Intergroup Relationships in Western Black-and-White Colobus, <Emphasis Type="Italic">Colobus polykomos polykomos</Emphasis>

We investigated patterns of intergroup relationships in western black-and-white colobus, Colobus polykomos, in Taï National Park, Côte d'Ivoire, between 1993 and 1999. They live in one-male multifemale units, and demonstrate male dispersal and occasional dispersal by females. Solitary males and all-male bands are absent or very rare. Our aim was to investigate the function of female and male aggression during intergroup interactions. The species is particularly interesting because, in contrast to predictions from socioecological models, female aggression occurs during intergroup interactions in combination with female dispersal. Home ranges of neighboring groups overlapped considerably and groups lacked an area of exclusive access. Intergroup interactions occurred once every 6.6 observation days. Encounters were either peaceful (12%), or involved displays and threats (25%) or chases and fights (63%). Females interacted in 74% and males in 98% of aggressive intergroup encounters. We found little to no indication that male and female aggression correlated with the presence of food, importance of a location, or presence of infants or receptive females. However, females were more often aggressive during the months when the group depended strongly on seeds from Pentaclethra macrophylla. We also observed forays by males to other groups. Forays occurred on average once every 20 observation days. In 75% of the forays, the intruding male chased members of the target group. In 25% of the forays 1–3 females joined their male but females never attacked the target group. Our main study group was the target of such forays significantly more often when young infants were present in the group than when not. We conclude that female aggression between groups was related to food procurement and that male forays might be related to infanticide.     

Stress and the costs of extra-territorial movement in a social carnivore

Costs associated with extra-territorial movement are believed to have favoured the evolution of delayed dispersal and sociality across a range of social vertebrates, but remain surprisingly poorly understood. Here we reveal a novel mechanism that may contribute substantially to the costs of extra-territorial movement: physiological stress. We show that subordinate male meerkats, Suricata suricatta, exhibit markedly elevated faecal glucocorticoid metabolite levels (a non-invasive measure of hypothalamic–pituitary–adrenal axis activity) while conducting extra-territorial prospecting forays. While brief increases in glucocorticoid levels are unlikely to be costly, chronic elevations, arising from prolonged and/or frequent forays, are expected to compromise fitness through their diverse negative effects on health. Our findings strongly suggest that prolonged extra-territorial movements do result in chronic stress, as the high glucocorticoid levels of prospectors do not diminish on longer forays and are no lower among males with greater prospecting experience. A generalized ‘stress’ of extra-territorial movement may therefore have strengthened selection for delayed dispersal and sociality in this and other species, and favoured the conduct of brief forays over extended periods of floating. Our findings have implications too for understanding the rank-related distribution of physiological stress in animal societies, as extra-territorial movements are often conducted solely by subordinates.     

Early growth,dominance acquisition and lifetime reproductive success in male and female cooperative meerkats

In polygynous species, variance in reproductive success is higher in males than females. There is consequently stronger selection for competitive traits in males and early growth can have a greater influence on later fitness in males than in females. As yet, little is known about sex differences in the effect of early growth on subsequent breeding success in species where variance in reproductive success is higher in females than males, and competitive traits are under stronger selection in females. Greater variance in reproductive success has been documented in several singular cooperative breeders. Here, we investigated consequences of early growth for later reproductive success in wild meerkats. We found that, despite the absence of dimorphism, females who exhibited faster growth until nutritional independence were more likely to become dominant, whereas early growth did not affect dominance acquisition in males. Among those individuals who attained dominance, there was no further influence of early growth on dominance tenure or lifetime reproductive success in males or females. These findings suggest that early growth effects on competitive abilities and fitness may reflect the intensity of intrasexual competition even in sexually monomorphic species.     

Determinants of reproductive success in dominant female meerkats

1. In cooperative societies with high reproductive skew, selection on females is likely to operate principally through variation in the probability of acquiring dominant status and variation in reproductive success while dominant. Despite this, few studies of cooperative societies have investigated the factors that influence which females become dominant, and/or their reproductive output while in the dominant position. 2. Here we use long-term data from a wild meerkats population to describe variation in the breeding success of dominant female meerkats Suricata suricatta and investigate its causes. 3. Female meerkats compete intensely for breeding positions, and the probability of acquiring the breeding role depends upon a female's age in relation to competitors and her weight, both at the time of dominance acquisition and early in life. 4. Once dominant, individual differences in breeding success depend principally on the duration of dominance tenure. Females remain for longer in the dominant position if they are heavier than their competitors at the start of dominance, and if the number of adult female competitors at the start is low. 5. Female breeding success is also affected by variation in fecundity and pup survival, both of which increase with group size. After controlling for these effects, female body weight has a positive influence on breeding rate and litter size, while the number of adult female competitors reduces litter survival. 6. These findings suggest that selection for body weight and competitive ability will be high in female meerkats, which may moderate their investment in cooperative activities. We suggest that similar consequences of competition may occur among females in other cooperative societies where the benefits of attaining dominance status are high.     

Sex differences in responsiveness to begging in a cooperative mammal

In species where young are provisioned by both parents, males commonly contribute less to parental care than females, and are less responsive to variation in begging rates. Similar differences in the care of young occur among adults in cooperative breeders, but fewer studies have investigated whether these are associated with differences in responsiveness. Here, we present results from a playback experiment investigating responsiveness to begging in the meerkat (Suricata suricatta), a cooperatively breeding mammal. Although increased begging rate raised the feeding rate of adults of both sexes, there was no consistent tendency for females to be more responsive than males. However, when we examined changes in the proportion of food items found that were fed to pups (generosity), we found that females were more responsive than males to increased begging rate. These results can be explained in terms of sex differences in dispersal: in meerkats, females are philopatric and receive considerable benefits from investing in young, both directly, by increasing group size, and indirectly, by recruiting helpers if they inherit the breeding position. In addition, they emphasize that generosity provides a more sensitive measure of responsiveness to begging than feeding rate, as it accounts for variation in foraging success.     

Male-biased Adult Sex Ratios and their Significance for Cooperative Breeding in Dhole,Cuon alpinus,Packs

Two free-ranging packs of dholes (Asiatic wild dog, Cuon alpinus) were monitored for a period of 6 yr (Sep. 1990-Sep. 1996) in the Mudumalai sanctuary, southern India. Demographic data on age structure, litter-size, sex ratio and age and sex specific dispersal were collected. Behavioural data on social interactions and reproductive behaviour among pack members were obtained to determine the frequencies of dominant and subordinate behaviours shown by male and female pack members and a measure of each male's reproductive access to females. Behaviours displayed by pack members at dens were recorded to determine whether any age- or sex-specific role specialization existed during pup care. Tenures for dominant males and females within the pack were calculated to ascertain the rate of breeding vacancies occurring within packs. Approximate levels of genetic relatedness within packs were determined by studying pedigrees. In most years one study pack had a male-biased adult sex ratio. This was caused by almost two-fold higher dispersal of adult females over adult males. A considerable variance existed in the percentage of sub-adults dispersing from the two packs. Differences existed in the frequencies of dominant and subordinate behaviours shown by males. For males, dominance ranks and ranks based on submissive behaviours were not correlated with frequencies of reproductive behaviours. Subordinate males also displayed reproductive behaviours. In packs, dominant males had lower tenures than dominant females indicating that among males breeding vacancies arose more quickly. The litter size was found to be negatively correlated with the age of the breeding female. There were no significant differences across individuals of varying age or sex classes in the display of pup care behaviours. Significant differences did exist among individual adults. Genetic relatedness among packs tended to vary temporally as a consequence of possible mating by subordinate animals and immigration of new males into the pack. In conclusion, it appears that males delay dispersal and cooperate within their natal packs because of the variety of reproductive strategies they could pursue within. A combination of ecological constraints and the difficulties of achieving breeding status within non-natal packs may make early dispersal and independent breeding less beneficial.     

Hormonal correlates of dominance in meerkats (Suricata suricatta)

In cooperatively breeding meerkats (Suricata suricatta), individuals typically live in extended family groups in which the dominant male and female are the primary reproductives, while their offspring delay dispersal, seldom breed, and contribute to the care of subsequent litters. Here we investigate hormonal differences between dominants and subordinates by comparing plasma levels of luteinizing hormone (LH), estradiol and cortisol in females, and testosterone and cortisol in males, while controlling for potential confounding factors. In both sexes, hormone levels are correlated with age. In females, levels of sex hormone also vary with body weight and access to unrelated breeding partners in the same group: subordinates in groups containing unrelated males have higher levels of LH and estradiol than those in groups containing related males only. When these effects are controlled, there are no rank-related differences in circulating levels of LH among females or testosterone among males. However, dominant females show higher levels of circulating estradiol than subordinates. Dominant males and females also have significantly higher cortisol levels than subordinates. Hence, we found no evidence that the lower levels of plasma estradiol in subordinate females were associated with high levels of glucocorticoids. These results indicate that future studies need to control for the potentially confounding effects of age, body weight, and access to unrelated breeding partners before concluding that there are fundamental physiological differences between dominant and subordinate group members.     

Dispersal influences genetic and acoustic spatial structure for both males and females in a tropical songbird

Animals exhibit diverse dispersal strategies, including sex‐biased dispersal, a phenomenon common in vertebrates. Dispersal influences the genetic structure of populations as well as geographic variation in phenotypic traits. Patterns of spatial genetic structure and geographic variation may vary between the sexes whenever males and females exhibit different dispersal behaviors. Here, we examine dispersal, spatial genetic structure, and spatial acoustic structure in Rufous‐and‐white Wrens, a year‐round resident tropical bird. Both sexes sing in this species, allowing us to compare acoustic variation between males and females and examine the relationship between dispersal and song sharing for both sexes. Using a long‐term dataset collected over an 11‐year period, we used banding data and molecular genetic analyses to quantify natal and breeding dispersal distance in Rufous‐and‐white Wrens. We quantified song sharing and examined whether sharing varied with dispersal distance, for both males and females. Observational data and molecular genetic analyses indicate that dispersal is female‐biased. Females dispersed farther from natal territories than males, and more often between breeding territories than males. Furthermore, females showed no significant spatial genetic structure, consistent with expectations, whereas males showed significant spatial genetic structure. Overall, natal dispersal appears to have more influence than breeding dispersal on spatial genetic structure and spatial acoustic structure, given that the majority of breeding dispersal events resulted in individuals moving only short distances. Song sharing between pairs of same‐sex animals decreases with the distance between their territories for both males and females, although males exhibited significantly greater song sharing than females. Lastly, we measured the relationship between natal dispersal distance and song sharing. We found that sons shared fewer songs with their fathers the farther they dispersed from their natal territories, but that song sharing between daughters and mothers was not significantly correlated with natal dispersal distance. Our results reveal cultural differences between the sexes, suggesting a relationship between culture and sex‐biased dispersal.     

Female "dispersal" in hamadryas baboons: transfer among social units in a multilevel society

Unlike most cercopithecines, hamadryas baboons (Papio hamadryas hamadryas) are characterized by female-biased dispersal. To clarify this pattern within the context of their hierarchical social system (comprising one-male units, clans, bands, and troops), we report here 7 years of data on female transfers among social units in wild hamadryas baboons in Ethiopia. Female tenure in one-male units (OMUs) ranged from 1 to 2,556 days (N = 208) and survival analysis revealed a median tenure length of 1,217 days (40 months). Changes in OMU membership consisted almost exclusively of takeovers by males, not voluntary transfer. Of 130 takeovers, 67% occurred within the band and 33% across bands, and, of the 22 takeovers for which we have clan membership data, 77% occurred within, not between, clans. These results reinforce the notion that hamadryas female dispersal is not analogous to sex-biased dispersal in other taxa, because (1) at least in Ethiopian populations, females do not disperse voluntarily but are transferred, often forcibly, by males; (2) only dispersal between bands will promote gene flow, whereas females are most often rearranged within bands; (3) hamadryas females undergo social dispersal but not usually locational dispersal; and (4) while male hamadryas are far more philopatric than females, they have been observed to disperse. It thus appears that the ancestral baboon pattern of female philopatry and male dispersal has evolved into a system in which neither sex is motivated to disperse, but females are forcibly transferred by males, leading to female-mediated gene flow, and males more rarely disperse to find females.     

Breeding and juvenile survival among slender-tailed meerkats (Suricatu suricatta) in the south-western Kalahari: ecological and social influences

Reproductive success of co-operatively-breeding slender-tailed meerkats ( Suricata suricatta ) in the Kalahari was monitored over four breeding seasons and 26 band years, concentrating on three focal bands. Breeding was strongly seasonal, with peak numbers of births coinciding with maximum rainfall between January and March. Breeding seasons were extended, and more litters were produced, during the years with above average rainfall. For dominant females, the mean annual rate of litter production was 1.9 ± 0.8 litters per year and short interbirth intervals (mean = 90 %pL 18 days) indicated that females came into oestrus within three weeks of giving birth. Births were synchronous within but not between bands. Major known causes of kitten mortality were cold weather and predation, and losses mainly occurred between three and five weeks of age. Sixty-seven percent of juveniles survived between emergence from the den at three or four weeks old and attainment of effective foraging independence at 12 weeks. There were no significant effects of birth date within the season upon litter size or survival. Several incidents of apparent infanticide were recorded. These meerkats were highly co-operative and adult band members assiduously guarded and provisioned the kittens. Nevertheless, regression analysis showed that neither rainfall (an index of prey availability) nor band size variables accounted for much variance in juvenile survival to 12 weeks, which appeared to be heavily influenced by chance events such as flash floods. In contrast, rainfall between January and March had a significantly positive effect on the total number of juveniles produced during the breeding year.     

Dominance and Mating in a Communal Polygynandrous Bird: Cooperation or Indifference towards Mating Competitors?

The effect of dominance on the mating behaviour of both females and males was investigated for the communally breeding pukeko (Porphyrio porphyrio melanotus). Pukeko are polygynandrous with most groups consisting of three to 7 breeding males and one or two breeding females that lay eggs in a single nest. Linear dominance hierarchies are well established within breeding groups. Alpha females were involved in more copulations than beta females, but this difference did not appear to be a result of dominance status. There was also no evidence of egg tossing or egg destruction by breeding females. There was no positive correspondence between dominance rank of males and their frequency of sexual activity or copulations during the prelaying or laying periods. Dominants did interrupt matings by subordinates, but both qualitative and quantitative analyses of behaviour found no evidence of mate guarding. Furthermore dominants were no more likely to interrupt a copulation involving a subordinate male than they were to passively observe it or not react at all. This led us to suggest that males tended to behave indifferently toward mating competitors. As a result, multiple paternity is predicted to be high in pukeko groups. We also question the standard interpretation that males are behaving cooperatively in communal groups rather than coactively.     

Mate inspection patterns in the female beaugregory damselfish (<Emphasis Type="Italic">Stegastes leucostictus</Emphasis>)

We observed female beaugregory damselfish (Stegastes leucostictus) as they interacted with males to see whether their movements fit the predictions of different mate-search models. We established high-quality and low-quality groups with low variance in breeding site quality and a medium-quality, high-variance breeding site group and compared focal observations of female nonforaging forays in each group. Only 8% of 137 forays monitored resulted in spawning events. Eighty-nine percent of the forays were less than 240 s and were within 7 m of the focal female’s territory. Average foray times and straight-line foray distances did not differ for females traveling between males in each group. Females entered high-quality breeding sites at a higher rate than low-quality sites and round-trip distances were also greatest in high-quality groups, indicating that females were following a more convoluted path. We interpret these results to mean that (a) female beaugregories separate the tasks of mate assessment and mate choice by conducting information-gathering forays between mating events, (b) forays are energetically costly and therefore usually short in duration and overall distance traveled, (c) information collection takes approximately the same time for high-quality and low-quality mates, but (d) females will alter their foray patterns in high-quality areas to collect additional information. These data are also consistent with the hypothesis that females optimize their search by employing a tiered process of assessment during information-gathering forays, first using an adjustable threshold to accept or reject each male based on his courtship and subsequently a fixed threshold to assess the quality of each breeding site.     

Sex-biased dispersal: a result of a sex difference in breeding site availability

Sex-biased dispersal is often explained by assuming that the resource-defending sex pays greater costs of moving from a familiar area. We hypothesize that sex-biased dispersal may also be caused by a sex bias in breeding site availability. In avian resource-defense mating systems, site availability is often more constrained for females: males can choose from all vacant sites, whereas females are restricted to sites defended by males. Using data on breeding dispersal of a migratory passerine, we show that average number of available breeding options and availability of the previous year's territory was greater for males than females. The female bias in site unavailability may explain the female bias in probability of breeding dispersal because there was no sex bias in dispersal among birds with their previous year's territory available. We suggest that sex biases in the availability of breeding options may be an important factor contributing to observed variation in sex-biased dispersal patterns.     

Sex-biased dispersal in western lowland gorillas (Gorilla gorilla gorilla)

We explored two hypotheses related to potential differences between sexes in dispersal behaviour in western lowland gorillas (Gorilla gorilla gorilla). Direct observations suggest that immature females have more opportunities to move between breeding groups than immature males. The distribution of kin dyadic relationships within and between groups does not, however, support this hypothesis. At larger geographical scales, dispersal is likely to be easier for males than females because of the solitary phase most blackbacks experience before founding their own breeding group. However, previous work indicates that males settle preferentially close to male kin. By specifically tracing female and male lineages with mitochondrial and Y-chromosomal genetic markers, we found that male gorillas in the 6000 km2 area we surveyed form a single population whereas females are restricted to the individual sites we sampled and do not freely move around this area. These differences are more correctly described as differences in dispersal distances, rather than differences in dispersal rates between sexes (both sexes emigrate from their natal group in this species). Differences in resource competition and dispersal costs between female and male gorillas are compatible with the observed pattern, but more work is needed to understand if these ultimate causes are responsible for sex-biased dispersal distances in western lowland gorillas.     

The disperser dilemma in cooperatively breeding birds

In most cooperatively breeding birds, individuals do not breed with their natal group members. In order to breed, they have either to disperse into another group or wait for an opposite-sex individual to join their group. In most of these species, females disperse more than males. We develop a dynamic game-theoretic model to account for this asymmetry. When males are physically larger/heavier than females, this allows them to effectively welcome female immigrants into their natal group and overcome the local females' opposition more than vice versa. The model further assumes that the dispersal decision is not confined to a restricted time window, but is rather based on acquired information and responsive to opportunities. The model predicts that (i) females disperse more than males, and (ii) females are willing to tolerate more risks in dispersal than do males. The latter prediction is supported inter alia by the fact that in many cooperatively breeding birds, females disperse at a younger age, and further away from their natal group as compared to dispersing males.     

Correlates and Consequences of Dominance in a Social Rodent

In harem‐polygynous societies, body condition is often correlated with dominance rank. However, the consequences of dominance are less clear. High‐ranking males do not inevitably have the highest reproductive success, especially in systems where females mate with multiple males. In such societies, we expect male reproductive success to be more highly skewed than female reproductive success, but reproductive skew in females can still arise from rankings established within matrilineal societies. Dominance can also impact life‐history decisions by influencing dispersal patterns in yearlings. To better understand the function of dominance in harem‐polygynous societies, we studied the causes and consequences of dominance in yellow‐bellied marmots (Marmota flaviventris), a social rodent with skewed male reproductive success and female reproductive suppression. Specifically, we examined body condition as a predictor and the probability of breeding, number of offspring, and dispersal as outcomes of dominance. Additionally, we looked at variation in dominance between males and females and adults and yearlings, because marmots can engage in distinct interactions depending on the type of individuals involved. We found that marmots in better body condition have higher dominance rank than those in poorer condition. In addition, adults are dominant over yearlings. Within yearlings, dominance does not influence dispersal, but those in better body condition are less likely to disperse. Within all adults, individuals in better condition produce more offspring per year. Within adult males, more dominant males have greater reproductive success. Despite previous evidence of reproductive suppression in females, we found no effects of dominance rank on female reproductive success in the current study. The function of dominance in female marmots remains enigmatic.     

Determinants of breeding dispersal in the sand lizard, Lacerta agilis, (Reptilia, Squamata)

We studied determinants of breeding dispersal (the distance that an animal shifts its mean home range co-ordinate between reproductive events) in an individually marked population of sand lizards (Lacerta agilis) in south west Sweden during 1987–1991. Female breeding dispersal was not determined by age, size, body condition, or number of partners, and females and males that dispersed further did not experience a higher mortality. However, females with a low reproductive output dispersed further than females that reproduced more successfully, and males that lost in bodily condition dispersed further than males that better maintained body condition. We also looked for relationships between age-differences and band-sharing similarity (DNA fingerprinting = DFP) in three categories of lizards — all females, all males, and males and females — to establish whether males would be likely to mate with close kin. Age-difference was strongly correlated with band-sharing in only one category, males and females. When males were older than females this relationship was not significant. However, when females were older than males, age-difference was strongly correlated with band-sharing. Furthermore, females that were old enough to be the mothers of the courting males shared significantly more bands with these males than did the younger females, including the females of the same age as the males. Although parental-specific DFP bands necessary for establishing paternity among our adult lizards were inaccessible to us, we suggest that our circumstantial evidence strongly suggests that some males mate with their mothers. Males that were more closely related with their neighbours also moved further when we controlled statistically for age and mating success. We suggest that by mating with many partners males not only increase their mating success, but also increase the probability of mating with females with ‘good genes’: mean heterozygosity of parents (as revealed by micro-satellites) were strongly correlated with offspring survival.     

Dispersal, philopatry, and genetic relatedness in a social carnivore: comparing males and females

A balance must be maintained between the proportion of individuals dispersing and the proportion remaining philopatric such that inbreeding and resource competition are minimized. Yet the relative importance of dispersal and philopatric behaviour is uncertain, especially for species with complex social systems. We examine the influence of dispersal on genetic relationships of a white-nosed coati ( Nasua narica : Procyonidae) population from Panama. Field studies of the coati indicate a social system in which all females are highly philopatric and live in bands while all adult males become solitary at maturity, but do not disperse from the home range of their natal band. Based on analyses of multilocus DNA fingerprints, we confirm that female philopatry is the rule, long-distance dispersal is rare, and that relatedness between most bands is low. However, some new bands result from fission events and these bands retain relatively high relatedness to one another for several years. Adult males inhabiting the home range of a band are closely related to band members. In contrast, males and band members whose ranges do not overlap are unrelated or only slightly related. Adult males are also more closely related to other males whose home ranges they overlap extensively than to males whose home ranges they overlap only slightly. These results indicate that males initially disperse from their natal bands to reduce resource competition and not to avoid inbreeding. Inbreeding avoidance, if it occurs, results from more extensive range movements by males during the mating season.     

Competition, breeding success and ageing rates in female meerkats

Competition between females is particularly intense in cooperatively breeding mammals, where one female monopolises reproduction in each group. Chronic competition often affects stress and may therefore have long-term consequences for fitness, but no studies have yet investigated whether intrasexual competition has effects of this kind and, in particular, whether it affects rates of reproductive senescence. Here, we use long-term data from a wild population of meerkats to test whether reproductive success and senescence in dominant females are affected by the degree of intrasexual competition experienced prior to dominance acquisition. Females that experienced greater competition had lower breeding success and higher rates of reproductive senescence. Furthermore, females that were evicted from the group more frequently as subordinates had lower breeding success when dominant. We conclude that the intense intrasexual competition between females in cooperatively breeding groups may carry fitness costs over a longer period than is usually recognised.     

Male immigration triggers increased growth in subordinate female meerkats

There is increasing evidence that some vertebrates can adjust their growth rate in relation to changes in the social context that affect their probability of breeding. Here, we show that, in meerkats (Suricata suricatta), which are singular cooperative breeders, subordinate females increase in body mass after their father is replaced as the dominant male in their natal group by an immigrant male, giving them regular access to an unfamiliar and unrelated mating partner, while their brothers showed no similar increase nor did subordinate females living in other stable groups (where male immigration did not occur did) in this time period. Moreover, subordinate females showed a greater increase in growth rate when their father was succeeded by an unfamiliar immigrant male than when he was replaced by a familiar male who was already resident. These results suggest that female meerkats can adjust their rate of growth to changes in the kinship composition of their groups that provide them with increased access to unrelated breeding partners, which may occur in other mammals as well when breeding opportunities change.