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.     

Is kin cooperation going on undetected in marine bird colonies?

In multitudinous breeding colonies, kin interactions could go unnoticed because we are unaware of the kinship among adults we observe. Evidence of cooperation and competition between close adult kin in a blue-footed booby colony was sought by analyzing patterns of natal dispersal and proximity of nests. Male and female recruits nested closer to their own natal sites than to their parents’ current sites. Males (only) dispersed less far when both parents were present than when no parent or one parent was present, but not selectively close to fathers versus mothers when these were divorced. Neither parental presence nor parental proximity affected breeding success of recruits of either sex. Although distances between the nests of simultaneously recruiting broodmates were unrelated to their sex, males dispersed 13.1 m less when a sister was present than when a brother was present. Neither sex was affected in its dispersal distance by the presence or hatching order/dominance of a broodmate. Neither sex was affected in its breeding success by the presence versus absence of a broodmate, although female success increased with proximity of their brothers. Parents and sisters may actively or passively help males establish their first territories near their natal sites and nearby brothers may help females in their first breeding attempts; otherwise, boobies do not influence each other’s natal dispersal and first breeding success. It appears that boobies do not nest selectively close to or far from their parents, offspring, or broodmates. Why there is apparently so little cooperation and altruism between close adult relatives in booby colonies is puzzling.     

Sociogenetic structure in a free-living nocturnal primate population: sex-specific differences in the grey mouse lemur (Microcebus murinus)

The grey mouse lemur (Microcebus murinus) has a dispersed social structure, within which female sleeping associations are common. These sleeping associations have been hypothesised to confer anti-predatory and thermoregulatory benefits, especially when rearing offspring. The genetic composition of these associations was determined using microsatellite markers to test predictions derived from kin selection theory. 161 (99 males, 62 females) individual M. murinus belonging to a free-living population in Ampijoroa, north-western Madagascar, were genotyped and observed over a total of 13 months distributed over the dry seasons of 3 successive years (1995-1997). Kin selection theory predicts that these female associations should consist of closely related members, and that female philopatry and male natal dispersal should characterise the dispersal pattern within this species. These predictions were confirmed by the data. Five out of six female sleeping groups consisted of one or more closely related dyads. Females that slept alone did not have close female kin in the vicinity or within the population at all. Closely related female dyads lived in significantly closer proximity than closely related male dyads and closely related male-female dyads showed intermediate proximity. In combination with the result that females possessed significantly more relatives within the population than males, these findings support the behavioural hypotheses of female philopatry and male natal dispersal. Matrilinear grouping patterns and sex-biased dispersal are therefore genetically established in a dispersed primate social organisation for the first time. The results further indicate that several generations of mouse lemurs live together within a given area, implying both an effective mechanism of kin recognition to avoid father-daughter incest and the potential for social learning to ensure individual recognition.     

Social organization of woodchucks (Marmota monax)

Summary The social organization of woodchucks (Marmota monax) in southeast Ohio was studied at two sites, at one for two 2 years (1979–1980) and the other for 3 years (1981–1983). Spatial organization was determined by trapping and radio tracking. The home ranges of adult females did not overlap in the early spring but during late spring and summer there was some overlap (<10%) as females expanded their home ranges. Adult females tended to occupy the same home range in consecutive years. Some adult males occupied well-defined home ranges that did not overlap the home ranges of other males but did overlap extensively the home range of one to three adult females. These males tended to occupy the same home range in consecutive years. Infants used the same home range of their dam until about 2–3 months of age when most males and females apparently dispersed. About 35% of the juvenile females did not disperse until their second spring, just before their mother's new litter first emerged from their burrow. The average social group consisted of an adult male with two female kin groups comprising an adult female, an offspring (usually female) of the previous year, and the young of the year. Interactions within the kin group and with the adult male were relatively frequent and generally amicable. Interactions between kin groups both within and between different social groups were relatively rare and agonistic. The social organization of woodchucks in Ohio differs from that described in previous studies of woodchucks elsewhere and from that predicted by current models proposed by others on the evolution of social organization of marmots.     

Movements of dispersing flying squirrels in relation to siblings and parents

Knowledge of kin interactions can be informative in explaining the processes underlying dispersal. By dispersing, relatives can avoid kin competition for resources or mates and prevent inbreeding. We investigated sibling movements in relation to each other and parents before and after dispersal in radio-collared juvenile flying squirrels (Pteromys volans L.). Before dispersal, most siblings moved in different areas and were not in contact while exploring the surroundings of natal area. After dispersal, all siblings settled far away from each other, as they dispersed different distances and to random directions compared to each other. No clear effect of litter size or presence of same-sex siblings was observed on dispersal. Time spent in the same nest with the mother did not differ between dispersing sons and daughters. Mother did not force dispersers to leave the natal nest; instead, the mother often moved away from the nest before offspring. Father and offspring were not observed to be in contact before dispersal. Dispersal direction was unaffected by the location of fathers’ home range. We conclude that after dispersal interaction between relatives is limited in flying squirrels, but we did not find any indication that interaction between relatives before dispersal is determining dispersal decision. Siblings’ settlement far away from each other was already indicated by the movement within the natal home range, but, interestingly, the latter could not be used to predict dispersal patterns of sexes.     

There is a limbo under the moon: what social interactions tell us about the floaters’ underworld

The ultimate and proximate causes of natal dispersal have been extensively investigated, but the behaviour of dispersers in relation to social interactions has been largely neglected. Here, we investigated the social organisation of floating individuals during their dispersal by analysing the behaviour of 40 radio-tagged eagle owls Bubo bubo during the wandering and stop phases of dispersal. Unexpectedly, eagle owl floaters mixed with conspecifics independently of their sex, age, phase of dispersal, birthplace, health status and habitat features, showing an ‘underworld’ of interactions characterised by the absence of obvious social organisation or short-term strategies. Non-breeding owls were not transient floaters that occurred at numerous sites for short periods of time but rather had fairly stable home ranges: they attempted to settle as soon as possible within well-defined home ranges. The spatial distribution pattern of floaters and high rates of home range overlap support the prediction that floating individuals are not spatially segregated, challenging the expectation that dominance by size, age and/or health status may determine the exclusive use of some portions of the dispersal area. Finally, (1) the short distances among conspecifics and the extensive home range overlaps allowed us to discard the possibility that neighbouring floaters represent a real cost during dispersal and (2) floater interactions showed a lack of clear mechanisms for avoidance of kin competition among offspring or inbreeding.     

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     

Subordinate males are more likely to help if unrelated to the breeding female in cooperatively breeding white-browed scrubwrens

Subordinates in communally breeding groups of birds usually help to provision nestlings, but in some species helping-at-the-nest is facultative. In species in which groups usually contain relatives, subordinates either always feed young or are more likely to do so when breeding dominants are close relatives, suggesting that benefits of helping collateral kin are important. In other species, adult group members are unrelated to each other and males may only feed young if they have gained paternity, showing that cooperation is related to the mating system. The white-browed scrubwren, Sericornis frontalis, is a communally breeding species in which most groups consist of a simple pair or a dominant pair with a subordinate male. Subordinate males either fed nestlings in a given nest at a rate comparable to the dominants, or did not feed them at all. Breeding groups usually formed through natal philopatry of males, so that about 80% of subordinates were closely related to one or both members of the dominant pair. However, because of death and dispersal, 54% of subordinates were unrelated to the resident female. Although subordinates with their mother fed nestlings in 48% of cases, they fed offspring in 75% of cases if their mother had been replaced by an unrelated female, suggesting that their decision to help is influenced by the opportunity to mate with the female. Supporting this conclusion, relatedness to the dominant male did not affect subordinate behaviour, and genetic studies showed that subordinates often gained paternity if unrelated to the female. Thus, paradoxically, provisioning nestlings is related to the opportunity for mating in a society in which there is natal philopatry and subordinates are usually related to one or both members of the dominant pair. Received: 25 January 1997 / Accepted after revision: 1 May 1997     

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.     

Relatedness and avoidance of inbreeding in the lizard, Tiliqua rugosa

Two potential mechanisms for reducing the level of inbreeding, sex-biased dispersal and kin avoidance, were examined in the Australian sleepy lizard, Tiliqua rugosa. The home range centres, and the genotypes at four polymorphic microsatellite DNA loci were determined for adult lizards in a 70-ha study area near Mount Mary, South Australia. From estimates of genetic relatedness, females were as closely related to other females as they were to males, both within the whole study area, and within home ranges. Similarly, males were as closely related to other males as they were to females. This suggests that dispersal in the population is not sex-biased. Sleepy lizards form monogamous pairs during the spring. Partners were less closely related to each other than to other potential partners in the home range area. This suggests active choice of unrelated partners. The mechanism for recognising related from unrelated individuals is unknown, but the behaviour could reduce inbreeding. Received: 7 November 1998 / Accepted: 30 May 1999     

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.     

Multiple socioecological factors influence timing of natal dispersal in kangaroo rats

Natal dispersal is an important event in the life history of many species. Timing of natal dispersal can influence survivorship and subsequent reproductive success. A variety of individual proximal factors determine if and when offspring disperse from the natal territory by influencing the costs of dispersing and the benefits of delaying dispersal. I examined the influence of multiple factors on dispersal age in the banner-tailed kangaroo rat (Dipodomys spectabilis), a solitary species lacking extreme sex-biased dispersal. I used an information theoretic approach to compare Cox proportional hazards regression models of dispersal age for 121 offspring over a 3-year period consisting of low and high population densities. The top-ranked models indicated that dispersal age was influenced by a combination of socioecological factors related to resource competition, environmental conditions, kin competition, and a lesser extent sex. Circumstances that likely reduced the probability of successful dispersal such as intense resource competition at high population density and being born earlier in the breeding season when environmental conditions were poor lead to longer delays in natal dispersal. Offspring in larger litters also dispersed earlier possibly to avoid competition with kin. Sex was weakly supported in top models but may only influence dispersal age at high population densities. These results suggest that the proximal factors that trigger dispersal are influenced by a combination of multiple effects related to the costs of dispersing and the benefits of remaining at home, even in species that do not form long-term social groups or have extreme sex-biased dispersal.     

Sociality reduces individual direct fitness in a communally breeding rodent,the colonial tuco-tuco (<Emphasis Type="Italic">Ctenomys sociabilis</Emphasis>)

In many social vertebrates, remaining in the natal group leads to at least short-term reductions in the direct fitness of philopatric animals. Among communally breeding rodents, the direct fitness costs of philopatry appear to increase as the frequency of successful natal dispersal decreases, suggesting a functional link between constraints on natal dispersal and the reproductive consequences of sociality. To explore this relationship empirically, I documented patterns of direct fitness among female colonial tuco-tucos (Ctenomys sociabilis), which are group-living subterranean rodents from southwestern Argentina. Demographic data suggest that successful natal dispersal is rare in this species, leading to the prediction that natal philopatry in C. sociabilis is associated with significant reductions in individual direct fitness. Using data obtained during 1996–2001, I compared the direct fitness of females that dispersed from their natal group and bred alone as yearlings to that of females that lived and bred in their natal group as yearlings. Philopatric yearlings reared significantly fewer young to weaning than did disperser (lone) yearlings. Although neither survival to a second breeding season nor the estimated lifetime number of pups reared to weaning differed between dispersal strategies, the annual direct fitness of group-living females was 23–40% less than expected, suggesting that philopatric animals experienced a substantial direct fitness cost by remaining in their natal group. These data yield important insights into the adaptive bases for group living in C. sociabilis and suggest that constraints on natal dispersal are an important factor favoring group living in this species.Communicated by J. Wilkinson     

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.     

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.     

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     

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.     

Proximate causes of natal dispersal in female yellow-bellied marmots, Marmota flaviventris

We investigated factors influencing natal dispersal in 231 female yearling yellow-bellied marmots (Marmota flaviventris) using comprehensive analysis of 10 years (1983-1993) of radiotelemetry and 37 years (1963-1999) of capture-mark-recapture data. Only individuals whose dispersal status was verified, primarily by radiotelemetry, were considered. Univariate analyses revealed that six of the 24 variables we studied significantly influenced dispersal: dispersal was less likely when the mother was present, amicable behavior with the mother and play behavior were more frequent, and spatial overlap was greater with the mother, with matriline females, and with other yearling females. Using both univariate and multivariate analyses, we tested several hypotheses proposed as proximate causes of dispersal. We rejected inbreeding avoidance, population density, body size, social intolerance, and kin competition as factors influencing dispersal. Instead, our results indicate that kin cooperation, expressed via cohesive behaviors and with a focus on the mother, influenced dispersal by promoting philopatry. Kin cooperation may be an underappreciated factor influencing dispersal in both social and nonsocial species.     

The association of female kin in the arctic ground squirrel Spermophilus parryii

Summary The association of kin in arctic ground squirrels (Spermophilus parryii) was studied near Haines Junction, Yukon Territory, during the summers of 1977, 1978, and 1979. Males dispersed in this population, thus only females were likely to live near relatives.Close female kin (sisters, mothers/daughters) had greater overlap of home ranges and interacted more amicably and less agonistically than did less closely related females. Closely related females clumped their young at emergence, whereas more distantly related females did not; there was little indication of clumping of pre-emerged young. The overlap of home ranges of distant relatives (known genetic relatives that had not associated in a natal burrow) was intermediate to that of close relatives and non-relatives. The types of interactions between distant relatives were more similar to those between non-relatives than between close relatives.I conclude that female arctic ground squirrels exhibit nepotism. Females may benefit from associations with relatives by sharing watching for predators once juveniles become conspicuous. Male arctic ground squirrels commit infanticide and several females may be more effective at protecting their young from infanticidal males than females living alone. I suggest that clumping of young by close relatives may provide a mechanism allowing distantly related females to identify each other.Address for offprint requests