The growth pattern of chimpanzees: Somatic growth and reproductive maturation inPan troglodytes

Growth of chimpanzees reared at the Kumamoto Primates Park of Sanwa Kagaku Kenkyusho Co. Ltd. was studied cross-sectionally from the viewpoints of somatic growth and reproductive maturation. Distance and velocity curves were expressed using spline function method. Males showed adolescent growth acceleration in body weight, with a peak at 7.86 yrs of age, but not in trunk length. Females showed continuous rapid growth from mid-juvenile to adolescent phase in both body weight and trunk length, but no isolated adolescent spurt. The Sanwa chimpanzees matured at about 12.5 yrs of age for females and 15.0 yrs for males. The mean adult weights and trunk lengths were 53.2 kg and 507.8 mm for males and 42.7 kg and 481.6 mm for females. The Sanwa chimpanzees had similar growth patterns to those of the Yerkes chimpanzees, although they showed a slight delay in infancy, and a higher growth rate from the early juvenile phase onwards. Growth patterns in these two laboratories may be regarded as “normative” for laboratory-reared chimpanzees. They matured earlier than wild chimpanzees by more than two years. The major reason for the retarded maturation in wild chimpanzees is the delay of growth from infant to the early juvenile phases (0–4 yrs of age), probably owing to a limited nutritional supply from the mother. Development of the testes comprised three phases: slow growth from infant to juvenile (until 6.4 yrs); rapid growth around adolescence (until 9.2 yrs); and adult (mean testicular volume, 187 cm³). Setting the nutritional standard at 2,000–2,600 Cal/day (= Kcal/day) per adult, calories were considered for captive chimpanzees in each age class.     

Do bonobos copulate more frequently and promiscuously than chimpanzees?

The copulatory activities of bonobos (Pan paniscus) of Wamba, Zaire, were compared with those of chimpanzees (P. troglodytes schweinfurthii) of Mahale, Tanzania. The copulation rates of adult male bonobos were equal to or lower than those of adult male chimpanzees. The copulation rates of adult female bonobos were approximately equal to those of adult female chimpanzees who were in maximal genital swelling, but it should be much higher than those of the adult female chimpanzees throughout the birth interval. The copulation rates of adolescent male bonobos were lower than those of adolescent male chimpanzees, whereas the copulation rates of adolescent female bonobos were much higher than those of adolescent female chimpanzees. It was suggested that the bonobos of Wamba did not copulate more promiscuously than did the chimpanzees of Mahale. The female bonobos may show “receptivity”, whereas female chimpanzees may show rather “proceptivity”.     

Age differences in social interactions of young males in a chimpanzee unit-group at the Mahale Mountains National Park,Tanzania

The behavior and social interactions of young male chimpanzees were studied in relation to their age change. The data were obtained at the Mahale Mountains National Park, during a four-month period in 1986. Early adolescent males, becoming independent of their mothers, spend a long time near adults of both sexes. Late adolescent males are not tolerated by the senior males. Although such animals do not stop traveling together with their seniors, they are separated from the other members including the males of their own age class, and each of them lives a relatively lonely life. Where seniors are not nearby, they perform charging displays in front of estrous females. Young adult males tend to remain in the proximity of the alpha male, and can associate with their seniors without pant-grunting. Although some young adult males dominate over some senior males, increasingly performing charging displays, they do not appear to be permitted to associate intimately with their seniors; they are not yet considered to have attained social maturity. Prime and senior males are strongly bonded with one another, being able to associate intimately with those of their own or senior age classes including the alpha male. A young adult male's rise in rank is not connected with joining the “adult male-cluster,” nor does a senior male's decline necessarily means his dropping out from the cluster: the social position of male chimpanzees cannot be understood solely from their agonistic dominance rank. The alpha male plays a leading part in integrating the males of the unit-group. Young adult males and their seniors tend to associate most frequently with him, and all the males of the early adolescent or senior age classes pay attention to his movements.     

Tool-using and -making behavior in wild chimpanzees at Bossou,Guinea

The behavior of wild chimpanzees at Bossou, Guinea, was studied from November 1976 to May 1977 recognizing each chimpanzee without artificial feeding. During the study period some tool-using and tool-making behavior was observed, as follows: (1) Although water drinking using a “leaf-sponge” was not seen, that using a “leaf-spoon” was observed for taking water from the hollow of a tree. (2) “Termite fishing” was not seen in this group although there were many termite hills in the moving range of the chimpanzees. They dug termites from the hollow of a tree by pounding with a small stick. Similar use of a stick was made for digging up the resin from a tree. (3) “Aimed throwing” was frequently observed in adult males for attacking an observer, and in adolescents and juveniles as mischief against an observer or for their own play. (4) “Nut cracking” with a pair of stones was seen for removing the ovule from palm-seeds. Particular stones were repeatedly used by many chimpanzees for a long period. (5) “Branch hauling” represented difficult work. Patient and inventive manufacture of proper sticks was necessary for capturing branches which they were unable to reach normally. Local variations in the tool-using patterns and manufacturing ability of chimpanzees are discussed.     

Predation on mammals by chimpanzees in the montane forest of Kahuzi, Zaire

The rate of predation on mammals by chimpanzees was determined from carcasses and from fecal specimens found on fresh trails during a 16-month period in the montane forest of Kahuzi-Biega National Park, Zaire. A unit-group of semi-habituated chimpanzees, composed of 22 – 23 individuals including 8 adult or adolescent males, appeared to kill about 18 – 30 mammalian prey (16 – 28Cercopithecus monkeys) per year, if the multiple kills by chimpanzees were not considered. A juvenile l'Hoest's monkey was recorded for the first time as the prey of chimpanzees in this study. Predation occurred in the late dry and the early rainy seasons, when the diversity of ripe fruits was the highest during the year. The Kahuzi chimpanzees tended to kill mammals less frequently but to killCercopithecus monkeys more frequently than chimpanzees in other habitats. The absence of red colobus monkeys, which are the most frequent prey in Gombe, Mahale, and Tai, might be responsible for the low predation rate. However, the estimated rate of predation onCercopithecus monkeys is the highest record among various chimpanzee habitats. At least 11 – 18% of theCercopithecus population seemed to be lost annually as a result of being killed by chimpanzees. Chimpanzees may be the most important predators on these monkeys in the absence of leopards at Kahuzi. The examination of fecal samples and carcasses suggested that adult (probably male) or adolescent chimpanzees tended to eat juvenile or subadult monkeys most frequently, as is also seen for chimpanzees in Gombe, Mahale, and Tai.     

Comparing infant and juvenile behavior in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): a preliminary study

The dichotomy between the two Pan species, the bonobo (Pan paniscus) and chimpanzee (Pan troglodytes) has been strongly emphasized until very recently. Given that most studies were primarily based on adult individuals, we shifted the “continuity versus discontinuity” discussion to the infant and juvenile stage. Our aim was to test quantitatively, some conflicting statements made in literature considering species differences between immature bonobos and chimpanzees. On one hand it is suggested that infant bonobos show retardation in motor and social development when compared with chimpanzees. Additionally it is expected that the weaning process is more traumatic to chimpanzee than bonobo infants. But on the other hand the development of behaviors is expected to be very similar in both species. We observed eight mother–infant pairs of each species in several European zoos. Our preliminary research partially confirms that immature chimpanzees seem spatially more independent, spending more time at a larger distance from their mother than immature bonobos. However, the other data do not seem to support the hypothesis that bonobo infants show retardation of motor or social development. The development of solitary play, environmental exploration, social play, non-copulatory mounts and aggressive interactions do not differ between the species. Bonobo infants in general even groom other group members more than chimpanzee infants. We also found that older bonobo infants have more nipple contact than same aged chimpanzees and that the weaning process seems to end later for bonobos than for immature chimpanzee. Additionally, although immature bonobos show in general more signs of distress, our data suggest that the weaning period itself is more traumatic for chimpanzees.     

Captive breeding, paternity determination, and genetic variation in chimpanzees (Pan troglodytes) in the Australasian region

DNA “fingerprinting” using polymorphic (CA)-repeat microsatellite markers was used to quantify the level of genetic variation present in chimpanzees (Pan troglodytes) in the Australasian region. These markers were also used to determine the paternity of chimpanzees born at Taronga Zoo over a 20-year period. The results suggested that the dominant male in the colony was responsible for siring most, but not all, of the offspring. Where the dominant male was excluded from paternity, the sire was identifiable if all candidate males were available for typing. This enabled us to prove the captive origin of offspring born in the colony during this period. Thus, microsatellite analysis was a useful tool for assignment of familial relationships and improving genetic management of breeding colonies.     

Longitudinal changes of dominance rank among the females of the Koshima group of Japanese monkeys

The changes of dominance rank among female Japanese monkeys of the Koshima group over a period of 29 years from 1957 were studied. The dominance rank order was relatively stable in the early population growing phase, while large scale-changes of dominance rank order occurred successively in the phase of population decrease brought about by the severe control of artificial feeding after 1972. Nevertheless, the rank order of several females of the highest status was stable. Furthermore, the reproductive success of these highest status females was high (Mori, 1979a;Watanabe et al., in prep.). Divergence of the dominance rank order fromKawamura's rules (Kawamura, 1958) was observed in the following respects: (1) Some females significantly elevated their rank depending on the leader males. (2) If mothers died when their daughters were still juveniles or nulliparous, the dominance rank of some of these offspring females was significantly lower than the mother's one. However 55% of daughters which lost their mothers at a young age inherited the mother's rank. (3) Dominance among sisters whose mother had died when at least one of the daughters was under 6 years old followed the rule of youngest ascendancy in 60% (Kawamura, 1958), and in 80% when both of the daughters were nulliparous at the mother's death. The mean rate of aggressive interactions for each female with subordinates to her was calculated by dividing the total aggressive interactions between the female in question and her subordinates by the number of subordinate females to the female in question. A female which showed a high rate of aggressive interactions with her subordinates was categorized as an “Attacker”, and a female showing a lower rate was categorized as a “Non-attacker”. Similarly, categories of “Attacked”, and “Non-attacked” were distinguished by using the rate of aggressive interactions with dominant females. Several females which were once categorized in one category in a year were repeatedly categorized in the same category over different years. The “Attacked” tended to be females of higher rank, and “Non-attackers” tended to be females of lower rank. “The second-higher-status females”, were “Attacked”, and their rank was unstable. In particular, females of lower rank within the lineage of the highest rank suffered this kind of severe status. Most of the daughters of these females showed a sharp drop of rank, and died when they were still at a young age, i.e. “the second-higher-status females” displayed low fitness. “Non-attackers” were significantly “Non-attacked”; i.e. they were females which showed a non-social attitude. Females which underwent a drop of rank tended to be “Non-attackers”. The most important factor which determined the females' rank was the memory of their dominance relations under the influence of their mother [dependent rank (Kawai, 1958)] in their early life during development. This finding corresponds well with the results in baboons obtained byWalter (1980); the target females of aggressive interactions by adolescent females were determined by the rank of the mothers when these adolescent females were born.     

Enhancement of juvenile Caribbean spiny lobsters: an evaluation of changes in multiple response variables with the addition of large artificial shelters

Shortage of natural crevice shelters may produce population bottlenecks in juvenile Caribbean spiny lobsters (Panulirus argus), a socially gregarious species. We conducted a field experiment to test enhancement of a local population of juvenile P. argus with the addition of artificial shelters (“casitas”) that mimic large crevices (1.1 m² in surface area and 3.8 cm in height). Changes in density and biomass of juvenile lobsters 15–50 mm carapace length (CL) were assessed with a multiple before-after control-impact (MBACI) analysis. Separate analyses were also conducted on small (15–35 mm CL) and large (35.1–50 mm CL) juveniles to assess size-related effects. First, we carried out 13 lobster surveys on nine fixed 1-ha sites over a shallow reef lagoon (“before” period). Then, we deployed ten casitas in each of five sites and left four sites as controls, and conducted 22 further surveys (“after” period). Deployment of casitas resulted in a sixfold increase in juvenile density (76% contributed by small and 24% by large juveniles) and a sevenfold increase in biomass (40 and 60%, respectively). Capture–recapture results revealed that enhancement was achieved not by promoting individual growth but by increasing survival, persistence, and foraging ranges of small and large juveniles. Casitas both mitigated shortage of natural shelter and increased sociality, allowing for cohabitation of smaller, more vulnerable juveniles with larger conspecifics that have greater defensive abilities. Casitas may help enhance local populations of juvenile P. argus in Caribbean seagrass habitats, typically poor in natural crevice shelters. The use of MBACI and the simultaneous assessment of multiple interrelated response variables may be a powerful analytical approach to test shelter limitation in other species and to examine the function of structural habitat in other systems.     

Responses of wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) toward seismic aftershocks in the Mahale Mountains National Park,Tanzania

This is the first report documenting the responses of wild chimpanzees (Pan troglodytes schweinfurthii) to seismic activities. During our long-term fieldwork in Mahale Mountains National Park, Tanzania, a high-intensity earthquake with a Richter magnitude of 6.8 occurred at 15:19 hours local time on 5 December 2005. During the main tremor, the chimpanzees displayed the “wraa” call, “scream,” and “pant bark” or “bark” vocalizations. Many mild aftershocks followed the main tremor, and the wild chimpanzees displayed a variety of responses to these. In several cases, they climbed trees or stopped activities such as grooming, moving, and feeding. These responses are similar to those previously reported in nonhuman primates. During the observations, a unique behavior, one never reported before was exhibited by a female chimpanzee. She placed her right palm on the ground giving the impression she was inspecting the trembling of the ground.     

Heterochrony and sexual dimorphism in the skull of the Liberian chimpanzee (Pan troglodytes verus)

Allometric methods and theory derived from principles of relative growth provide new and powerful approaches to an understanding of the nature and development of sexual dimorphism among living primates. The Frankfurt collection of Liberian chimpanzee skulls and mandibles provides a large skeletal sample from a single natural population of wild shot animals, including individuals of all ages and both sexes, and allows investigation of allometric and heterochronic patterns of sexual dimorphism. Univariate, bivariate, and multivariate analyses are utilized in this study in order to ascertain the ontogenetic nature of male-female differences in the skull of the Liberian chimpanzee. The results of univariate and multivariate analyses indicate that, while overall levels of sexual dimorphism in the chimpanzee skull are small, the greatest differences are in dimensions of the viscerocranium, while neurocranial dimensions and orbital size tend to be less dimorphic. Bivariate regressions of 21 cranial variables against basicranial length document positive allometry in many facial and mandibular dimensions, and isometry or negative allometry for most neurocranial dimensions. The data confirm previous work in chimpanzees and other anthropoid primates suggesting that males and females are “ontogenetically scaled” in most cranial traits. That is, males and females share the same cranial growth trajectories, although ending up at different points. Both rate and time hypermorphosis are suggested as underlying causes of ontogenetic scaling in the Liberian chimpanzee.     

Predation on mammals by the chimpanzee (Pan troglodytes)

With respect to prey selectivity and predation frequency, chimpanzees (Pan troglodytes) show local differences as well as diachronic variability within the same population. When data on predation from three long-term studies at Mahale, Gombe, and Tai are compared, some differences and similarities emerge; Mahale is more like Gombe than Tai in regard of prey selection but features of hunting at Tai with respect to predation frequency are not conspicuous. The most responsible factor for diversity in prey selectivity is a distinct “prey image” maintained by chimpanzees of different populations, although it is necessary to clarify in future studies why and how such tradition develops. Relative body size of chimpanzees to prey species and/or the degree of cooperation among members of a hunting party may explain the variability in prey size selected at each site, the latter influencing the frequency of successful hunts at the same time. Although various degrees of habituation and different sampling methods including artificial feeding might have obscured the real differences, recent data from the three populations do not seem to be biased greatly by such factors. Nevertheless, it is still difficult to make strict comparisons due to the lack of sufficient standardized data across the three populations on the frequency of hunting and predation. It is suggested that the size or demographic trend of a chimpanzee unit-group, especially the number of adult males included, necessarily influences its hunting frequency as well as its prey profile. It is also suggested that factors which bring these males together into a party (e.g. fruit abundance, swollen females, conflict between unit-groups etc.) strongly affect theactual hunting and kill rates. Other possible factors responsible for the local differences are forest structure (e.g. tree height), skilful “hero” chimpanzees, and competition with sympatric carnivorous animals. A total of at least 32 species have been recorded as prey mammals of chimpanzees from 12 study sites and the most common prey mammals are primates (18 species), of which 13 species are forest monkeys. Forest monkeys, colobine species in particular, are often the most common victims of the predation by chimpanzees at each site. We may point out a tendency toward selective hunting for the forest monkeys in terms of the selectivity of prey fauna among all three subspecies of chimpanzees, including populations living in drier environment. The mode of chimpanzee hunting seems to correspond to the highest available biomass of gregarious, arboreal monkeys in the forest, colobine species in particular. In contrast, bonobos (P. paniscus) are less carnivorous than chimpanzees, only rarely preying on a few species of small mammals. The sharp contrast of the two allied species in their predatory tendencies appears to have something to do with the differences in the structure of primary production between their habitats.     

Longitudinal analysis of length growth in the chimpanzee (Pan troglodytes)

The adolescent growth spurt in linear dimension in humans is considered to be unique among mammals, but few comparative studies have been done, even on chimpanzees. Growth of the summed length of crown to rump, thigh, and leg was studied longitudinally in 12 chimpanzees. We took body weight growth and reproductive maturation into consideration. Reproductive maturation was monitored by the swelling of sexual skin and menarche in females, and by testicular development in males. We applied two relationships found in humans between body length growth and the environment to the chimpanzees. The first relationship was the robustness of the growth spurt, meaning that the spurt is absent only in individuals under the most severe environmental pressure. Subjects maturing in a favorable or even mediocre environment are anticipated to show the growth spurt. The second relationship was catch-up growth, where, when the environment is ameliorated, growth may be accelerated to attain the target size. Catch-up growth at the end of the juvenile period may mimic the adolescent growth spurt. Results showed that subjects living under favorable conditions did not exhibit a growth spurt, and that it was only the subjects who had delayed growth in the juvenile period that showed a spurt in adolescence, the period when reproductive maturation occurred. Although we have concluded that chimpanzees do not have an adolescent growth spurt, except in cases of catch-up growth, this does not mean that they have a different growth pattern from that of humans. The absence of a growth spurt may be associated with adaptations to chimpanzee patrilineal society, where adolescent males are incorporated into the adult hierarchy at a low rank.     

Influence of chimpanzee predation on the red colobus population at Ngogo, Kibale National Park, Uganda

Frequent hunting of red colobus monkeys (Procolobus rufomitratus) takes place at all long-term chimpanzee (Pan troglodytes) study sites where both species are present. Red colobus are the most commonly selected prey of chimpanzees even when other monkey species are more abundant. In particular, the chimpanzee community at Ngogo, Kibale National Park, Uganda, preys heavily on red colobus monkeys: the chimpanzee hunting success rate is extremely high, and chimpanzees kill many individuals per successful hunt. Census data had suggested that the red colobus population is declining and that predation by chimpanzees may be contributing to this decline. In this paper, I address the impact of hunting on the red colobus population at Ngogo. To test the hypothesis that chimpanzee hunting is sustainable, I am using demographic data collected on red colobus monkeys over a period of 3 years, as well as fecundity and mortality data from previous studies of this species. I apply matrix models and vortex analyses using a sensitivity analysis approach to project future population development. Results show that current rates of hunting are not sustainable, but that chimpanzees are neither more “noble”, nor more “savage” than humans are, but that they also hunt to ensure maximum benefit without regard for the consequences for the prey population.     

Leaf-grooming by a wild chimpanzee in Mahale

During the course of systematic observations of the leaf-grooming behavior by the chimpanzees (Pan troglodytes) of the Mahale Mountains National Park, Tanzania, I recovered a louse from a leaf “groomed” by an adult male chimpanzee after a typical leaf-grooming session. During the leaf-grooming session I observed a small object on his lower lip. He picked up a leaf, transferred the small object from his lip to the leaf, folded the leaf and crushed the folded side of the leaf with his thumb. I present this observation as further evidence of the “squashing ectoparasites” hypothesis for leaf-grooming.     

A study on the social structure and dispersal patterns of hamadryas baboons living in a commensal group at Taif,Saudi Arabia

Three levels of hamadryas social structure—the one male unit (OMU), the band, and the troop—have been observed at all sites studied, but a fourth—the clan—has been observed at only one site, Erer-Gota, Ethiopia, during a longitudinal check of the dispersion of identified individuals. The clan is important since it appears to provide the basis for male philopatry, although comparative data is needed from other sites to confirm this. We studied a huge commensal group of hamadryas baboons (over 600 animals) in Saudi Arabia. We put ear tags on baboons between 1998 and 2004 and analyzed social structure, relying on the interactions of these tagged animals by focusing especially on their dispersal patterns from OMUs. OMU membership tended to be looser than that of the Ethiopian hamadryas. Females tended to shift between OMUs on an individual basis in our study group, whereas the collapse of an OMU was a major occasion of adult female transfer in Ethiopia. We found neither stable bands (a “band” in our study group was defined as a regional assemblage of OMUs) nor clans that lasted for several years. Some OMUs moved and transferred into neighboring areas over both the short and long term. Further, some post-adolescent males appeared to move out of the study area. The ratio of adult females in an OMU in our study group was larger than for any other documented study site, and this may be the reason for enhanced female transfer between OMUs. A large proportion of the adolescent females showed no clear membership to OMUs, and no “initial units” (commonly observed in Ethiopia) were discernible. The ease with which young males acquired adult females at the study site must have disrupted the formation of a clan, a “male-bonded society.”     

Decline in energy storage in the Antarctic minke whale (<Emphasis Type="Italic">Balaenoptera bonaerensis</Emphasis>) in the Southern Ocean

The annual trend in energy storage in the Antarctic minke whale was examined using catch data from all 18 survey years in the Japanese Whale Research Program (JARPA). Regression analyses clearly showed that blubber thickness, girth and fat weight have been decreasing for nearly 2 decades. The decrease per year is estimated at approximately 0.02 cm for mid-lateral blubber thickness and 17 kg for fat weight, corresponding to 9% for both measurements over the 18-year period. Furthermore, “date”, “extent of diatom adhesion”, “sex”, “body length”, “fetus length”, “latitude”, “age” and “longitude” were all identified as partially independent predictors of blubber thickness. The direct interpretation of this substantial decline in energy storage in terms of food availability is difficult, since no long-term krill abundance series is available. However, an increase in the abundance of krill feeders other than minke whales and a resulting decrease in the krill population must be considered as a likely explanation.     

Peering in mature, captive bonobos (Pan paniscus)

“Peering”—close-proximity staring at the mouth of another—was observed in ten (three males and seven females) mature (at least 7 years old) bonobos (Pan paniscus) living in three social groups at the San Diego Zoo and Wild Animal Park. Instantaneous scan samples, taken at 2-min intervals, over a three-and-a-half year period, yielded 617 observations of peering (1.4 per observation hour). Food was exchanged in only 15 of these scans. Peering was most often performed by younger animals and was primarily directed toward older females (“matrons”). In a given dyad, the animal more likely to peer at the other was also more like to both peer and be peered at if they frequently groomed and infrequently displayed aggression at a given female. An adolescent male showed the highest frequency of peering when living with two older females, but dropped to adult male levels when later housed with two younger (albeit mature) females. A reversal in which animal was more likely to peer, follow, and groom occurred in one female dyad, after the birth of the younger animal's first infant. After a similar birth in the other group, no such changes were observed. We discuss how these and related findings, in conjunction with what is known of the social structure of this species, suggest that one possible function of peering in bonobos may be as a signal acknowledging female status.     

Reproductive patterns in mantled howler monkeys: Estrus,mate choice and copulation

The present study was undertaken to evaluate non-random mating patterns in two groups of mantled howler monkeys in two tropical dry forest habitats. Sexual dimorphism, female estrus stage, male dominance rank, sexual solicitations and copulations were assessed. Males are significantly larger than females, but female weight varies more than male weight. The length of female estrus cycles is comparable in both habitats, but females in the more strongly seasonal habitat demonstrate greater estrus synchrony relative to their numbers. Males solicit potential mates more frequently than females, a pattern explained by the relatively high rate of sexual solicitation by high-ranking males. Females in “peak” estrus solicit “alpha” males, while females in other stages of estrus solicit males equally by rank. Intersexual aggression occurs rarely, and “forced copulations” are attempted but, apparently, are unsuccessful. Sexual solicitations by “alpha” males and “peak” estrus females are most likely to lead to copulation, and “alpha” males are more likely to copulate than “gamma” males. In general, latencies from first solicitation to copulation are expensive in time, especially for high-ranking males. Estimated annual reproduction success favors high-ranking males, and results indicate that male and female mating behavior is mutually coordinated and controlled.     

Mating patterns,mate choice,and birth season heterosexual relationships in free-ranging rhesus macaques

Birth season adult heterosexual nonkin relationships of 50 free-ranging female rhesus macaques (Macaca mulatta) in two social groups at Cayo Santiago, Puerto Rico were examined using focal follow (289 hr) and ad lib data. Eighty-eight percent of subjects had at least one relationship characterized by particularly high frequencies of spatial proximity, grooming, or both. These were designated “friendships.” Males intervened in aggressive interactions more frequently on behalf of Friends than non-Friends. Female aggressive support of males was extremely rare. Higher-ranking males experienced more friendships than lower-ranking males. High-ranking females had higher-ranking Friends than low-ranking females. Older females had higher-ranking Friends than younger females. Females groomed high-ranking Friends more than they were groomed by them, whereas they groomed low-ranking Friends less than they were groomed by them. In one social group, high-ranking females were more likely than low-ranking females to groom their Friends more than they were groomed by them. Males were more responsible than females for spatial proximity maintenance in 9 of 14 Friend dyads for which sufficient data were available. Neither male nor female dominance rank affected responsibility for proximity maintenance in Friend dyads. Eight of 24 females had friendships with males with whom they had completed copulations during their conception peri-ovulatory period of the preceding mating season. Two of 19 females completed peri-ovulatory copulations with Friends during the following mating season. Friendship was not correlated with either of two demonstrated female mate choice indicators: (1) proximity maintenance during estrus; or (2) cooperation with male “hip-grasp” courtship attempts. Males directed “muzzle-up” courtship signals at lower rates toward Friends than toward non-Friends. These and other investigators' results indicate that (1) protection from aggression is the primary benefit to female rhesus macaques of birth season heterosexual relationships; (2) the most effective protectors are in greatest demand as Friends; and (3) friendship has no effect or an inhibitory effect on mate choice in this species. Benefits to males of friendships were not apparent from this study but may include coalitional support against lower-ranking males.