Geographic patterns of natal dispersal in barn swallows <Emphasis Type="Italic">Hirundo rustica</Emphasis> from Denmark and Spain

Dispersal is an important factor that determines the degree of gene flow and, hence, the degree of differentiation among populations. Using two long-term datasets on natal philopatry and short-distance dispersal in barn swallows Hirundo rustica from Denmark and Spain, we evaluated the fitness costs and benefits and test a number of predictions about the functional significance of dispersal. The proportion of philopatric individuals was more than six times larger in Spain than in Denmark, with a higher rate of philopatry in males than in females. Dispersal propensity decreased in both populations during the course of the study. Males from the more philopatric Spanish population lived longer when philopatric rather than dispersing while that was not the case for either sex of the less philopatric Danish population. There were large differences in dispersal propensity among cohorts and breeding sites, suggesting that sites differed in their suitability as sites for immigrants. We found no evidence consistent with the mate competition hypothesis suggesting that males in better condition or with larger condition-dependent secondary sexual characters were more likely to be philopatric. These findings suggest that there is a high degree of intraspecific variation in dispersal propensity between populations, probably relating to local differences in costs and benefits of philopatry and dispersal.     

Fitness costs of dispersal in red foxes (Vulpes vulpes)

The costs of dispersal are an important factor promoting natal philopatry, thereby encouraging the formation of social groups. The red fox, Vulpes vulpes, exhibits a highly flexible social system and one that is thought to represent a possible stage in the evolution of more complex patterns of group-living. Although the potential benefits accruing to philopatric offspring have previously been studied in this species, the potential costs of dispersal have received less attention. We contrasted survival rates, nutritional status, injuries and reproductive output of dispersing and non-dispersing male and female foxes in an urban population to assess the relative costs of dispersal versus natal philopatry. Mortality rates were not significantly higher for dispersing foxes, either in the short- or long-term. There was no evidence of increased nutritional stress in dispersing individuals. Dispersing individuals did, however, exhibit greater levels of wounding, although this did not appear to affect survival. Dispersing females were more likely to miss a breeding opportunity early in their reproductive lifespan. In contrast, both dispersing and non-dispersing males were unlikely to breed in their first year. We conclude that the major fitness component in females affected by dispersing is age at first reproduction.     

Philopatry and mother-daughter associations in bushy-tailed woodrats: space use and reproductive success

Social groups in mammals are often based on overlapping generations of philopatric females, yet few researchers have examined fitness consequences of associations between females and their daughters. We examined survival and reproductive consequences of sharing a natal site with one's mother or daughter in wild bushy-tailed woodrats (Neotoma cinerea). Bushy-tailed woodrats reside on discrete rocky outcrops, and females tend to spend their entire reproductive life on their natal outcrop. Mothers and daughters that co-existed on the natal outcrop were closely associated in space. Juvenile females were more likely to survive their first winter to become breeding yearlings at their natal site if their mother was also present at the natal site during the winter. Juveniles survived equally well with or without their mother where densities of adult females were experimentally reduced, suggesting that females enhance their daughter's survival by facilitating their access to limited resources. Yearling females exhibited no reproductive costs from association with their mothers at the natal site during the breeding season; in fact, yearlings that bred at the natal site simultaneously with their adult mothers tended (P = 0.07) to experience fewer reproductive failures than did those that bred in the absence of their mothers. Mothers did not experience any detectable costs when sharing a natal site with a daughter. We conclude that bushy-tailed woodrats exhibit stable, fitness-enhancing associations among mothers and their philopatric daughters.     

Natal philopatry in bannertailed kangaroo rats

Summary I describe prolonged retention of offspring in natal home ranges, or natal philopatry, in the bannertailed kangaroo rat Dipodomys spectabilis. Though weaning occurs at about one month of age, offspring shared natal burrows with their mothers for three to seven months, and 39% of surviving offspring remained within natal home ranges through reproductive maturity. Males as well as females were philopatric. Data on the availability of burrows and on patterns of resettlement suggest that natal philopatry in this species may be a means of providing juveniles with access to essential resources, in this case food caches and large complex burrow systems, that are not readily available outside natal home ranges.     

Genetic relatedness and space use in a behaviorally flexible species of marmot, the woodchuck (Marmota monax)

Solitary species show several patterns of space use and relatedness. Individuals may associate randomly or may live near female or male kin, often as a result of natal philopatry or dispersal patterns. Although usually described as solitary or asocial, woodchucks (Marmota monax) are behaviorally flexible marmots that exhibit greater sociality in some populations than others. I examined relationships between kinship, geographic distance, and home range overlap, as well as dispersal and philopatry, to determine the extent to which kin associated spatially. I used a combination of microsatellite DNA analysis, long-term behavioral observations, and radiotelemetry to test predictions that females, but not males, would associate with kin. Indeed, woodchucks lived closer and shared a greater proportion of their home range with more closely related animals. Overlap of females' and males' home ranges was positively correlated with kinship, and male–female dyads shared more area with closer kin. Most juveniles delayed dispersal beyond their first summer. Females often remained philopatric and settled near their natal range. Although males often dispersed as yearlings, some males also established territories within or immediately adjacent to their natal home ranges. A combination of factors can explain these spatial patterns, including high population density associated with the study site's location within a suburban environment, high dispersal costs, and abundant food. Thus, despite their asocial and solitary reputation, woodchucks displayed spatial patterns seen in other, more social species of ground-dwelling sciurids.     

Sex-specific fitness consequences of dispersal in Siberian jays

In most birds, natal dispersal is female-biased, but the selective pressures leading to this pattern have rarely been explored with comprehensive data on lifetime reproductive success. In territorial birds, the benefit of philopatry should be higher for males than for females when males establish territories for which knowledge about the local environment is important. As females may use male characteristics for mate choice, and hence indirectly for territory choice, the benefit from the direct knowledge of the local environment may be lower for females than males. We tested this hypothesis using data from a long-term study of group living corvids, the Siberian jays (Perisoreus infaustus). In this species, the socially dominant offspring delay dispersal while the sub-dominant offspring leave the family group directly after reaching independence. Our results show that natal dispersal distance (a proxy for local knowledge) was related to sex and dispersal timing (a proxy for “quality”): Females and early dispersers traveled further on average than males and delayed dispersers. Furthermore, dispersal distance and timing were negatively related to the number of recruits produced over an individual’s lifetime in males, but not in females. Hence, the results support the hypothesis that the female-biased natal dispersal found in this and other bird species may come about through higher lifetime reproductive success of philopatric males than females.     

Philopatry and nest founding in a primitively social bee,Halictus rubicundus

Summary Observations of inter- and intraseasonal dispersal patterns in the primitively social sweat bee Halictus rubicundus in New York reveal considerable philopatry in both sexes. Females overwinter away from the nest aggregation, in diapause for 8–11 months, and return to dig new nests close to the site of their natal nests (typically within 50 cm). Nests are all founded by siggle females (haplometrosis). Clumping of nests may reflect patterns of soil vegetation. Workers rarely enter or take up residence in non-natal nests. Males commonly return to both natal and non-natal nests, and actively patrol vegetation near the natal aggregation for females. These philopatric tendencies, in combination with the haplometrotic mode of nest founding, should have important effects on population structure, particularly in terms of population subdivision, mate choice, and intracolony interactions. The resulting population structure may be conducive to the evolution and maintenance of social behavior.     

Natal philopatry and breeding area fidelity of long-billed curlews (Numenius americanus): Patterns and evolutionary consequences

Summary Long-billed curlews (Numenius americanus) appear unique among scolopacid shorebirds so far studied in possessing a significant sex bias in natal philopatry. We resighted 9 curlews at least attempting to breed that were color-banded as chicks; 8 of these were males. Male curlews also cooperate extensively with neighbors in mobbing potential chick predators. This mutualistic behavior may have evolved through kin selection among philopatric males. If so, we would expect such an evolutionary consequence to lead to a similar sex bias in breeding area fidelity. Yet our resightings of colorbanded adults over 4 consecutive years indicate that males and females were equally likely to return to previous nesting territories. Excessive disturbance such as capture and nest loss within a single breeding season was correlated with the likelihood of breeding dispersal by females but not males. This suggests potentially stronger breeding area fidelity of males.     

Socially induced delayed primiparity in brown bears Ursus arctos

Reproductive suppression through behavioral or physiological means is common in group-living and cooperative breeding mammals, but to our knowledge it has not been shown in wild large carnivores other than those with a clear form of social organization. Brown bear (Ursus arctos) females form matrilinear assemblages with related females using a common and largely exclusive area. Behavioral reproductive suppression might develop due to a hierarchical system among females within a matrilinear assemblage or due to inbreeding avoidance, because male brown bears can overlap with their daughters. We tested whether natal dispersal influenced the age of primiparity. We predicted that emigrant females, geographically removed from maternal or paternal influence, would reproduce earlier than philopatric females. The average age of primiparity was 4.3 years in females that dispersed outside their mother’s home range (n=8) and 5.2 years in philopatric females (n=10). Only the overlap with the mother’s home range, and not body size, body mass, growth, local population density, or overlap with the father’s home range, had a significant influence on the age of primiparity. The ultimate role of reproductive suppression for brown bears is likely to avoid inbreeding or to minimize resource competition. Due to the low risk of inbreeding and frequent exposure of young females to unrelated males, we conclude that resource competition within female hierarchies causes reproductive suppression in young females.     

Why do female Belding's ground squirrels disperse away from food resources?

We examined the effects of food provisioning on the natal dispersal behavior of Belding's ground squirrels (Spermophilus beldingi). We provided extra food to adult and yearling females in their maternal territories during pregnancy and lactation, and to offspring of these females in their natal areas for 6 weeks after weaning. We used unprovisioned young of unprovisioned mothers as controls. Provisioning influenced the probability of dispersal from the natal area by female but not male S. beldingi. All surviving male S.␣beldingi dispersed by 55 weeks of age, regardless of whether they and their mothers received extra food. By contrast, we observed a significant trend, beginning 3 weeks after weaning and continuing through the yearling year, for a greater proportion of provisioned than control female S. beldingi to emigrate from the natal area. Competition for food did not appear to influence natal dispersal of females. However, overall population density, density of females weaning litters, and rates of aggression and vigilance among these females, were higher in provisioned than control areas, suggesting that competition for non-food resources was unusually intense in provisioned areas. We propose that juvenile female, but not juvenile male, S. beldingi may emigrate from the natal site to increase access to areas with low densities of conspecifics. Together with findings of earlier workers, our results suggest that spatial and temporal distributions of environmental resources are important influences on the dispersal behavior of female ground squirrels. Received: 28 February 1996 / Accepted after revision: 16 October 1996     

Costs of delayed dispersal and alloparental care in the fungus-cultivating ambrosia beetle <Emphasis Type="Italic">Xyleborus affinis</Emphasis> Eichhoff (Scolytinae: Curculionidae)

Body reserves may determine the reproductive output of animals, depending on their resource allocation strategy. In insects, an accumulation of reserves for reproduction is often obtained before dispersal by pre-emergence (or maturation) feeding. This has been assumed to be an important cause of delayed dispersal from the natal nest in scolytine beetles. In the cooperatively breeding ambrosia beetles, this is of special interest because in this group delayed dispersal could serve two alternative purposes: “selfish” maturation feeding or “altruistic” alloparental care. To distinguish between these two possibilities, we have experimentally studied the effect of delayed dispersal on future reproductive output in the xyleborine ambrosia beetle Xyleborus affinis. Females experimentally induced to disperse and delayed dispersing females did not differ in their body condition at dispersal and in their founding success afterwards, which indicates that females disperse independently of condition, and staying adult females are fully mature and would be able to breed. However, induced dispersers produced more offspring than delayed dispersers within a test period of 40 days. This suggests that delayed dispersal comes at a cost to females, which may result primarily from alloparental care and leads to a reduced reproductive output. Alternatively, females might have reproduced prior to dispersal. This is unlikely, however, for the majority of dispersing females because of the small numbers of offspring present in the gallery when females dispersed, suggesting that mainly the foundress had reproduced. In addition, “gallery of origin” was a strong predictor of the reproductive success of females, which may reflect variation in the microbial complex transmitted vertically from the natal nest to the daughter colony, or variation of genetic quality. These results have important implications for the understanding of proximate mechanisms selecting for philopatry and alloparental care in highly social ambrosia beetles and other cooperatively breeding arthropods.     

Natal dispersal and philopatry in prairie voles (Microtus ochrogaster) in relation to population density,season, and natal social environment

Summary We used intensive livetrapping to examine natal dispersal and philopatry in prairie voles (Microtus ochrogaster). The majority of male (70.0%) and female prairie voles (75.1 %) remained at the natal nest until death. Those males and females that did disperse left home at about the same age (45–55 days) and moved similar distances (28–33 m). Dispersal was more common (1) from small natal groups than from large natal groups, (2) following disappearance of parents, (3) during the breeding period than during the nonbreeding period, and (4) at low population densities than at high densities. Dispersal was not associated with level of competition for mates within natal groups, and dispersers did not differ from nondispersers in body weight. Our data do not support competition for mates or resources as important factors influencing natal dispersal in prairie voles. The absence of sex differences in dispersal tendency or distance, and our fording that more than half of dispersers had become reproductive before leaving the natal nest, lead us to suggest that inbreeding avoidance is not a primary function of dispersal in this species. Dispersal was, however, more common when potential mates within the natal group were relatives than when they were nonrelatives. Although not tested here, if family members avoid mating with one another through patterns of mate choice, then some animals may leave home in search of mates. The precise benefits associated with philopatry in prairie voles remain to be identified. Correspondence to: B. McGuire at her present address     

Female philopatry and its social benefits among Bornean orangutans

Female philopatry in mammals is generally associated with ecological and sometimes social benefits, and often with dispersal by males. Previous studies on dispersal patterns of orangutans, largely non-gregarious Asian great apes, have yielded conflicting results. Based on 7?years of observational data and mitochondrial and nuclear DNA analyses on fecal samples of 41 adult Bornean orangutans (Pongo pygmaeus wurmbii) from the Tuanan population, we provide both genetic and behavioral evidence for male dispersal and female philopatry. Although maternally related adult female dyads showed similar home-range overlap as unrelated dyads, females spent much more time in association with known maternal relatives than with other females. While in association, offspring of maternally related females frequently engaged in social play, whereas mothers actively prevented this during encounters with unrelated mothers, suggesting that unrelated females may pose a threat to infants. Having trustworthy neighbors may therefore be a social benefit of philopatry that may be common among solitary mammals, thus reinforcing female philopatric tendencies in such species. The results also illustrate the diversity in dispersal patterns found within the great-ape lineage.     

Competitive regimes and female bonding in two species of squirrel monkeys (Saimiri oerstedi and S. sciureus)

Summary Ecological and behavioral data from long-term field studies of known individuals in two closely related squirrel monkey species (Saimiri oerstedi and S. sciureus) were used to examine hypotheses about the source of variation in female bonding among group-living primates. Social relationships in species which live in cohesive groups are thought to depend on the nature of competition for resources. S. oerstedi and S. sciureus both live in large groups and are subject to intense predation. Direct feeding competition both between and within groups is extremely low in S. oerstedi; in this species female relationships are undifferentiated, no female dominance hierarchy is evident and females disperse from their natal group. S. sciureus also experiences very low levels of between-group competition, but within-group direct competition for resources is frequent; this species demonstrates differentiated female relationships, a female dominance hierarchy, and female philopatry. The correlated ecological and social variables found in these two congeners further minimize the minor effects of phylogenetic differences and emphasize the importance of food distribution in determining social characteristics. Offprint requests to: S. Boinski     

Fitness consequences of dispersal decisions for male mountain gorillas (Gorilla beringei beringei)

The fitness consequences of dispersal decisions are difficult to quantify, especially for long-lived species with complex social systems. To calculate those consequences for male mountain gorillas, from the perspective of both subordinate and dominant males, we used behavioral and demographic data obtained over 30 years from the Virunga Volcano population to develop an agent-based model that simulates the life history events, social structure, and population dynamics of the species. The model included variables for birth rates, mortality rates, dispersal patterns, and reproductive skew. The model predicted an average lifetime reproductive success (LRS) of 3.2 for philopatric males (followers) and 1.6 for emigrants. The benefits of philopatry were most sensitive to opportunities for social queuing and to female transfer preferences, but philopatry remained the best strategy over a wide range of group conditions and hypothetical simulations. The average LRS for dominant males was 4.5 when a subordinate stayed and 4.6 when the subordinate emigrated. The dispersal decision of the subordinate male had little impact on the fitness of the dominant male because it came relatively late in the dominant males reproductive life span, and it changed his group composition only incrementally. The fitness consequences for the dominant male were most sensitive to the degree of reproductive skew. Since subordinates suffer a fitness loss when they leave a group, they should accept whatever reproductive restraint is needed to avoid eviction, and the dominant male does not need to offer concessions for them to stay. The dominant male may offer reproductive concessions for other reasons, such as peace incentives or to confuse paternity, or he may not have complete control of reproduction within his group.Communicated by D. Watts     

What are the strengths and limitations of direct and indirect assessment of dispersal? Insights from a long-term field study in a group-living bird species

Molecular methods of assessing dispersal have become increasingly powerful and have superseded direct methods of studying dispersal. Although now less popular, direct methods of studying dispersal remain important tools for understanding the evolution of dispersal. Here, we use data from Siberian jays Perisoreus infaustus, a group-living bird species, to compare natal dispersal distances and rates using visual mark–recapture, radio-tracking and microsatellite data. Siberian jays have bimodal natal dispersal timing; socially dominant offspring remain with their parents for up to 5 years (delayed dispersers), while they force their subordinate brood mates to leave the parental territory at independence (early dispersers). Early dispersers moved about 9,000 m (visual mark–recapture, radio-tracking) before settling in a group as a non-breeder. In contrast, delayed dispersers moved about 1,250 m (visual mark–recapture only) and mainly moved to a breeding opening. Dispersal distances were greater in managed habitat compared to natural habitat for both early and delayed dispersers. Molecular estimates based on 23 microsatellite loci and geographical locations supported distance estimates from the direct methods. Our study shows that molecular methods are at least 22 times cheaper than direct methods and match estimates of dispersal distance from direct methods. However, molecular estimates do not give insight into the behavioural mechanisms behind dispersal decisions. Thus, to understand the evolution of dispersal, it is important to combine direct and indirect methods, which will give insights into the behavioural processes affecting dispersal decisions, allowing proximate dispersal decisions to be linked to the ultimate consequences thereof.     

Genetic reconstruction of breeding patterns in gundis (Rodentia: Ctenodactylidae)

Current mating system theory predicts that the number of females breeding in a group will depend on the number of females in the group and the accessibility of unrelated males, whereas the number of males breeding in a group will depend on the ability of males to control access to reproductive females. By combining information on group composition with genetic data, I determined whether breeding patterns in a rock-dwelling rodent, Ctenodactylus gundi, were concordant with these expectations. C. gundi breeding patterns varied from facultative monogamy to uni-male polygyny to multi-male polygyny. The number of reproductive individuals of each sex in a group increased with group size. Whereas communal breeding among related females tended to increase female reproductive success, males breeding in the same group were unrelated to other group members and seemed to compete over access to matings. While some males were assigned offspring from neighboring social groups, most group-living males successfully monopolized the reproduction of females within their group. There was no evidence that females had multiply sired litters, although some bred with alternative males in separate breeding attempts. Although numerous individuals were not assigned as parents or offspring, genetic information enabled me to determine that most unassigned individuals were philopatric group members, whereas only a few were unrelated immigrants into their current social group. Together, these results provide evidence that C. gundi social groups represent fairly distinct reproductive units whose breeding patterns are dependent on group size and composition in accordance with theoretical predictions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Sex-biased dispersal in a long-lived polygynous reptile (Crocodylus johnstoni )

We compared natal dispersals of freshwater crocodiles (Crocodylus johnstoni) against the prediction of male dispersal bias for a polygynous mating system. The crocodiles inhabited a linear series of pools and we calculated the net distances from natal pools to recapture locations some 12–18 years later, at maturity. Philopatry was assessed in terms of adult social distances. A female social distance was 0.46 pools and a male social distance was 1.0 pool. By these criteria, both sexes showed low levels of philopatry (7–12%). However, individuals of both sexes dispersed from the natal site long before they were sexually mature. Divergence in dispersal patterns by sex occurred after the maturity threshold, as males dispersed two to three times farther than females. Intrasexual competition by males is resolved by a size-based hierarchy. The displacement of small males from local mating access is a probable cause of the longer dispersals undertaken by males. Competition, rather than inbreeding avoidance, is driving dispersal in this population of freshwater crocodiles. Received: 8 May 1998 / Accepted after revision: 26 June 1998     

Sex-specific effects of the local social environment on juvenile post-fledging dispersal in great tits

An individual’s decision to disperse from the natal habitat can affect its future fitness prospects. Especially in species with sex-biased dispersal, we expect the cost–benefit balance for dispersal to vary according to the social environment (e.g., local sex ratio and density). However, little is known about the social factors affecting dispersal decisions and about the temporal and spatial patterns of the dispersal process. In our study, we investigated experimentally the effects of the social environment on post-fledging dispersal of juvenile great tits by simultaneously manipulating the density and sex ratio of fledglings within forest plots. We expected young females in the post-fledging period mainly to compete for resources related to food and, as they are subordinate to males, we predicted higher female dispersal from male-biased plots. Juvenile males compete for vacant territories already in late summer and autumn; thus, we predicted increased male dispersal from high density and male-biased plots. We found that juvenile females had a higher probability to leave male-biased plots and had dispersed further from male-biased plots in the later post-fledging phase when juvenile males start to become territorial and more aggressive. Juvenile males were least likely to leave male-biased plots and had smallest dispersal distances from female-biased plots early after fledging. The results suggest that the social environment differentially affected the costs and benefits of philopatry for male and female juveniles. The local sex ratio of individuals is thus an important social trait to be considered for understanding sex-specific dispersal processes.     

Maternal investment and sex-allocation in the Galapagos fur seal,Arctocephalus galapagoensis

Summary Maternal investment and sex-allocation were measured in a large, sexually dimorphic mammal, the Galapagos fur seal (Arctocephalus galapagoensis). The sex ratio at birth was 1.06. Males were always heavier than females and, at least initially, grew faster. Growth was variable from year to year suggesting energetic constraints on maternal investment. Sucking time conrrelated with milk intake. Mothers suckled yearling and 2-year-old sons more than daughters of the same age. Age at weaning appeared to be the same in both sexes or even slightly greater in males. No sex differences was found in mortality prior to weaning or in post-weaning dispersal. Birth rates of females with yearlings or 2-year-olds were significantly lower than those of females with no dependent young. Mothers invested more in sons than in daughters until weaning. It is unlikely that higher post-weaning investment in daughters balances the higher pre-weaning investment in sons. Data on sex ratio at birth, different growth rates, and weaning age of the sexes are typical of otariid seals as a group. The results of this study fit Maynard Smith's (1980) model of the evolution of sex allocation better than Fisher's (1930).