Male and Female Reproductive Success in <Emphasis Type="Italic">Macaca sylvanus</Emphasis> in Gibraltar: No Evidence for Rank Dependence

The relationship between social rank and reproductive success is one of the key questions for understanding differences in primate social group structures. We determined the paternity of 18 infants in a social group of Barbary macaques (Macaca sylvanus) born over a period of 6 yr in the provisioned, free-ranging colony in Gibraltar. We successfully used 13 pairs of primers of variable microsatellite loci to amplify DNA from blood and hair samples and applied the computer programs CERVUS 2.0 and KINSHIP 1.3 to assign paternity to 13 candidate males. We collected data for 19 females that had given birth to 66 infants over a period of 7 yr. We used paternity analyses and female birth records to test the hypothesis that social rank is correlated with reproductive success. Results showed that numbers of paternities and maternities were equally distributed among all reproducing individuals in the social group regardless of rank. Subadult males reproduced as often as adult males. High-ranking females did not start to reproduce earlier than low-ranking females. Interestingly, there was a tendency toward a positive correlation between the ranks of mothers and the ranks of the corresponding fathers. It might be concluded either that a correlation between social rank and reproductive success is generally absent in Barbary macaques or that artificially favorable environmental conditions in Gibraltar preclude any correlation between social rank and reproductive success.     

Conspecific reproductive success affects age of recruitment in a great cormorant, Phalacrocorax carbo sinensis, colony.

Few studies have addressed the proximate factors affecting the age at which individuals of long-lived bird species are recruited into the breeding population. We use capture-recapture analysis of resightings of 16 birth cohorts of colour-ringed great cormorants, Phalacrocorax carbo sinensis, in a Danish colony to assess the evidence for two hypotheses: conspecific attraction (earlier recruitment when the colony is large) and conspecific reproductive success (earlier recruitment following years of high breeding success). For both males and females, conspecific reproductive success was the most important covariate explaining the interannual variation in age of recruitment; colony size was also important for females. These covariates explained nearly 60% of the year-to-year variation for both sexes. The age of recruitment increased for cohorts born after 1990, and this increase was correlated with a decline in breeding success in the colony; we interpret this as an indirect and delayed density-dependent effect. Females were recruited earlier than males (mean age of recruitment for cohorts born before 1990: 2.98 years versus 3.53 years); the most plausible reason for this is a skewed sex ratio in favour of males in the adult population. Recruitment of males may thus, to some extent, be constrained by the availability of females. This study provides the first evidence that conspecific reproductive success can affect the age at which individual birds start to breed.     

Female competition in chimpanzees

Female chimpanzees exhibit exceptionally slow rates of reproduction and raise their offspring without direct paternal care. Therefore, their reproductive success depends critically on long-term access to high-quality food resources over a long lifespan. Chimpanzee communities contain multiple adult males, multiple adult females and their offspring. Because males are philopatric and jointly defend the community range while most females transfer to new communities before breeding, adult females are typically surrounded by unrelated competitors. Communities are fission–fusion societies in which individuals spend time alone or in fluid subgroups, whose size depends mostly on the abundance and distribution of food. To varying extents in different populations, females avoid direct competition by foraging alone or in small groups in distinct, but overlapping core areas within the community range to which they show high fidelity. Although rates of aggression are low, females compete for space and access to food. High rank correlates with high reproductive success, and high-ranking females win direct contests for food and gain preferential access to resource-rich sites. Females are aggressive to immigrant females and even kill the newborn infants of community members. The intensity of such aggression correlates with population density. These patterns are compared to those in other species, including humans.     

Hierarchical organization of femaleMacaca radiata in captivity

Females of several species of macaques form cohesive matrilineal units in which all members share a collective status. The relationship between rank and kinship inMacaca radiata has not previously been studied. Analysis of observations of social interactions in a large and stable captive group ofM. radiata and longitudinal study of kinship and reproductive success indicate that with few exceptions a matrilineal dominance hierarchy exists in that group. Four young, upwardly mobile females are responsible for the exceptions. Contrary to the pattern noted in other species of macaques, several adult females outrank their daughters. Old age and deteriorating physical condition of mothers appear to be associated with mother-daughter rank reversals. The age and lineage size of females when they entered the group have had a lasting impact. Females who entered the group as adults have achieved higher rank and greater reproductive success than females who entered the group as juveniles without relatives. This research was conducted at the California Primate Research Center in Davis, supported by USPHS grant RR00169.     

Variation in selection pressure acting on body size by age and sex in a reverse sexual size dimorphic raptor

Body size strongly influences fitness, with larger individuals benefiting in terms of both greater productivity and survivorship; for reverse sexual size dimorphic (RSD) species, this relationship may be more complex. We examined the selection pressures acting on body size in male and female Merlins Falco columbarius to assess whether larger or smaller individuals of this RSD species were favoured in terms of survival and breeding performance. For males and females there were clear links between body size and survival but the exact relationship varied by sex. Among males, birds that survived each year class were larger than those that died and yearlings were on average smaller than older birds, but there were no measurable differences among adult males (age 2+). Among females, larger individuals aged 1 and 2 years were more likely to survive, but this size‐based pattern was not apparent in older age classes. Size early in life predicted the lifespan in male Merlins but not as strongly as for females and not for the largest individuals. Reproductive performance based on brood size was not associated with body size in either males or females, but there was a weak positive relationship between female body size and lifetime reproductive success. Selection appears to favour larger males and females but there is no indication that the population is evolving towards bigger individuals, perhaps in part due to selection against the largest birds. Increased survival may allow larger and higher quality individuals to occupy higher quality territories as they age and thereby to accrue greater lifetime reproductive success in the process.     

Two girls for every boy: The effects of group size and composition on the reproductive success of male and female white-faced capuchins

Many factors have been hypothesized to affect the size and adult sex ratios of primate groups and these, in turn, have been argued to influence birth rates. Using park-wide census data collected on a population of capuchins over a 25-year period, we examined whether group size and adult sex ratio affect the per capita reproductive success of male and female white-faced capuchins (Cebus capucinus) in Santa Rosa National Park, Costa Rica. We found that the reproductive success of females (measured as the observed minus the expected ratio of immatures to adult females in the group) decreased with increasing group size, whereas that of males was independent of group size. The proportion of adult males residing in groups had significant, yet contrasting effects on males and females. Male reproductive success was negatively associated with the proportion of males residing in groups whereas female reproductive success increased with the proportion of males. The latter finding supports the intersexual conflict hypothesis, which suggests that a conflict of interest occurs between males and females over adult sex ratios. The effects of group size and composition on the reproductive success of capuchins, a male-dispersed omnivorous species, are similar to those reported for howlers, a bisexually-dispersed folivorous species. One common factor between these taxa is that groups with low ratios of males to females are at greater risk of takeovers and resultant infanticide. Our results suggest that regardless of dietary preference and dispersal pattern, the threat of infanticide can constrain primate group size and composition.     

Reproductive success in relation to dominance rank in the absence of prime-age males in Barbary macaques

In some primate species dominance rank of males is correlated with reproductive success, whereas in other species this relationship is inconsistent. Barbary macaques (Macaca sylvanus) live in a promiscuous mating system in which males are ranked in a dominance hierarchy that influences their access to females. High-ranking males usually monopolize fertile females during their estrous period and show increased mating activities. Subadult males generally rank below adult males. For Barbary macaque females in the Gibraltar colony, there was no correlation between dominance status and reproductive success. Paternity data for 31 offspring collected over four consecutive breeding seasons were used to test whether male social rank was associated with reproductive success and whether reproductive success was mainly confined to a small number of males. Genetic variation was assessed using 14 microsatellite markers for a dataset of 127 individuals sampled in all five social groups of the Gibraltar colony. Paternity analysis was conducted for offspring in one social group only, where all in-group males were sampled. Eighty-three percent of the offspring could be assigned to an in-group candidate father; none of the extra-group males appeared to have sired an infant. Male dominance rank was not found to contribute to the observed variation in male reproductive output. Fifty-nine percent of the offspring was sired by two low-ranking males, whereas the two top-ranking males sired one-fifth. A highly significant correlation was found for male age and dominance rank. Reproductive success of subadult males might be explained by the gap in the age distribution of male group members. These missing prime males are usually regarded as serious competitors for older males. Subadult males may have gained easier access to females in their absence. In addition, the presence of inbreeding avoidance mechanisms, which might also have overpowered possible rank effects, cannot be excluded.     

Male-female affiliative relationships in naturally occurring ringtailed lemurs (Lemur catta) at the Beza-Mahafaly Reserve,Madagascar

Affiliative behavior between adult male and female ringtailed lemurs was examined as part of a project concerning male affiliation with conspecifics of all age/sex classes. Males in three social groups were studied over a 12 month period. Male-female preferred partnerships existed, and were variable according to reproductive season. Dominance rank, age, or tenure of the male did not appear to affect either the number of partnerships or frequency of affiliative behaviors that males had with females. However, males residing in groups with fewer males exhibited both higher frequencies of affiliative interactions with females and were nearest neighbors to females more often than males living in a group containing more males. Females were found to be responsible for proximity maintenance of male-female dyads in the majority of cases. Neither reproductive season nor seasonal availability of food resources strongly affected the frequency of affiliative interactions between males and females, it is proposed that an important aspect of successful group membership for male ringtailed lemurs relates to the development of social relationships with adult females. Males can benefit from such relationships in terms of greater centrality to the spatial core of the group, which can result in enhanced predator protection, greater opportunities for social contact, and potentially greater access to estrous females. © 1996 Wiley-Liss, Inc.     

Sexually dimorphic head sizes and reproductive success in the sleepy lizard Tiliqua rugosa

In 1993, 458 males and 346 females of the largve Australian skink, Tiliqua rugosa , were captured in a study area near Mt. Mary, South Australia. Females were significantly longer than males, although there was broad overlap in snout-vent length measures. Males had significantly longer and broader heads than females of equivalent snout-vent length. In the spring some, but not all, lizards formed monogamous pairs. Pairing was used as an indirect indicator of reproductive success. When all adult males were considered there was no significant difference in head size between those found paired or unpaired. However, among small adults, paired males had signficantly broader heads than unpaired males. This supports the hypothesis that head size is under sexual selection. Individuals with wider heads could be more successful in male-male combat where jaws are the major offensive weapon. Younger, smaller males with a wise head could gain mates at an earlier age. Females showed a different pattern. In all females, and most strongly amongst larger size classes, paired females had significantly larger heads than unpaired females. An explanation is that larger heads somehow reduce the chance that a female will skip a year of reproduction, although the mechanism is not clear.     

Modeling Developmental Class Provides Insights into Individual Contributions to Infant Survival in Callitrichids

Cooperative breeders live in social groups in which individuals in an age–sex class vary in reproductive development due to reproductive dominance by a few individuals in each group. Among callitrichids, adult males have been implicated in driving group reproductive output, but uneven sampling efforts, the underlying effects of group size, and pseudoreplication at the group and species levels are confounding variables in these analyses. We examined the drivers of group reproductive output in callitrichids by 1) conducting a meta-analysis of published studies of callitrichid group composition; 2) assigning developmental class based on reproductive morphology; and 3) using multivariate modeling to test whether the proportion of individuals of each developmental class predicts the presence and the number of surviving offspring among free-ranging Weddell’s saddleback tamarins (Leontocebus weddelli) and emperor tamarins (Saguinus imperator) in Peru. The meta-analysis revealed that the number of adult females and group size, but not the number of adult males, are significantly correlated with group reproductive output. Statistical models of the new dataset revealed that the proportion of primary breeding males, primary breeding females, and group size predicted whether groups had surviving infants, and that only the proportion of primary breeding females and group size predicted the number of surviving infants. Thus, primary breeding males appear to be necessary for groups to raise any infants, but a higher proportion of primary breeding females and a larger group size increase group reproductive output overall.     

Spacing pattern in a social group of stray cats: effects on male reproductive success

Reproductive consequences of male spacing patterns have received relatively little attention in nonterritorial mammals, in particular in group-living species, where most studies have focused on the relation between social rank and reproductive success. We investigated the effects of spacing pattern on male reproductive success within a social, nonterritorial, promiscuous population of stray cats, Felis catus. Male home ranges overlapped home ranges of many females, consistent with a promiscuous mating system. Furthermore, males with the largest home ranges included the most female home ranges; they successfully reproduced with these females and had the highest reproductive success. Home range size predicted male reproductive success even when controlling for the effect of social rank. However, males also reproduced with females whose home range did not overlap their home range, suggesting that males can make quick excursions outside their home range to find new mating opportunities. We conclude that, in group-living situations, a male's ability to maintain a large home range may be one of the principal causes of variation in mating success in the stray cat.     

Age-related and individual differences of reproductive success in male and female barbary macaques (Macaca sylvanus)

Age-related and individual differences in longterm reproductive success were analyzed in two social groups of free-ranging Barbary macaques. Maternity data were obtained from continuous birth records and paternity was determined with oligonucleotide-fingerprinting. The fathers of 246 of 286 investigated individuals could be identified. They were born during a 14-year period and represented 73 and 34% of all known offspring from the females of the study groups B/F and C, respectively. Only these infants were considered when comparing male reproductive success with that of females. The necessary adjustment of the female data resulted in small deviations from the true values in one group, but substantially increased individual differences in female fertility in the second group. Subadult males, 4.5 – 6.5 yrs old, had a much lower reproductive success than adult males (7.5 – 25 yrs old) and same-aged females. Reproductive success of adult males was not significantly affected by age, while females invariably ceased reproduction during the first half of the third decade of life. Males were more likely than females to leave no offspring, unless they survived 9 – 10 yrs of age. The number of years with breeding opportunities was important for male reproductive success but less significant than that for females. Reproductive success of several males during the 14-year study period was similar to or even exceeded that possible for a female in her whole lifetime. Variance of male reproductive success significantly exceeded that of females in both study groups.     

Familiarity breeds progeny: sociality increases reproductive success in adult male ring-tailed coatis (Nasua nasua)

The ring-tailed coati (Nasua nasua) is the only coati species in which social groups contain an adult male year round, although most males live solitarily. We compared reproductive success of group living and solitary adult male coatis to determine the degree to which sociality affects reproductive success. Coati mating is highly seasonal and groups of female coatis come into oestrus during the same 1-2 week period. During the mating season, solitary adult males followed groups and fought with the group living male. This aggression was presumably to gain access to receptive females. We expected that high reproductive synchrony would make it difficult or impossible for the one group living male to monopolize and defend the group of oestrous females. However, we found that group living males sired between 67-91% of the offspring in their groups. This reproductive monopolization is much higher than other species of mammals with comparably short mating seasons. Clearly, living in a group greatly enhanced a male's reproductive success. At the same time, at least 50% of coati litters contained offspring sired by extra-group males (usually only one offspring per litter); thus, resident males could not prevent extra-group matings. The resident male's reproductive advantage may reflect female preference for a resident male strong enough to fend off competing males.     

Group size and composition influence male and female reproductive success in black howler monkeys (Alouatta pigra)

It has been argued that grouping patterns might influence the reproductive performance of individuals. Increasing group size results in greater travel costs and competition over depletable food resources, which could lead to reduced individual reproductive success. However, in groups with an increasing number of males, female reproductive success is predicted to augment because larger male groups might better protect immatures from infanticidal attacks. In contrast, male reproductive success is predicted to decrease with number of males in a group because fertilization cannot be shared between males. In this paper, we test these predictions on the Mesoamerican black howler monkey (Alouatta pigra) with data on group size and composition for 120 groups from eight populations of black howler monkeys existing in eight protected forests in Mexico and Guatemala. Male and female reproductive success were calculated as a deviation of the observed number of infants (or immatures) from the expected number of infants (or immatures), relative to the number of males and females in a group. Results indicate that both male and female reproductive success decreased with group size. Male reproductive success decreased with an increasing number of males in a group and with increasing proportion of males relative to females in a group. Decreased female reproductive success was associated with increasing number of females in a group, and female reproductive success had a tendency to increase with increasing number of males in a group. These results suggest that in black howler monkeys, living in larger groups might negatively affect the reproductive success of each member. Our findings are similar to those reported for a population of a sister species, Alouatta palliata, living in larger groups.     

Relationships of reproductive traits and body size with attainment of sexual maturity and age in Blanding's turtles (Emydoidea blandingi)

To place associations among body size, age at maturity, age, and reproductive traits of a long-lived organism in the context of current life history models based on the concept of norms of reaction, we examined data from a mark-recapture study of Blanding's turtles (Emydoidea blandingi) in southeastern Michigan during 24 of the years between 1953 and 1988. Females matured between 14 and 20 years of age. Both the smallest and largest adult females in the population were reproducing for the first time in their lives. This result suggests that a combination of differences in juvenile growth rates and ages at maturity, and not indeterminate growth, are the primary cause of variation in body size among adults. Body size variation among individuals was not related to age at sexual maturity. Females that had slower growth rates as juveniles matured later at similar mean body size compared to those with more rapid growth that matured at an earlier age. As a result, a linear model of age at sexual maturity with growth rates of primiparous females between hatching and maturity was significant and negative (R² = 0.76). Frequency of reproduction of the largest and smallest females was not significantly different. Clutch size did not vary significantly with age among either primiparous or multiparous females. Clutch sizes of primiparous females and multiparous females were not significantly different. However, older females (>55 years minimum age) reproduced more frequently than did younger females (minimum age <36 y).     

扩散或不扩散：雄性太行山猕猴繁殖成功比较研究

典型的猕猴（Macaca multta）社群为多雄多雌型,雌性留群并形成母系单元,雄性多在亚成年或成年期离开出生群,而群内成年雄性多为外部迁入个体。雄性的迁出被认为可降低近亲繁殖或提升繁殖成功。然而,诸多野外数据显示,少数本群出生雄性个体会居留于出生群一年或数年。尽管驱动雄性离群的因素较复杂（如社会关系、近亲回避、繁殖成功）,但繁殖成功的差异可能是驱动雄性离群的主导因素。为探讨居留于出生群是否影响雄性的繁殖成功,于2010年3月至2014年1月,在太行山猕猴国家级自然保护区王屋山地区,以一群野生太行山猕猴为研究对象,采用非损伤取样法并结合分子生物学方法,分析了群内出生和迁入成年雄性个体的繁殖成功（以子代数量评估）。研究发现：(1)群内51只子代个体中有36只个体可以匹配到其遗传学父亲;(2)4个迁入雄性繁衍了34个子代,仅1只群内出生雄性ZM繁衍了2只子代,但群内出生的雄性BB未匹配到子代。本研究提示,迁入雄性较本群出生雄性的繁殖成功较高,即迁移有助于繁殖成功的提升。     

Age,reproductive value,and dominance-related behaviour in vervet monkey females: cross-generational influences on social relationships and reproduction

Behavioural interactions among captive socially-living adult female vervet monkeys were analysed to determine whether (1) the motivation to maintain and increase, dominance rank declines with age as individual reproductive value declines, or (2) the cross-generational inheritance of dominance rank causes adult females to continue to involve themselves in dominance-related behaviours for the benefit of their daughters, even though their own reproductive value, is declining. The results indicate that adult females do not reduce their involvement in aggressive and dominance-related behaviours as they age. There was strong support for the hypothesis that older adult females maintain affiliative and supportive ties with their adult daughters, and that the daughters benefit from this relationship both in terms of social relationships and reproductive success. Young adult females without mothers were the recipients of more aggression, were less likely to be defeded when they were attacked and were less likely to challenge the rank of others compared to females with mothers living in the group. Young adult females with mothers produced significantly more surviving offspring and had a lower level of infant mortality than comparable adult females without mothers.     

The effects of dominance rank and group size on female lifetime reproductive success in wild long-tailed macaques,Macaca fascicularis

Demographic changes were recorded throughout a 12-year period for three social groups ofMacaca fascicularis in a natural population at Ketambe (Sumatra, Indonesia). We examined the prediction that females' lifetime reproductive success depended on dominance rank and group size. Average birth rate was 0.53 (184 infants born during 349 female years). For mature females (aged 8–20 yr) birth rate reflected physical condition, being higher in years with high food availability and lower in the year following the production of a surviving infant. High-ranking females were significantly more likely than low-ranking ones to give birth again when they did have a surviving offspring born the year before (0.50 vs 0.26), especially in years with relatively low food availability (0.37 vs 0.10). Controlled comparisons of groups at different sizes indicate a decline in birth rate with rroup size only once a group has exceeded a certain size. The dominance effect on birth rate tended to be strongest in large groups. Survival of infants was rank-dependent, but the survival of juveniles was not. There was a trend for offspring survival to be lower in large groups than in mid-sized or small groups. However, rank and group size interacted, in that rank effects on offspring survival were strongest in large groups. High-ranking females were less likely to die themselves during their top-reproductive years, and thus on average had longer reproductive careers. We estimated female lifetime reproductive success based on calculated age-specific birth rates and survival rates. The effects of rank and group size (contest and scramble) on birth rate, offspring survival, age of first reproduction for daughters, and length of reproductive career, while not each consistently statistically significant, added up to substantial effects on estimated lifetime reproductive success. The group size effects explain why large groups tend to split permanently. Since females are philopatric in this species, and daughters achieve dominance rank positions similar to their mother, a close correlation is suggested between the lifetime reproductive success of mothers and daughters. For sons, too, maternal dominance affected their reproductive success: high-born males were more likely to become top-dominant (in another group). These data support the idea that natural selection has favored the evolution of a nepotistic rank system in this species, even if the annual benefits of dominance are small.     

Male reproductive success in plateau pikas (Ochotona curzoniae): A microsatellite analysis

In many social species, competition and cooperation between group members may lead to a large variance in reproductive success among individuals, especially for adult male. From April to August 2002, we studied the adult male reproductive success of plateau pikas in Haibei Alpine Meadow Ecosystem Research Station, Chinese Academy of Science, using microsatellite analysis of paternity, mark-recapture and behavioral observations. Our result indicated that the reproductive success of adult males had a large variance. Its average was 6 individuals and range was from 0 to 13 individuals. One-third of males sired 63.22% offspring. There was a hierarchy system in polygynandrous families. Although the reproductive success of dominant males was higher than that of subordinate males, subordinate males still play an important role in the reproductive success of the population.     

The effect of male tenure and female mate choice on paternity in free-ranging Japanese macaques

In this study, the paternity of all the infants born in 2002 and 2003 in a free-ranging Japanese macaque (Macaca fuscata) group at Arashiyama in Kyoto, Japan, was analyzed in relation to males' age, dominance rank, and tenure and females' mate choice. The fathers of 20 out of 23 infants were determined by DNA analyses. Central adult (high-ranking) males sired two infants, whereas peripheral adult (low-ranking) males sired 14 infants. Young males sired only one infant. Among adult males, tenure was the most dominant factor that negatively affected male reproductive success. The mating behavior of females who gave birth was also analyzed. The number of male copulations in the peri-fertilization period was positively correlated with the number of infants that they sired. Females copulated with central males with a long tenure only when fertilization was unlikely or impossible. The females probably avoided insemination by males with a long tenure and selected males with a shorter tenure as their mating partners during the ovulation period.