Environmental conditions affect spatial genetic structures and dispersal patterns in a solitary rodent

The study of the spatial distribution of relatives in a population under contrasted environmental conditions provides critical insights into the flexibility of dispersal behaviour and the role of environmental conditions in shaping population relatedness and social structure. Yet few studies have evaluated the effects of fluctuating environmental conditions on relatedness structure of solitary species in the wild. The aim of this study was to determine the impact of interannual variations in environmental conditions on the spatial distribution of relatives [spatial genetic structure (SGS)] and dispersal patterns of a wild population of eastern chipmunks (Tamias striatus), a solitary rodent of North America. Eastern chipmunks depend on the seed of masting trees for reproduction and survival. Here, we combined the analysis of the SGS of adults with direct estimates of juvenile dispersal distance during six contrasted years with different dispersal seasons, population sizes and seed production. We found that environmental conditions influences the dispersal distances of juveniles and that male juveniles dispersed farther than females. The extent of the SGS of adult females varied between years and matched the variation in environmental conditions. In contrast, the SGS of males did not vary between years. We also found a difference in SGS between males and females that was consistent with male‐biased dispersal. This study suggests that both the dispersal behaviour and the relatedness structure in a population of a solitary species can be relatively labile and change according to environmental conditions.     

Fine-scale genetic structure and dispersal in cooperatively breeding apostlebirds

In cooperatively breeding species, restricted dispersal of offspring leads to clustering of closely related individuals, increasing the potential both for indirect genetic benefits and inbreeding costs. In apostlebirds (Struthidea cinerea), philopatry by both sexes results in the formation of large (up to 17 birds), predominantly sedentary breeding groups that remain stable throughout the year. We examined patterns of relatedness and fine-scale genetic structure within a population of apostlebirds using six polymorphic microsatellite loci. We found evidence of fine-scale genetic structure within the study population that is consistent with behavioural observations of short-distance dispersal, natal philopatry by both sexes and restricted movement of breeding groups between seasons. Global F(ST) values among breeding groups were significantly positive, and the average level of pairwise relatedness was significantly higher for individuals within groups than between groups. For individuals from different breeding groups, geographical distance was negatively correlated with pairwise relatedness and positively correlated with pairwise F(ST). However, when each sex was examined separately, this pattern was significant only among males, suggesting that females may disperse over longer distances. We discuss the potential for kin selection to influence the evolution and maintenance of cooperative breeding in apostlebirds. Our results demonstrate that spatial genetic structural analysis offers a useful alternative to field observations in examining dispersal patterns of cooperative breeders.     

Gene dispersal and outbreeding in a philopatric mammal

Extensive mark-recapture data from banner-tailed kangaroo rats, Dipodomys spectabilis, have shown that both males and females are highly philopatric and suggest the possibility of close inbreeding. However, indirect analyses based on genetic structure appear to contradict direct observations, suggesting longer dispersal distances. Using microsatellite genotypes from most members of a banner-tailed kangaroo rat population during five successive breeding seasons, we ask how relatedness is influenced by dispersal and how it in turn influences mating patterns. The data confirm that, because of philopatry, neighbours are often close relatives. However, patterns of parentage also show that the average distance between mates is large relative to natal dispersal distances and larger than the average distance between nearest opposite-sexed neighbours. Females' mates were often not their nearest male neighbour and many were less related than the nearest male neighbour. We detected multiple paternity in some females' litters; both sexes produce offspring with multiple mates within and between breeding seasons. At the population level, heterozygosities were high and estimates of F were low, indicating that levels of inbreeding were low. Using individual inbreeding coefficients of all juveniles to estimate their parents' relatedness, we found that parental relatedness was significantly lower than relatedness between nearest opposite-sexed adult neighbours. Thus in philopatric populations, long breeding forays can cause genes to move further than individuals disperse, and polyandry may serve to reduce relatedness between mates.     

Spatial association of genetically similar Atlantic salmon juveniles and sex bias in spatial patterns in a river

Atlantic salmon ( Salmo salar ) juveniles were electrofished along a 300-m stretch of a river to test for possible associations between genetic similarity and geographical distance between individuals. Multilocus DNA fingerprinting showed that genetically similar juveniles (1–4 years old) were found closer together in the river than less related individuals. However, the association between genetic similarity and geographical distance, although significant, was not strong. This may indicate that factors other than genetic relatedness influence the positioning in the river. A sex bias in the relationship between genetic similarity and geographical distance was caused by a difference between sexually mature and immature males. The study shows that sampling of salmon juveniles should be spread over a wide stretch of the river in order to avoid sampling relatives. Moreover, by including several year classes, the overall degree of genetic similarity is effectively reduced compared with sampling individuals of similar age.     

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.     

Evidence for size and sex-specific dispersal in a cooperatively breeding cichlid fish

African Great Lake cichlid populations are divided into thousands of genetic subpopulations. The low gene flow between these subpopulations is thought to result from high degrees of natal philopatry, heavy predation pressure, and a patchy distribution of preferred habitats. While predation pressure and habitat distribution are fairly straightforward to assess, data on dispersal distances and rates are scarce. In fishes, direct observations of dispersal events are unlikely, but dispersal can be studied using molecular markers. Using seven microsatellite loci, we examined dispersal in the cooperatively breeding cichlid fish, Neolamprologus pulcher. As this species is found in well-defined groups clustered into subpopulations, we could assess dispersal on a narrow (within subpopulation) and broad (between subpopulation) scale. While fish were generally more related to others in their own subpopulation than they were to fish from other subpopulations, large males diverged from this pattern. Large males were more related to other large males from different subpopulations than they were to large males from their own subpopulation, suggesting more frequent dispersal by large males. Across subpopulations, relatedness between large males was higher than the relatedness among large females; this pattern was not detected in small males and small females. Within a subpopulation, individuals appeared to be preferentially moving away from relatives, and movement was unrestricted by the physical distance between groups. Our results highlight the importance of examining multiple spatial scales when studying individual dispersal biases.     

Inferring colonization history from analyses of spatial genetic structure within populations of Pinus strobus and Quercus rubra

Many factors interact to determine genetic structure within populations including adult density, the mating system, colonization history, natural selection, and the mechanism and spatial patterns of gene dispersal. We examined spatial genetic structure within colonizing populations of Quercus rubra seedlings and Pinus strobus juveniles and adults in an aspen-white pine forest in northern Michigan, USA. A 20-year spatially explicit demographic study of the forest enables us to interpret the results in light of recent colonization of the site for both species. We assayed 217 Q. rubra seedlings and 171 P. strobus individuals at 11 polymorphic loci using nine allozyme systems. Plant genotypes and locations were used in an analysis of spatial genetic structure. Q. rubra and P. strobus showed similar observed levels of heterozygosity, but Q. rubra seedlings have less heterozygosity than expected. Q. rubra seedlings show spatial genetic clumping of individuals on a scale to 25 m and levels of genetic relatedness expected from the clumped dispersion of half-siblings. In contrast, P. strobus has low levels of genetic relatedness at the smallest distance class and positive spatial genetic structure at scales < 10 m within the plot. The low density of adult Q. rubra outside the study plot and limited, spatially clumped rodent dispersal of acorns is likely responsible for the observed pattern of spatial genetic structure and the observed heterozygote deficit (i.e. a Wahlund effect). We attribute weaker patterns observed in P. strobus to the longer dispersal distance of seeds and the historical overlap of seed shadows from adults outside of the plot coupled with the overlap of seed shadows from younger, more recently established reproductive adults. The study demonstrates the utility of long-term demographic data in interpreting mechanisms responsible for generating contemporary patterns of genetic structure within populations.     

不同生活史阶段长爪沙鼠尿标记行为的遗传和空间距离效应

气味标记普遍存在于哺乳动物类群中,它能向同类传达领地占有、社会等级及繁殖状态等多种信息。2014年4月至10月,我们在内蒙古锡林郭勒正蓝旗对长爪沙鼠 (Meriones unguiculatus) 野外种群进行了标志重捕,采用中立场的方式测定了不同社群间同性别长爪沙鼠的尿标记行为。通过建立洞群样地坐标后用勾股弦方法计算群间距离,并用微卫星DNA分子标记估算个体间的亲缘系数,以探讨野生动物标记行为是否受个体间空间和遗传距离的影响。广义线性混合效应分析显示,雄鼠间的尿标记强度在繁殖期和贮食期受空间距离或遗传距离的影响均不显著。繁殖期雌鼠间的尿标记强度受空间或遗传距离的影响亦不明显,但在贮食期,遗传距离对雌鼠间标记的强度有显著影响,且遗传距离与空间距离对标记强度有明显的交互作用。本研究结果暗示亲缘关系和空间距离对长爪沙鼠的尿标记行为具有一定影响,这与长爪沙鼠的繁殖或贮食行为相适应。     

Estimates of relatedness and inbreeding in goose strains from DNA fingerprints

The purpose of this study was to determine if DNA fingerprints (DFPs) could be used to estimate relatedness and inbreeding of strains of geese and to compare three methods of calculating relatedness indices. Strains included a control and selected strain from each of the Chinese and Synthetic (Chinese, Hungarian and Pilgrim) breeds. DFP patterns for each strain were based on individual DNA samples from six females, or on pooled DNA from 15 females different from those used for individual samples. Three relatedness indices were used, namely, genetic distance, modified Rogers distance and band sharing. All relatedness indices showed a closer relationship of strains within than between breeds. Correlation coefficients among relatedness indices were higher based on pooled DNA (r ≥|0·97|) than those based on individual DNA (r ≥|0·741). Inbreeding estimates were higher for selected compared with control strains. It appears that the use of DFPs to estimate relatedness, regardless of index used, and inbreeding can be valuable for studying geese where there is a limited breeding history.     

Fine-scale study finds limited evidence of kin clustering in a gray seal colony

Microsatellites were used to investigate fine‐scale spatial and temporal genetic structure of a gray seal breeding colony, using samples collected throughout the colony (1997) and more extensive local sampling (2000–2002). Previous behavior of breeding seals on North Rona, including philopatry and high breeding site fidelity, suggested female kin may cluster together. However, low F_ST scores indicated no genetic differentiation between the major breeding aggregations in different regions of the colony. Nevertheless, the pairwise relatedness of mothers within regions was significantly higher than the relatedness of mothers between regions. Evidence of kin clustering occurred within one region in the colony. Within this region, mothers' pairwise relatedness decreased significantly with increasing distance between pupping sites. Pairwise relatedness of neighboring females within this region was also higher than expected. Conversely, in the other regions mothers that were considered likely to have social interactions, based on their spatial and temporal proximity, were not more related to each other than random. Therefore, sustained philopatry appears sufficient to produce differences in relatedness at a within‐colony scale (>500 m), but not at finer scales. This suggests that long‐term associations of mothers on North Rona detected previously are unlikely to be formed solely by close kin.     

Group size,composition, and reproductive success in wild common marmosets (Callithrix jacchus)

Data from published sources about size and composition of wild common marmoset groups (Callithrix jacchus) were analyzed to see if the number of juveniles in a group is closely related to the number of other group members. Mean group size was 8.7 members including 4.4 adults (1.8 females, 2.5 males), 2.9 subadults, and 1.4 juveniles. The number of juveniles was significantly positively correlated to the number of adult males. Groups with one or two adult males had significantly fewer juveniles (mean: 1:1 juveniles) than groups containing more than two adult males (mean: 2.0 juveniles). Apart from a different number of subadults, results showed obvious similarities between common marmosets and tamarins of the genus Saguinus in size and composition of subgroups of adults as well as the key role of adult males in mediating the reproductive success of a breeding female. Common marmoset females seem to gain direct fitness benefits in increased reproductive success from the presence of a larger number of adult males. Whether or not other group members get fitness benefits depends on the reproductive strategy of adult males (monogamy vs. polyandry), their kinship, and on the genetic relationship of nonbreeders to the offspring of the breeding female. © 1995 Wiley-Liss, Inc.     

The timing of gonadal development and moult in three raptors with different breeding seasons: effects of gender, age and body condition

Differences between species in breeding seasons are thought to be mediated through differences in their reproductive physiology. Little is known about how the timing and duration of gonadal maturation varies between raptor species, how the timing of moult relates to the gonadal cycle, whether the timing and degree of sexual maturation varies between juveniles and adults or whether body condition has a significant effect. To address these questions, data on gonadal size and moult for adults and juveniles of both sexes of three raptor species were extracted from the Predatory Bird Monitoring Scheme (based on birds found dead by members of the public). The three species, Sparrowhawk Accipiter nisus, Kestrel Falco tinnunculus and Barn Owl Tyto alba, have different ecologies – diurnal bird predator, diurnal mammal predator and nocturnal mammal predator, respectively. All are single‐brooded but have different breeding seasons. The duration of gonadal maturation was markedly different between the species. Barn Owls showed the earliest maturation and the latest gonad regression, and Sparrowhawks the latest maturation and earliest gonad regression. Kestrels were intermediate. In males of all species, the testes remained fully mature throughout their respective breeding seasons. In females, the ovaries remained partially mature throughout the breeding season. Moult started slightly earlier in Sparrowhawks than in Kestrels and coincided with gonadal regression in the two species. Although females of the two species started to moult earlier than males, moult duration was similar between the sexes. Barn Owls showed no distinct annual pattern of moult. In juveniles of all three species, the gonads were smaller than in adults throughout spring and started to mature later. Gonad size in birds that had starved tended to be smaller than in birds dying from other causes, but did not influence the difference in gonad mass between adults and juveniles and between seasons. Body condition had no effect on moult. Whilst ecology has led to the evolution of different breeding seasons, differences between species, and between adults and juveniles, are mediated through adaptive differences in their reproductive physiology.     

Socio-genetic correlates of unbiased sex dispersal in a population of black capuchin monkeys (<Emphasis Type="Italic">Sapajus nigritus</Emphasis>)

Most social mammal species exhibit male-biased dispersal. Sex bias in dispersal leads to a higher degree of relatedness among individuals of the philopatric sex, thus an atypical dispersal pattern might lead to deviations in the typical within-group kinship structure. Kinship, in turn, influences patterns of social interactions, as widely evident by kin-biased behaviors. We investigated the link between dispersal, relatedness structure, and sociopositive interactions established by adult females of black capuchin monkeys (Sapajus nigritus) living in a population that experiences female dispersal, an unusual pattern for capuchin monkeys. The study was conducted in Parque Estadual Carlos Botelho (PECB), within the Brazilian Atlantic Forest. We addressed dispersal and relatedness patterns by genotyping 20 adults of 3 groups across 9 microsatellite loci. We also sampled the monkeys’ behavior and compared spatial association frequencies and rates of grooming among same- and opposite-sex dyads. There was no difference between males and females in genetic parameters; both males and females show low coefficients of relatedness indicating that neither sex is consistently philopatric. The mean pairwise coefficient of relatedness for co-resident females was not higher than that for co-resident males. Compared to other populations of capuchin monkeys, female bond was weak, as evident by lower spatial association frequencies, reduced rates of grooming and lack of correlation between coefficients of relatedness and measures of dyadic sociopositive interactions. Our findings thus confirm that female dispersal is a habitual process in the capuchin population of PECB, and that, as expected, dispersal by females strongly influences the relatedness structure of the population as well as the affiliative relationships among female groupmates.     

Fine-scale social and spatial genetic structure in Sitka black-tailed deer

The spatial extent of Sitka black-tailed deer (Odocoileus hemionus sitkensis) populations below the regional scale is relatively unknown, as is dispersal between populations. Here, we use noninvasive samples to genotype 221 Sitka black-tailed deer in three watersheds on Prince of Wales Island, Alaska, separated by a maximum of 44 km, using traditional and spatial genetic approaches. We find that despite geographic proximity, multiple lines of evidence suggest fine-scale genetic structure among the three study sites. The 2 most geographically distant watersheds differed significantly in genetic composition, suggesting an isolation-by-distance pattern. Within study sites, deer exhibited spatial genetic structure within a radius of 1,000 m. Based on a reduced sample of known-sex individuals, females exhibited positive spatial genetic structure within a radius of 500 m but males showed no structure. Moreover, females were more likely to be related to their 5 nearest female neighbors, regardless of distance, than were males. Evidence indicates dispersal by both sexes although it may be more common, or dispersal distances are greater, in males. Nonetheless, analysis of assignment indices and comparison of sex-specific correlograms found no evidence of sex-biased dispersal between watersheds. Patterns of spatial relatedness and connectivity suggest limited dispersal among Sitka black-tailed deer, creating genetic structure on a fine spatial scale, perhaps as small as the watershed.     

Patterns of infant handling and relatedness in Barbary macaques (<Emphasis Type="Italic">Macaca sylvanus</Emphasis>) on Gibraltar

Among papionin primates, the Barbary macaque (Macaca sylvanus) shows the most extensive interactions between infants and group members other than the mother. Two different types of interactions occur: (1) long-lasting dyadic interactions between a handler and an infant, and (2) brief triadic interactions between two handlers involving an infant. Previous investigations showed that infant handling by males is best explained as use of infants to manage relationships with other males. In contrast, no adaptive explanation for infant handling by females emerged. Here, we compared the infant-handling pattern between subadult/adult males and subadult/adult females in a free-ranging group of 46 Barbary macaques on Gibraltar to test whether the relationship management hypothesis also applies to female handlers. We further investigated the infant-handling pattern of juveniles and used microsatellite markers to estimate relatedness between infant handlers and the infant’s mother. We found that males, females and juveniles all participated extensively in triadic interactions using infants of above-average related females. In contrast, only males and juveniles were highly involved in dyadic interactions with infants of related females, while females rarely handled infants other than their own. The pattern of infant handling was entirely compatible with the predictions of the relationship management hypothesis for males and mostly so for females. Moreover, our genetic analysis revealed that males and females differ in their partner choice: while females preferred to interact with related females, males had no significant preference to interact with related males. We further discuss the observed above-average relatedness values between infant handlers and the infant’s mother in the light of kin-selection theory.     

Demographic parameters and events in wild moustached Tamarins (Saguinus mystax)

This paper examines demographic events in the context of population structure and genetic relationships in groups of wild moustached tamarins (Saguinus mystax). We used a combination of long-term behavioral observations and genetic data from a total of eight groups from a population in northeastern Peruvian Amazonia. The mean group size was 6.0 (range = 4-9), including 2.5 adult males and 1.8 adult females. Within-group relatedness was generally high (r = 0.3), and most nonbreeding individuals were either natal or closely related to the respective same-sex breeder. The mean annual persistence of adults in the groups was 70% and 68% for males and females, respectively, and the reproductive tenure of one breeding pair lasted for at least 6 years. Migrations predominantly occurred after stability-disrupting events such as the immigration of new individuals and/or the loss of breeding individuals, or when groups were rather large. Migrations of both breeding and nonbreeding males and females occurred. Our results show that the hypothesis of Ferrari and Lopes Ferrari [Folia Primatologica 52:132-147, 1989] that tamarins live in smaller and less stable groups with lower relatedness compared to marmosets does not generally hold true. In contrast, we found that tamarin groups can consist of predominantly related individuals, and are stable as well. It is also apparent that a single demographic event can produce a chain of subsequent complex demographic changes.     

Cryptic behaviours, inverse genetic landscapes, and spatial avoidance of inbreeding in the Pacific jumping mouse

Although the behaviour of individuals is known to impact the genetic make-up of a population, observed behavioural patterns do not always correspond to patterns of genetic structure. In particular, philopatric or dispersal-limited species often display lower-than-expected values of relatedness or inbreeding suggestive of the presence of cryptic migration, dispersal, or mating behaviours. I used a combination of microsatellite and mark-recapture data to test for the influence of such behaviours in a dispersal-limited species, the Pacific jumping mouse, within a semi-isolated population over three seasons. Despite short dispersal distances and a low rate of first generation migrants, heterozygosities were high and inbreeding values were low. Dispersal was male-biased; interestingly however, this pattern was only present when dispersal was considered to include movement away from paternal home range. Not unexpectedly, males were polygynous; notably, some females were also found to be polyandrous, selecting multiple neighbouring mates for their single annual litter. Patterns of genetic structure were consistent with these more inconspicuous behavioural patterns. Females were more closely related than males and isolation by distance was present only in females. Furthermore, detailed genetic landscapes revealed the existence of strong, significant negative correlations, with areas of low genetic distance among females overlapping spatially with areas of high genetic distance among males. These results support the hypothesis that the detected cryptic components of dispersal and mating behaviour are reducing the likelihood of inbreeding in this population through paternally driven spatial mixing of male genotypes and polyandry of females.     

Temporal genetic structure and relatedness in the Tufted Duck Aythya fuligula suggests limited kin association in winter

Conspecific aggregation of waterfowl in winter is a common example of animal flocking behaviour, yet patterns of relatedness and temporal substructure in such social groups remain poorly understood even in common species. A previous study based on mark‐recapture data showed that Tufted Ducks Aythya fuligula caught on the same day were re‐caught together in subsequent winters more often than expected by chance, suggesting stable assortments of ‘socially familiar’ individuals between wintering periods. The genetic relationships within these social groups were not clear. Based on 191 individuals genotyped at 10 microsatellite markers, we investigated the temporal genetic structure and patterns of relatedness among wintering Tufted Ducks at Lake Sempach, Switzerland, in two consecutive winters. We found no evidence of genetic differentiation between temporal groups within or between winters. The average levels of relatedness in temporal groups were low and not higher than expected in random assortments of individuals. However, Mantel tests performed for each sex separately revealed significant negative correlations between the pairwise relatedness coefficients and the number of days between the capture dates of pairs of wintering Tufted Duck in males and females. This pattern suggests the presence of a small number of co‐migrating same‐sex sibling pairs in wintering flocks of Tufted Ducks. Our findings provide one of the first genetic analyses of a common duck species outside the breeding season and contribute to the understanding of social interactions in long‐distance migratory birds.     

Relatedness and demography of African forest elephants: inferences from noninvasive fecal DNA analyses

African forest elephants (Loxodonta cyclotis) are genetically and morphologically distinct from their savannah counterparts, but their biology remains poorly understood. In this study, I use noninvasive fecal DNA analyses to examine the relatedness structure and historical demography of forest elephants at 2 sites in SW Gabon, central Africa. Pairwise relatedness values calculated between 162 elephant individuals genotyped at 8 microsatellite loci were significantly higher within spatially associated dung piles than between random pairings for one site. First- and second-order relatives were most commonly detected among dung piles from adult female pairs and adult females and juveniles. Pairwise relatedness estimates suggested that, like savannah elephants, forest groups are largely composed of adult females, their sisters, and juvenile offspring. Associations between males, and groups containing juveniles from multiple related females, were detected but at much lower frequency. Analysis of mitochondrial d-loop sequences from 70 elephant individuals identified 2 haplogroups in SW Gabon.     

Facultative dispersal by juvenile males in the cooperative stripe-backed wren

We present genetic and demographic data documenting juveniledispersal in the cooperatively breeding stripe-backed wren (Campylorhynchusnuchalis) of Venezuela. Parentage and DNA fragment-sharing analysesrevealed 12 cases in which juveniles were unrelated to othergroup members. Of these 12 foreign juveniles, (1) all were males,(2) eight of 12 had been found with breeding pairs lacking helpersrather than with groups containing helpers, and (3) four outof seven of those observed as adults courted or sired offspringwith the dominant females in their new groups despite the strongincest avoidance of this species. Furthermore, juvenile maleshad a significant tendency to disappear from natal groups intheir first year, and singleton juveniles observed with pairsafter the breeding season were mostly males. These data supportthe hypothesis that foreign juveniles were dispersers from intactgroups and not products of conspecific brood parasitism or adoptionfollowing group dissolution. We suggest that unassisted pairsmight accept juvenile males into their groups as helpers toincrease their future reproductive success and that dispersersthemselves might leave large natal groups in which their helpingis superfluous to join small groups of nonrelatives in whichthey might soon reproduce.