for general gregariousness, adult males as a class showed no specific preference for associating with each other. Anestrous females preferred each other as party members, and estrous females avoided each other. These results are consistent with previous findings that adult males are more gregarious than females. They diverge from the standard picture of chimpanzee society, however, by suggesting a mutual affinity among anestrous females, but not among adult males as a class.     

Relationship tenure differentially influences pair-bond behavior in male and female socially monogamous titi monkeys (Callicebus cupreus)

Pair-bonded primates have uniquely enduring relationships and partners engage in a suite of behaviors to maintain these close bonds. In titi monkeys, pair bond formation has been extensively studied, but changes across relationship tenure remain unstudied. We evaluated differences in behavioral indicators of pair bonding in newly formed (~6 months paired, n = 9) compared to well-established pairs (average 3 years paired, n = 8) of titi monkeys (Callicebus cupreus) as well as sex differences within the pairs. We hypothesized that overall males would contribute more to maintenance than females, but that the pattern of maintenance behaviors would differ between newly formed and well-established pairs. Each titi monkey (N = 34) participated in a partner preference test (PPT), where the subject was placed in a middle test cage with grated windows separating the subject from the partner on one side and an opposite-sex stranger on the other side. During this 150-min behavioral test, we quantified four key behaviors: time in proximity to the partner or stranger as well as aggressive displays toward the partner or stranger. Overall, we found different behavioral profiles representing newly formed and well-established pair-bond relationships in titi monkeys and male-biased relationship maintenance. Males spent ∼40% of their time in the PPT maintaining proximity to the female partner, regardless of relationship tenure. Males from well-established bonds spent less time (14%) near the female stranger compared to males from newly formed bonds (21%) at the trend level. In contrast, females from well-established bonds spent less (23%) time near the male partner in the PPT compared to females from newly formed bonds (47%). Aggressive displays were more frequent in newly formed bonds compared to well-established bonds, especially for females. Scan sampling for homecage affiliation showed that newly formed pairs were more likely to be found tail twining than well-established pairs.     

Aggressive behavior and dominance hierarchies in female sable antelope,Hippotragus niger: Implications for captive management

Social interactions of adult female sable antelope, Hippotragus niger, were observed at the National Zoological Park's Conservation and Research Center. Aggressive behavior was quantified for two herds containing 8 and 11 females. Aggression was lowest at calving, and highest near the time of conception. Dominance hierarchies were strictly linear and stable over time. Dominance rank was highly correlated with age, with the oldest females achieving highest rank. High-ranking females initiated more aggression and were more likely to behave aggressively towards a newly introduced adult female. The newly introduced individual was the victim of prolonged, intense group chases and was permanently excluded from the herd. Low-ranking females directed more aggression at yearling herd members than did high-ranking females. Temporary absences by herd members resulted in long-lasting, substantial increases in received aggression upon their return. Because of their aggressive nature, sable antelope present a challenge to captive management. © 1993 Wiley-Liss, Inc.     

Nutritional quality of gorilla diets: consequences of age,sex, and season

We tested the effects of age, sex, and season on the nutritional strategies of a group of mountain gorillas (Gorilla beringei) in the Bwindi Impenetrable National Park, Uganda. Through observations of food intake of individual gorillas and nutritional analyses of dietary components over different seasons and environments, we estimated nutrient intake and evaluated diet adequacy. Our results suggest that the nutritional costs of reproduction and growth affect nutrient intake; growing juveniles and adult females ate more food and more protein per kilogram of metabolic body mass than did silverbacks. The diets of silverback males, adult females, and juveniles contained similar concentrations of protein, fiber, and sugar, indicating that adult females and juveniles did not select higher protein foods than silverbacks but rather consumed more dry matter to ingest more protein. Juveniles consumed more minerals (Ca, P, Mg, K, Fe, Zn, Mn, Mo) per kilogram of body mass than adult females and silverback males, and juveniles consumed diets with higher concentrations of phosphorous, iron, and zinc, indicating that the foods they ate contained higher concentrations of these minerals. Seasonally, the amount of food consumed on a dry weight basis did not vary, but with increased frugivory, dietary concentrations of protein and fiber decreased and those of water-soluble carbohydrates increased. Energy intake did not change over the year. With the exception of sodium, gorillas ate diets that exceeded human nutrient requirements. A better understanding of the relative importance of food quantity and quality for different age–sex classes provides insights into the ways in which gorillas may be limited by food resources when faced with environmental heterogeneity.     

Inter-male associations in a wild Japanese macaque troop on Yakushima Island,Japan

Adult male association and its annual change were studied in a wild population of Japanese macaques (Macaca fuscata yakui) on Yakushima Island, Japan. Unlike many other Japanese macaque troops, adult troop males frequently maintained proximity and exchanged grooming with one another in both the mating and non-mating seasons, and the dominance relationship rarely appeared in such inter-male associations. The few cases of agonistic interactions occurred mostly when estrous females or food resources were immediately concerned. Although troop males were very intolerant to newly appeared solitary males (new males) during the mating season, close associations were formed between troop males and new males as soon as the mating season terminated. The consort of new males and lower-ranking troop males with estrous females was frequently disturbed, but these males could copulate no less frequently than higher-ranking males. A comparison among macaque species suggests the existence of two forms of inter-male association: (1) the frequent association based on the symmetrical exchange of social behaviors; and (2) the infrequent and asymmetrical association related to the dominance relationship. The form of inter-male association seems to be influenced by whether or not males can keep close associations with females throughout the year.     

Food calling by captive bonobos (Pan paniscus): An experiment

We examined (i) whether bonobos display a specific food-calling behavior when discovering a hidden food resource, (ii) whether the presence of competitors affects this behavior, and (iii) whether food quantity or gender influences its appearance. We carried out experiments (n = 108) within a captive group of eight bonobos at the Animal Park Planckendael (Mechelen,Belgium). We hid highly preferred food items (n = 7 or 25) in their enclosure and recorded vocal behavior and interactions between discoverer and group members. As a control, we gave the same number of items to the individuals when isolated from the group, a situation without potential food competition (n = 38). The only vocalization frequently uttered by the discoverer was the food peep. They uttered food peeps significantly more often when no food competition was possible. The amount of food had no significant influence on whether food peeps were uttered. The same applies to the individuals’ identity or gender. Although the costs of food calling behavior seemed much higher for males, both sexes uttered food calls to the same extent. We hypothesize thai males signal food presence in order to attract potential mates and are willing to give up the discovered food resource in return for sex: sex for food exchange. In contrast, females may vocalize to attract coalition partners. Through these coalitions, they can monopolize food resources vis-à-vis males. It is also possible that females have less reason to suppress food calk, since they are dominant to males. This study suggests that bonobos are able to give shaded signals about their environment and have the potential to communicate this information in order to promote their sexual strategy.     

Social scratch: Another custom in wild chimpanzees?

Chimpanzees (Pan troglodytes schweinfurthii) in the Mahale Mountains National Park, Tanzania, scratch other individual's bodies while they groom them. This behavioral pattern of “social scratch” is another example of locality-specific social behavior, or custom, as it is not found in the Gombe National Park, Tanzania, about 150 km north of Mahale, nor has it been reported from any other sites of chimpanzee study. Frequency of social scratch was correlated with frequency of social grooming, but not with frequency of self-scratch. Frequencies of social scratch per grooming bout among adult and adoles-cent males, and from lactating females to infants or juveniles, were high, and among males, higher-ranking males especially received more. These facts indicate some social function of the behavior. Social scratch was directed mostly to the dorsal side of the body. However, when lactating females social scratched to infants or juveniles, they scratched other body parts. Social scratch was not lateralized to left or right. We present four hypotheses on the functional origin and on the learning process of this cultural behavioral pattern.     

Sexual behavior across ovarian cycles in wild black howler monkeys (Alouatta pigra): male mate guarding and female mate choice

We studied two multimale–multifemale groups of black howler monkeys (Alouatta pigra) during a 14‐month study (June 2006–July 2007) in Palenque National Park, Mexico to evaluate the ways in which their sexual behavior changes across ovarian cycles. We analyzed 231 fecal samples, collected every 2.2±1.4 days from five females. For four females, estradiol and progesterone profiles revealed an average (±SE) cycle length of 18.3±1.4 days. Copulations occurred significantly more frequently during the periovulatory period (POP), defined as the estimated day of ovulation ±3 days (N=18). This was largely the result of cycling females soliciting sexual interactions during their POPs. Females directed their solicitations significantly more often toward “central” males of their group, who had close spatial associations with females at other times, compared with “noncentral” males, who did not associate closely with females. Central males rarely solicited sexual interactions, but instead monitored the females' reproductive status by sniffing their genitals, and maintained significantly closer proximity to females during their POPs, suggesting male mate guarding when conceptions are most likely to occur. Our findings indicate that the reproductive strategies of black howler central males and females coincide, highly skewing mating opportunities toward central males. Black howler females, however, occasionally choose to copulate with noncentral resident males or extra‐group males during their POPs, undermining the ability of central males to monopolize all reproductive opportunities. Am. J. Primatol. 71:153–164, 2009. © 2008 Wiley‐Liss, Inc.     

Social interactions regulate resource utilization in a Tephritidae fruit fly

Previous studies have suggested that social interactions (e.g., the actions and reactions elicited by the interaction of co-specific individuals) induce individual fruit flies (Tephritidae) to ingest more food, especially protein-rich food. Changes in feeding behavior related to social interactions have been associated with reproduction (e.g., when different sexes are present), reproductive facilitation (e.g., when two females interact) and stress and aggression (e.g., flies of the same sex, or crowdedness). The present study investigated the effect of social interaction on the feeding, longevity and resource management of the Ethiopian fruit fly, Dacus ciliatus. Single flies and pairs of flies (of the same or different sexes) were confined to a small arena (the PUB system), in which we measured the amount of liquid food ingested daily by each fly. In addition, we sampled flies of different ages, extracted and quantified their lipid and protein contents, and related individual metabolic contents to the ingestion of a fructose and protein hydrolysate solution. Results showed that individual ingestion was significantly higher in flies maintained in pairs than in flies kept as solitary individuals. The highest intake rates were observed for the female–female pairs. In general, females ingested significantly greater volumes than males. Lipid contents tended to decrease progressively with age in flies kept as solitary individuals, especially in female flies, while lipid levels decreased and then increased in flies maintained in pairs. Protein trends were similar, although less pronounced than the patterns observed for the lipids. The flies kept as solitary individuals lived significantly longer than those kept in pairs. A resource-management analysis points to a decreased metabolic rate in flies kept as solitary individuals, as compared to paired flies. Results are discussed in view of theories of resource management and survival strategies.     

Feeding ecology of the long-tailed macaque(Macaca fascicularis) in Kalimantan Tengah,Indonesia

I studied long- tailed macaque (Macaca fascicularis)feeding behavior and ecology as part of a larger behavioral ecological study at the Natai Lengkuas Station, Tanjung Puting National Park, Kalimantan Tengah, Indonesia. I collected data on feeding behavior via scan sampling of all visible individuals in the focal group (approximately 800 observation hours). I established vegetational plots and monitored them monthly to determine food availability and abundance. I found long- tailed macaques to be primarily frugivorous;leaves, flowers, insects and bark provided the remainder of the diet. They used at least 33 plant species as food sources, but >60% of the diet was provided by only 5 species. Based on previous vegetational analyses, these tree species were among the highest in relative density. However, selection ratios for 19 food species indicate that 13 of them were selected more often than expected. Long- tailed macaques appear to be selective feeders but can exploit a variety of food sources during periods of food scarcity.     

The behavior of postreproductive females in a wild population of toque macaques (Macaca sinica) in Sri Lanka

I compared the behavior of three old postreproductive females in a wild population of toque macaques (Macaca sinica)in Polonnaruwa with those of reproductive females via focal-animal sampling techniques. Postreproductives foraged less, slept more, and were less active overall than reproductive females were. They also had significantly lower rates of agonistic behavior, were more peripheral, and had lower frequencies of overall affiliative contact. Although postreproductives initiated contact with others as frequently as reproductives did, group members initiated contact with them significantly less than they did with reproductive females. Postreproductives associated more with adult females than reproductives did and less with adult and subadult males than high-ranking reproductives did. Juvenile and infant females associated more frequently with reproductive females of high or low rank than with postreproductives. Postreproductives resembled low-ranking reproductive females in giving less grooming to others than they received. This contrasts with high-ranking females, which gave more grooming to others than they received. The results suggest that old age and cessation of reproduction are evident through the manifestation of distinct behavioral characteristics in toque macaque females.     

Activity and ranging patterns of Colombian woolly monkeys in north-western Amazonia

In this study, I revise three aspects of the socioecology of woolly monkeys (genus Lagothrix) that might give us a better understanding of the patterns found in this species: (1) the association between temporal variation in fruit abundance and diet, activity, and ranging patterns; (2) the individual trade-offs associated with living in small or large groups, and (3) the relationship between social dominance and foraging success. Using behavioral and ecological data collected during 3 years in Tinigua Park, Colombia, I found that woolly monkeys tend to avoid open-degraded forests, where fruit production is generally lower than it is in mature forests. Diet and activity budgets were highly associated with temporal patterns of fruit production. Daily path length was positively correlated with group size and monthly fruit abundance, and negatively correlated with habitat quality. I found differences in activity budgets and the diet preferences of different age/sex classes. For example, adult males rest more and juveniles play more than other classes. Juveniles and adult females without infants look for arthropods more often than adult males and females with young infants, who showed the highest frequencies of fruit feeding. Dominant adult males were not consistently the most efficient foragers on fruits according to two different indexes. Most of these results are consistent with the expectations from strong intra-group competition for resources. However, females with infants received benefits during feeding similar to those of dominant adult males, which may be mediated by differential aggression from males to other group members (juveniles and females without infants).     

Female dominance rank and behavior during artificial menstrual cycles in social groups of rhesus monkeys (Macaca mulatta)

We recently reported (Michael & Zumpe: American Journal of Primatology 15:157–170, 1988) evidence for the hypothesis that copulations by pairs of rhesus monkeys are linked more closely to the ovulatory phase of the female's menstrual cycle when a male has access to several females in different cycle phases rather than access to females in the same cycle phase. We have now used data from this earlier study to assess the role of female dominance. Artificial menstrual cycles were induced with hormone injections in two of four ovariectomized females in eight social groups (each consisting of one male and four females). The artificial cycles were either synchronized or offset by 7-day increments. There were differences in the behavioral interactions of the higher-ranking (dominant) and lower-ranking (subordinate) hormone-treated females in each group. The amount of male sexual activity received by dominant and subordinate females differed little, but dominant females initiated significantly more sexual activity (P<.0001) than did subordinate females. Dominant females were significantly more aggressive (P<.0001) toward other females and received more direct and redirected aggression from males. Males generally initiated and maintained more social contacts with dominant than with subordinate females, and dominant females were less active than subordinates in initiating and maintaining social interactions with males. The findings provide new evidence for both female mate competition and mate choice by males for higher-ranking females.     

Influence of Ecological and Social Factors on Body Mass of Wild Chimpanzees

The chimpanzees (Pan troglodytes schweinfurthii) of Gombe National Park, Tanzania, were weighed regularly over a period of 33 yr, resulting in 1286 measurements on 31 males and 26 females aged 2–43 yr. Female growth slowed at 10 yr and that of males at 13 yr. Median adult body mass is 39 kg for males and 31.3 kg for females. Body mass varied between years. Chimpanzees were heaviest during a period of frequent banana provisioning. They were also heavier when community range size was large and population density within the range was low. Chimpanzees were heavier in the wet than in the dry season and body mass tracked rainfall in the preceding mo except for May in which mass was anomalously low. Dominance rank is significantly correlated with body mass for females but not males. High-ranking individuals tended to maintain more stable mass. Variability in body mass was greater for young and old individuals than for prime adults.