Male dominance rank and reproductive success in primate groups

The relationship between male dominance rank and reproductive success has been a topic of interest since the beginning of primatology. From a theoretical point of view the existence or absence of this relationship has great implications with respect to the meaning of dominance rank and more general of the social relationships between individuals within social groups. Until fairly recently mating behaviour has been used as an indicator of reproductive success, but these two variables need not be correlated. The relations between mating success, reproductive success, and dominance rank indicate whether selective mating is involved (different mating partners at different phases of fertility, for example through male contest or female choice). With the development of genetic techniques to determine reproductive success directly, it has now become possible to investigate these relations. In our study on wild long-tailed macaques (Macaca fascicularis) we find a relatively strong correlation between rank and reproductive success which is attributable to selective mating by the alpha male during fertile periods of the females. In most previous studies no such clear results have been obtained, and we discuss the differences in outcome in relation to the study conditions, group sizes, and possible differences between species in terms of reproductive strategies.     

Proximate and ultimate causes of reproductive suppression among female yellow baboons at Mikumi National Park,Tanzania

Attack coalitions, whereby two or more individuals attack another animal simultaneously, were studied in 53 sexually mature female baboons from one small and two large troops at Mikumi National Park, Tanzania. For 14 months, three observers recorded coalition behavior throughout the day, using focal group sampling on a behavior-dependent basis. Coalition rates were considerably greater in the two large troops than in the small troop, and primarily involved natal females rather than males. Coalition frequencies corresponded closely to measures of reproductive synchrony, both within and between troops. Involvement in attack coalition behavior depended on the reproductive states and ranks of both attackers and recipients. The first few females to give birth in the cohort received particularly high amounts of coalition aggression just before and immediately after giving birth. Receipt of attack coalitions was significantly associated with an increased number of cycles to conception and longer interbirth intervals, as well spontaneous abortion, premature birth, and prolonged gestation. A parallel study on survivorship patterns of immature baboons at Mikumi provided evidence for ultimate cause of the observed coalition patterns. These combined results suggest that attack coalition behavior is a form of reproductive competition whereby females attempt to suppress the reproduction of others at predictably competitive times and thereby reduce the competition their own infants face from the time they are weaned.     

Male rank and optimal lek size

Widemo and Owens presented a model that calculates the expectedcopulation rates of males on leks of a range of sizes. Theyclaim that a negative relationship between lek size and malemating skew will result in low-ranking males having greateroptimal lek sizes than higher ranking rivals. Widemo and Owensoffered no proof of their claim, and their model assumes thatthe rank of a male does not change as lek size increases, whereasin reality, rank may change as more males arrive. We presenta general model that allows rank to change as lek size increases.We show that the crucial determinant of whether optimal leksize increases with male rank is whether relative competitive differencesincrease with lek size. Contrary to the claim of Widemo andOwens, the relationship between skew and lek size has no directbearing on the optimal levels of aggregation of males of differentrank. We show that a negative relationship between skew andlek size can exist even when high-ranking males have the greatestoptimal lek sizes.     

Variability and stability in the rank relations of nonhuman primate females: Analysis by computer simulation

This paper summarizes results of simulation analyses aimed at elucidating the way in which two important dimensions of female rank orders—nepotism and age-gradedness—are affected by variation in individual strategies of rank acquisition on the one hand and by variation in population-wide demographic parameters (eg, birth rate, death rate, etc) on the other. Female rank orders in simulated primate groups were characterized by substantial variability on a year-to-year basis in both nepotism and age-gradedness owing to stochastic processes alone. Although, in general, groups characterized by strongly nepotistic rank orders were not also strongly age-graded, quantitative measures of these two rank-order dimensions for individual groups showed little stability over intervals of time exceeding 3 years. Surprisingly, changes in birth and death rates did not strongly affect either nepotism or age-gradedness so long as average group size remained constant. However, in populations characterized by expanding or declining group size, variation in demographic parameters did indeed exert a powerful influence on the structure of female rank orders. Both nepotism and age-gradedness were also strongly affected, and in quite different ways, by changes in the rules of rank acquisition by females. In sum, the present analyses suggest that differences between species in the degree of nepotism and age-gradedness characteristic of their female rank orders does not in and of itself constitute evidence that the females of those species use markedly different strategies of rank acquisition.     

Age at maturity in wild baboons: genetic, environmental and demographic influences

The timing of early life-history events, such as sexual maturation and first reproduction, can greatly influence variation in individual fitness. In this study, we analysed possible sources of variation underlying different measures of age at social and physical maturation in wild baboons in the Amboseli basin, Kenya. The Amboseli baboons are a natural population primarily comprised of yellow baboons (Papio cynocephalus) that occasionally hybridize with anubis baboons (Papio anubis) from outside the basin. We found that males and females differed in the extent to which various factors influenced their maturation. Surprisingly, we found that male maturation was most strongly related to the proportion of anubis ancestry revealed by their microsatellite genotypes: hybrid males matured earlier than yellow males. In contrast, although hybrid females reached menarche slightly earlier than yellow females, maternal rank and the presence of maternal relatives had the largest effects on female maturation, followed by more modest effects of group size and rainfall. Our results indicate that a complex combination of demographic, genetic, environmental, and maternal effects contribute to variation in the timing of these life-history milestones.     

Grooming Between Male Chimpanzees at Ngogo,Kibale National Park. II. Influence of Male Rank and Possible Competition for Partners

Allogrooming contributes to the development and maintenance of social relationships, including those that involve alliances, in many primate species. Variation in relatedness, dominance rank, and other factors can produce variation in the value of others as grooming partners. Several models have been developed to account for variation in the distribution of grooming in relation to dominance ranks. These start from the premise that individuals are attracted to high-ranking partners, but time limits, direct competition, and prior grooming engagement between high-ranking individuals can constrain access to them. Sambrook et al. (1995) formalized some of these models and showed the importance of taking group size variation into account when assessing them. Chimpanzees form multimale communities in which males are the philopatric sex. Males commonly associate and groom with each other; they also form dominance hierarchies and form alliances that influence dominance ranks and mating success. Both male rank and the rank distance between partners are significantly correlated with the distribution of grooming between males in an extremely large chimpanzee community at Ngogo, Kibale National Park, Uganda, that has more males than any other known community. High-ranking males had more grooming partners than mid- or low-ranking males. Grooming predominantly went up the dominance hierarchy, but was also concentrated among males that were close in rank. Rank and rank distance apparently both affected grooming independently of reciprocity in grooming and independently of the frequency with which males associated in temporary parties. However, the data do not clearly indicate how constraints on access to partners might have operated. Published data from a smaller chimpanzee community at Mahale show no rank or rank distance effect on male grooming. These results and earlier, conflicting findings on the association between dominance rank and grooming in male chimpanzees indicate that variation in group size, i.e., the number of males per community, probably influences the strength of any such effects, as happens for grooming between females in several cercopithecine species. Data on coalitions at Ngogo support the argument that high-ranking males are valuable social partners, and similarity in strategies of alliance formation may influence the distribution of grooming.     

Alternative male reproductive strategies in the peacock blenny

A 4-year study (1989–1992) of a population of Lipophrys pavo living in a brackish lagoon (Maugio, France) revealed three types of male during the spawning season (April to September): (i) nest-breeder males which are territorial, and have well-developed secondary sexual characteristics; (ii) non-functional males which were abundant only in 1991 when population density was very high and (iii) 1-year-old kleptogamic males with small body size which possess neither secondary sexual organs nor a testicular gland, although their testes are functional and hyperdeveloped (up to 10.5% of carcass weight). The percentage of kleptogamic males was fairly stable in the population (from 7.9 to 10.0%) and highly variable in their age class (from 8.5 to 51.3%). It is suggested that kleptogamy is a conditional strategy adopted by smaller males.     

Male rank,reproductive behavior,and reproductive success in free-ranging rhesus macaques

Paternity assessment through DNA fingerprinting by synthetic oligonucleotide probes was applied to one birth cohort in a social group of free-ranging rhesus macaques (Macaca mulatta) on Cayo Santiago. The 11 group males and 9 males from other groups were observed mating with the females. Paternity was determined for 11 of the 15 infants. Male dominance rank was not associated with reproductive success. High-ranking resident males (N=5) sired 27% of the infants born during a one-year study. Four of the 11 infants of known paternity were sired by males of other social groups. The four infants of unknown paternity were sired either by males not observed mating with the females or the low-ranking male who was not fingerprinted. Male dominance rank was not associated with reproductive activity during conception cycles. These results suggest that the effect of rank on male reproductive success is not a predictable correlation, but a conditional probability.     

Adult Male Scent-Marking in Lemur catta and Eulemur fulvus rufus

Scent-marking and olfactory communication are used extensively by prosimians and can provide spatial and temporal records of group movement and behavior. We compare rates of male scent-marking in relation to reproductive seasons, male dominance rank, and habitat use in two related prosimians: Lemur catta and Eulemur fulvus rufus. We collected scent-marking data on adult male Lemur catta at Beza-Mahafaly Reserve (dry forest), and on Eulemur fulvus rufus at Ranomafana National Park (rainforest), Madagascar. In Lemur, rates of overall scent-marking differed significantly by reproductive season, with higher rates occurring in mating and lactation/migration periods, whereas in Eulemur, reproductive season did not appear to affect scent-marking rates. Dominance rank of male Lemur catta did not affect rates of scent-marking. Among male Eulemur fulvus, dominance relations were not apparent; however, 2 of the 5 focal males scent-marked somewhat more frequently during the mating season and also experienced greater mating success. In Lemur catta, higher rates of scent-marking in the mating season may relate to indirect reproductive competition during a period of high aggression, while such mating competition was not as marked in Eulemur fulvus. Furthermore, higher rates of marking in resident male Lemur catta during male migration may correlate with vigilance toward immigrating males. Greater overall scent-marking rates in ring-tailed lemurs may relate to extensive intergroup home range overlap and no area of exclusive use, whereas the red-fronted lemur groups tended to forage in areas of their home range where little-to-no intergroup overlap occurred.     

Adaptation to grass-eating in gelada baboons

Gelada faecal samples were analyzed for nutritional content and for particle size, and compared with similar data forPapio baboons, cattle, and zebra. Particle size in gelada is similar to that for zebra, larger than that for cattle and smaller than that for baboons. Gelada and baboons are less efficient than ungulates at extracting protein from their diet. The data on energy extraction are less easy to interpret and appear to be confounded by dietary and seasonal factors. It is suggested that gelada may be too large to compete effectively with ruminants in low altitude grassland habitats under the climatic conditions that have prevailed in eastern Africa since the late Pleistocene.     

The association between rank and reproductive success of male rhesus monkeys

The paternity of 202 of 220 offspring of rhesus monkeys housed in six separate half-acre field cages at the California Primate Research Center was determined by genetic marker techniques. Reproductive success of the adult males was statistically significantly correlated (r = 0.76, P <0.0005) with rank. Marked systematic changes in reproductive success were, however, observed for two adult males during the three years covered by this study. Based upon these results and those of another study, it is argued that changes in rank position typically follow, rather than precede, changes in reproductive success.     

Female weight and reproductive condition in a population of olive baboons (Papio anubis)

The study reported here tested Altmann's prediction [Baboon Mothers and Infants. Cambridge, Harvard University Press, 1980] that lactating female baboons endure a weight loss. Data from 64 adult female olive baboons (Papio anubis) residing in six troops in Kenya revealed that reproductive condition was related to weight. Lactating females weighed less and pregnant females weighed more than cycling females. There was a negative correlation between the weight of cycling females and the number of months postweighing to their next conception. These results indicate that lactation in wild baboons imposes energy costs that result in lost weight. It is suggested that female baboons may have to surpass a minimum weight threshold prior to resumption of postlactational cycling and that nutritional status is more influential than rank in affecting female reproductive success.     

Body composition in baboons: Evaluating a morphometric method

The objective of this study was to determine whether noninvasive morphometric measurements of olive baboons (Papio cynocephalus anubis) can reliably predict lean body mass and fat mass in this species. Crown-rump length, triceps circumference, and skinfold measures at the neck, subscapular, suprailiac, and triceps sites were obtained prior to necropsy from 21 clinically normal infant baboons at 18 weeks of age and from 22 clinically normal adolescent baboons at 5 years of age. At necropsy, the lean body mass and body fat mass were measured using gravimetric methods. Pearson's correlations and regression analysis were used to test the relationship between direct measures of lean and fat mass obtained at necropsy with calculated estimates based on morphometric measures obtained prior to death. Our null hypothesis was that the morphometric measures (individually or in combination) were not related to direct measures of fat mass or lean mass. Non-linear and multiple regression estimators, by age and gender, were derived and provided increased predictability. Our results indicate that (1) morphometric measures can accurately predict lean body mass in male and female baboons; (2) morphometric measures used to predict lean body mass change with age; (3) morphometric measures are strongly associated with body fat mass at 18 weeks of age but are not as strongly associated with body fat mass in 5-year-old baboons; (4) triceps circumference provides the best single indicator of lean body mass for both genders and age periods; (5) baboons are like humans in that adolescent females tend to accumulate body fat while males of the same age tend to develop lean mass; and (6) combinations of these morphometric measurements explain between 70% and 100% of the variability and can be used to estimate lean and fat mass in baboons.     

Social relationships and ritualized greetings in adult male baboons (Papio cynocephalus anubis)

Greetings involving exchanges of ritualized sexual gestures are a common form of interaction among adult male baboons, although relatively little attention has been paid to them. In this study, we investigate how greetings reflect important aspects of the male's social relationships, including dominance rank, age/residence status, and cooperative tendencies. The results are based on over 600 greetings among 12 adult males recorded during a 4-month study of a troop of wild olive baboons near Gilgil, Kenya. Four of the adult males were older, lower-ranking, long-term residents, which frequently formed coalitions to take estrous females away from the eight young, higher-ranking males. Virtually all dyads greeted: greetings occurred more than twice as often as other types of male-male interactions; and nearly all greetings occurred in a neutral context, in which there was no resource at stake. The percentage of greetings completed, the frequency with which different gestures were employed, and the roles adopted by each male varied significantly across old-old, old-young, and young-young dyads. Greetings between young adult males were often interrupted or actively resisted, consistent with their unstable and ambiguous dominance relationships. Greetings between old-old dyads were usually completed and appeared consistent with their cooperative relationships. One pair of old males formed a stable, reciprocal coalition against young males, and this pair's greetings showed remarkable symmetry of roles. Greetings, we hypothesize, function to allow males to negotiate important aspects of their relationships, including cooperation.     

Agonism,ranking, and the social behavior of adult male baboons

An ingrained technique in primatological analysis is to rank individual animals in a hierarchical system that is usually but not always based on the outcome of agonistic encounters. An explication of group structre and function is then sought by correlation these rankings with aspects of behavior. When the behavior of the adult males in a free-ranging troop of olive baboons is examined, ranking is found to be inadequate to represent the behavioral variability existing between individuals. The relationships maintained by these animals is analyzed, emphasizing amounts of time devoted to specific activities, and it is recommended that affiliative rather than agonistic behavior be stressed in future studies.     

Dental eruption schedules of wild and captive baboons

Dental eruption schedules previously used to age wild baboons have in the past derived from studies of captive animals housed under standard conditions and fed standard laboratory diets. This paper reports for the first time eruption schedules derived from wild baboons, the yellow baboons (Papio hamadryas cynocephalus) of Mikumi National Park, Tanzania, and compares these schedules with those of other baboon subspecies inhabiting both similar and dissimilar environments. Eighteen males and twelve females from the Viramba groups, ranging in age from 21 to 103 months, were trapped, and dental impressions and notes were made of the state of eruption of each tooth. Eruption of all teeth were delayed at Mikumi relative to the baboon standards derived from the captive animals at the Southwest Foundation for Biomedical Research, San Antonio, Texas. Teeth of the canine-premolar 3 complex and third molars were most delayed, erupting up to a year and a half later than their counterparts from captive animals. Comparison with data on hamadryas baboons from Erer-Gota in Ethiopia revealed that both the hamadryas and yellow subspecies of baboons, with different genetic backgrounds and living under markedly different environmental conditions, followed the same schedule. This constancy of developmental schedules suggests that these Mikumi data may reasonably be used as standards for other wild baboon populations and that acceleration of dental maturation, as well us maturation of other somatic systems in captivity, is another manifestation of the short-term adaptive plasticity of the baboon species as a whole.     

Intertroop variation in the frequencies of ABO alleles in a population of olive baboons

Blood and saliva were obtained from 183 olive baboons captured on the Kekopey Ranch in the Central Rift Valley of Kenya. Samples were returned to the Laboratory of Biological Anthropology and tested for ABO antigens. A uniquely high frequency of the amorph is reported. Intertroop variation at this locus appears to be very high, although within the ranges reported by some previous investigators.     

Formula-fed and breast-fed baboons: Weight growth from birth to adulthood

The objective of this study was to compare the immediate and long-term effects of type of infant diet (formula-fed vs. breast-fed) on the weight growth of baboons. Longitudinal (serial) weight data were collected from 44 savannah baboons (Papio cynocephalus anubis) for the total preadult period of growth from birth to approximately 8 years of age. Fifty percent of the sample (11 females and 11 males) were fed a standard infant formula by a nursery care technician for the first 18 weeks of postnatal life. The remaining 11 females and 11 males were breast-fed by their mothers for an equivalent period of time. After 18 weeks of age the animals were removed from their infant environments (either nursery or mother-reared) and assigned to one of four age-cohort social groups. Each age-cohort group was balanced for infant treatment and gender. The animals remained in these social groups for the duration of the project and were provided nutritionally identical postweaning diets. No significant weight differences were found between the two rearing conditions for either gender during the initial 15 weeks of growth. Following this period, however, females of breast-fed environments averaged greater weight at all ages than their bottle-fed peers. The intensity of the adolescent growth spurt was also slightly greater among the female breast-fed group. No postinfant average weight-per-age disparity was observed between the male treatment groups.