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.     

Spring dispersal and agonistic behaviour of the Red-legged partridge (Alectoris rufa)

Spring behaviour, dispersal and population changes were studied in two populations of individually marked Red-legged partridges.
Young males took up home ranges close to their natal site but young females dispersed away. The incidence of agonistic behaviour between males, the extent of the spring decline in population density and the distance dispersed by males between the winter and breeding ranges were less in an area with more abundant nesting cover.
Half the neighbours of a typical male came from the same winter social group. These groups were based on family parties. Males from the same winter group showed less frequent agonistic behaviour to each other and had more overlapped home ranges than those from different groups.     

Stepping stones of life: natal dispersal in the group-living but noncooperative speckled warbler

In most cooperatively breeding birds the offspring of one sex, usually male, delays dispersal to remain on the natal territory and helps its parents to rear subsequent young. Thus delayed dispersal could be the first step in the evolution of cooperative breeding. We studied natal dispersal in a population of the group-living speckled warbler, Chthonicola sagittata, based on observations of a colour-banded population over 3 years. Unlike other group-living members of the Acanthizinae, all juvenile males in this population dispersed to settle on foreign territories as subordinates, which do not help rear the young. Speckled warblers showed all the life history traits that are thought to result in a saturated habitat and lead to delayed dispersal: they were sedentary, had high adult survival and had a male-biased sex ratio. However, they differed from other acanthizids in occurring at low density (0.18 birds/ha) on large breeding territories (6-12 ha), with a maximum of two males per territory. This may allow subordinates to live on foreign territories yet avoid aggression from dominants. A benefit of dispersal is that it provides an additional route to gaining a breeding vacancy. Dispersers can acquire vacancies on their new territory or on a neighbour's, but incest avoidance would be likely to constrain nondispersing males to neighbours' territories. A model of relative lifetime success showed that the survival benefits of natal philopatry are unlikely to outweigh this benefit of dispersal.     

Endocrine correlates of contrasting male mating strategies in the European badger (Meles meles)

In mammals, testosterone secretion is known to respond rapidly to changes in males' immediate social environment. However, such changes do not take testosterone levels below the baseline required to maintain spermatogenesis, and so do not usually affect males' fertility. In this paper, we show that individual males' patterns of testicular activity reflect their social roles in a group-living carnivore, the European badger ( Melcs metes ), leading to individual and population differences in the seasonal patterns of both testosterone secretion and, apparently, spermatogenesis. In one badger population, some males left their natal groups to become breeding males in neighbouring territories. These males had higher plasma testosterone levels, and appeared to sustain spermatogenesis for a longer period, than males which remained in their natal territories. In contrast, in a second (higher density) population, males rarely transferred away from their natal territories, and appeared not to defend access to groups of females. Instead, males made forays to mate with females in neighbouring territories. In this population males had a shorter period of testicular activity, and there were no differences in testicular activity between immigrant and natal males.     

Sex biased natal dispersal in a closed, saturated population of Seychelles warblers Acrocephalus sechellensis

The distances that individuals disperse, from their natal site to the site of first breeding and between breeding sites, have important consequences for the dynamics and genetic structure of a population. Nearly all previous studies on dispersal have the problem that, because the study area encompassed only a part of the population, emigration may have been confounded with mortality. As a result long-distance dispersers may have been overlooked and dispersal data biased towards short distances. By studying a virtually closed population of Seychelles warblers Acrocephalus sechellensis we obtained almost unbiased results on several aspects of dispersal. As in the majority of other avian species, natal dispersal distance was female biased in the Seychelles warbler. Female offspring also forayed further from the natal territory in search of breeding vacancies than male offspring. The sex bias in natal dispersal distance did, however, depend on local breeding density. In males, dispersal distance decreased as the number of territories bordering the natal territory increased, while in females, dispersal distance did not vary with local density. Dispersal by breeders was rare and, unlike in most species, distances did not differ between the sexes. We argue that our results favour the idea that the sex bias in natal dispersal distance in the Seychelles warbler is due to inbreeding avoidance and not resource competition or intrasexual competition for mates.     

Dispersal,kinship and inbreeding in an island population of the Great Tit

We investigated the kinship structure of an island population of the Great Tit (Parus major). Kinship of birds could be inferred by comparing their family trees. Dispersal was also studied to explain the observed pattern of kinship. On the island of Vlieland the tits breed in several wooded areas. Both males and females preferred to breed in their natal area; males did so more strongly than females. Hence gene flow between the areas is restricted. However, within the largest wooded area females showed random dispersal, while males showed a slight tendency to breed near their natal site. The degree of kinship of neighbouring birds is a suitable control group for the relatedness of partners that takes into account the effects of dispersal. In the largest wooded area, birds were on average equally related to their partner and to their neighbours. Moreover, the mean coefficient of kinship between male and female neighbours was equal to the average kinship in this part of the population. We conclude that mating is random with respect to kinship. There is no evidence for avoidance of inbreeding. It is unlikely that kin recognition plays an important role in the process of mate choice in this population of Great Tits. We suggest that ecological factors are the main causes for the observed patterns of dispersal and mating. On the island more female than male immigrants enter the population each year. Incidental data indicate an exchange of birds between the population studied and surrounding populations. Ancestries of immigrants are not known, and indeed a first analysis of all birds, including immigrants, showed that males were more closely related than females. However, differential immigration could not fully explain the observed difference in kinship. The presence of local adaptation in males is suggested as a possible additional cause.     

Sex-bias and timing of natal dispersal in cooperatively breeding Puff-throated Bulbuls Alophoixus pallidus

While natal dispersal can have a significant impact on population dynamics, it is typically difficult to quantify. We investigated timing of natal dispersal of the cooperatively breeding Puff-throated Bulbul Alophoixus pallidus in a tropical evergreen forest by modelling the probability of staying in or dispersing from their natal territory whilst taking into account the effects of sex, group size, and the presence of helper(s). Birds did not disperse until the beginning of and during the breeding season following the hatching year. Dispersal was strongly female-biased both in frequency and distance: most females (95%) dispersed away from their natal territories, and of those relocated, traversed 2–7 territories. In contrast, 50% of males remained in the natal territory as helpers in their second year, while relocated dispersing males crossed 1–2 territories. Natal dispersal was not influenced by either group size or the presence of helpers. Males that fledged earlier in the breeding season exhibited higher rates of philopatry than the males that fledged later, but no correlation between fledging date and philopatry was observed in females. The probability of staying in the natal territory during the second year was 0.58 ± 0.14 SE and 0.05 ± 0.04 for males and females, respectively. These findings may add to our understanding of how natal dispersal can reflect social patterns and kin structure in cooperative breeding species from a little-studied tropical forest region.     

The Cooperative Breeding System of the Red-cockaded Woodpecker

The breeding system of the red-cockaded woodpecker is described based on data collected over six years from a population of 500 marked individuals in the Sandhills of North Carolina. Male-female pairs were the most common social unit (59%), but 30% of social units contained one or more adult helpers, and 11% consisted of solitary males. Helpers were almost exclusively male: 27% of males remained in their natal group as helpers for at least one year, whereas only four (1%) females did. Most breeding females remained as breeders in the same group from one year to the next (56%), but a surprising number (12%) moved to another group. Many movements were related to incest avoidance or mate death, but 39% involved deserting a mate, usually following successful reproduction. We suggest that females sometimes are forced from groups by immigrants or other group members. The median distance of movements by adult females was only 1.3 km. In contrast to females, no breeding males switched groups. Survival of both breeding (76%) and helper (80%) males was higher than that of breeding females (69%). Males exhibited two distinct life-history strategies. Some remained as helpers on their natal territory for one or more years, and became breeders by inheriting breeding status in the natal group (17% per year) or by replacing a deceased breeder in a nearby group (13% per year, median distance moved 1.0 km). Other males dispersed from their natal group permanently during their first year. Some of these males were floaters at age one year, others were solitary, and a few became helpers in a non-natal group, but many were breeders. In contrast to males that first functioned as helpers, those that dispersed after fledging moved long distances (median dispersal distance 4.5 km), longer even than dispersing female fledglings moved (median distance 3.2 km). The habitat saturation model of the evolution of cooperative breeding is based on selection between the two life-history strategies exhibited by male red-cockaded woodpeckers. The model therefore may be tested directly with this species. Another indication that this model is appropriate for this species is the existence of a resource (cavity trees) that might provide an ecological basis for habitat saturation.     

Microgeographic socio-genetic structure of an African cooperative breeding passerine revealed: integrating behavioural and genetic data

Dispersal can be motivated by multiple factors including sociality. Dispersal behaviour affects population genetic structure that in turn reinforces social organization. We combined observational information with individual-based genetic data in the Karoo scrub-robin, a facultative cooperatively breeding bird, to understand how social bonds within familial groups affect mating patterns, cause sex asymmetry in dispersal behaviour and ultimately influence the evolution of dispersal. Our results revealed that males and females do not have symmetrical roles in structuring the population. Males are extremely philopatric and tend to delay dispersal until they gain a breeding position within a radius of two territories around the natal site. By contrast, females dispersed over larger distances, as soon as they reach independence. This resulted in male neighbourhoods characterized by high genetic relatedness. The long-distance dispersal strategy of females ensured that Karoo scrub-robins do not pair with relatives thereby compensating for male philopatry caused by cooperation. The observed female-biased strategy seems to be the most prominent mechanism to reduce the risk of inbreeding that characterizes social breeding system. This study demonstrates that tying together ecological data, such as breeding status, determining social relationships with genetic data, such as kinship, provides valuable insights into the proximate causes of dispersal, which are central to any evolutionary interpretation.     

Female-biased natal and breeding dispersal in an alpine lizard, Niveoscincus microlepidotus

We measured two aspects of dispersal in the alpine Australian scincid lizard, Niveoscincus micolepidotus : (1) natal dispersal, i.e. shift in home range over the lizard's first year of life, and (2) breeding dispersal, i.e. shifts of home ranges between breeding attempts as adults. On average, displacements were surprisingly small. Female neonates dispersed about twice as far as did males in the same cohort (means of 12 m vs. 6 m). A female's natal dispersal distance was not correlated with her body size or our estimate of physiological performance (sprint speed). However, larger, faster-running male neonates dispersed further than did smaller, slower males. As was the case for neonates, adult females moved significantly further between breeding seasons than did adult males (14.2 m vs. 9.6 m). Because of a female's long gestation period (more than 1 year), two groups of females occur simultaneously in the population, non-ovulated (i.e. with yolking folicles) and pregnant females (i.e. approaching parturition). Females that were not yet ovulated showed a markedly stronger dispersal in response to high reproductive effort (i.e. clutch size in relation to body condition) than did pregnant females. In adult males, body size was negatively correlated with dispersal distance, suggesting that although males have overlapping territories, they exhibit an increasing level of site tenacity with age and/or size. Thus, selection for the relatively more pronounced site tenacity in adult males may have resulted in the more marked philopatric behaviour compared to females also as neonates.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 277–283.     

Great bustard population structure in central Spain: concordant results from genetic analysis and dispersal study.

We found significant sex differences in the mtDNA genetic structure and dispersal patterns of great bustards in a population of 11 breeding groups, "leks", in central Spain. The analysis of genetic distances showed that the female population was divided into three groups of leks separated by ca. 50 km, whereas male haplotypes were randomly distributed among leks. Genetic distances among pairs of leks were positively correlated with geographical distances in females but not in males. While female haplotype distributions were homogeneous among leks at close distances, differences in male genetic structure were highly variable even between two close leks. These results from genetic analyses were concordant with those from a radiotracking study on natal dispersal. Natal dispersal distances were higher in males than in females. Also, the frequency of movement of a female between two leks was positively correlated with their genetic affinity and geographical proximity. In males, the frequency of movement was correlated with geographical proximity but not with genetic affinity. Males dispersed among genetically unrelated leks, contributing to keep nuclear genetic diversity in the population, whereas females tended to be philopatric. These results suggest that isolation-by-distance influences the distribution of maternal lineages at a regional level.     

Influential factors for natal dispersal in an avian island metapopulation

Dispersal is a key parameter in evolutionary, demographic and conservation theory, but the factors influencing dispersal between populations are rarely known, and the contribution of immigrants to population stability remains uncertain. Using dispersal data from nine island populations of song sparrows, we show that female and male immigrants responded differently to population structure: in females, immigration varied with adult sex ratio; whereas immigration by males was more influenced by population density. These patterns are consistent with the hypothesis that intra-sexual competition for breeding resources influenced recruitment patterns. Immigrants often constituted a substantial fraction of local population size, and in six cases immigration by females prevented the extirpation of that sex from the island. Breeding vacancies and extirpations may have been more likely in females because their apparent survival was lower than in males. Local recruitment and immigration varied markedly among islands, perhaps as consequence of island size and isolation. Overall, our results suggest that immigration varied with local demography in a sex-specific way, stabilized population numbers and reduced extinction rates in the smallest populations.     

Role of breeding and natal movements in lifetime dispersal of a forest‐dwelling rodent

The lifetime movements of an individual determine the gene flow and invasion potential of the species. However, sex dependence of dispersal and selective pressures driving dispersal have gained much more attention than dispersal at different life and age stages. Natal dispersal is more common than dispersal between breeding attempts, but breeding dispersal may be promoted by resource availability and competition. Here, we utilize mark–recapture data on the nest‐box population of Siberian flying squirrels to analyze lifetime dispersal patterns. Natal dispersal means the distance between the natal nest and the nest used the following year, whereas breeding movements refer to the nest site changes between breeding attempts. The movement distances observed here were comparable to distances reported earlier from radio‐telemetry studies. Breeding movements did not contribute to lifetime dispersal distance and were not related to variation in food abundance or habitat patch size. Breeding movements of males were negatively, albeit not strongly, related to male population size. In females, breeding movement activity was low and was not related to previous breeding success or to competition between females for territories. Natal philopatry was linked to apparent death of a mother; that is, we did not find evidence for mothers bequeathing territories for offspring, like observed in some other rodent species. Our results give an example of a species in which breeding movements are not driven by environmental variability or nest site quality. Different evolutionary forces often operate in natal and breeding movements, and our study supports the view that juveniles are responsible for redistributing individuals within and between populations. This emphasizes the importance of knowledge on natal dispersal, if we want to understand consequences of movement ecology of the species at the population level.     

Effects of breeding success, mate fidelity and senescence on breeding dispersal of male and female blue-footed boobies

1. Understanding the effects of individual and population factors on variation in breeding dispersal (the movement of individuals between successive breeding sites) is key to identifying the strategies behind breeders' movements. Dispersal is often influenced by multiple factors and these can be confounded with each other. We used 13 years of data on the locations, mates, breeding success and ages of individuals to tease apart the factors influencing breeding dispersal in a colonially breeding long-lived seabird, the blue-footed booby Sula nebouxii. 2. Breeding dispersal varied among and within years. Males dispersed further in years of higher population density, and late breeding males and females dispersed further than early breeders. This temporal variation related to changes in competition for territory was taken into account in all tests of individual factors influencing breeding dispersal. 3. Individuals that retained their mates from the previous year dispersed shorter distances than those that changed their mates. 4. The effect of previous breeding success depended on mate fidelity. Unsuccessful breeding induced greater dispersal in birds that changed their mates but not in birds that retained their mates, indicating that breeders who change mates may take their own previous breeding experience into account during habitat selection. Faithful individuals may have to stay close to their previous sites to encounter their mates. 5. Male divorcees dispersed over shorter distances than their former mates, possibly because males contribute more than females to establishing territories. 6. Dispersal of males and females declined with increasing age over the first 10-11 years of life, then increased in old age, possibly due to senescent decay in the ability to compete for mates and territories.     

Spatial structuring and dispersal in a high density population of the California vole Microtus californicus

Space use patterns of California voles are described from intensive live-trapping data obtained during the late breeding season in a peak density year. Operational sex ratios were strongly female biased. Breeding males had mutually exclusive home ranges that overlapped the home ranges of one to several females. The average male home range was larger than the average female home range and tended to encompass female home ranges in their entirety. Breeding females had home ranges that often overlapped extensively with the home ranges of other breeding females. The degree of overlap, however, tended to be either very high or very low, indicating that groups of females may act territorially among themselves. Family groups that may include members of sequential litters formed at high density. Males on experimental removal grids had larger home ranges than males on the high density grids and some became wanderers while most females had home ranges comparable to those on the high density grids.
Dispersers onto low density grids were not a random sample of the high density population; large breeding adults and small juveniles were under-represented. No sex differences in dispersal were noted. Immigration into the high density populations was probably negligible.     

Reverse sex-biased philopatry in a cooperative bird: genetic consequences and a social cause

The genetic structure of a group or population of organisms can profoundly influence the potential for inbreeding and, through this, can affect both dispersal strategies and mating systems. We used estimates of genetic relatedness as well as likelihood-based methods to reconstruct social group composition and examine sex biases in dispersal in a Costa Rican population of white-throated magpie-jays ( Calocitta formosa , Swainson 1827), one of the few birds suggested to have female-biased natal philopatry. We found that females within groups were more closely related than males, which is consistent with observational data indicating that males disperse upon maturity, whereas females tend to remain in their natal territories and act as helpers. In addition, males were generally unrelated to one another within groups, suggesting that males do not disperse with or towards relatives. Finally, within social groups, female helpers were less related to male than female breeders, suggesting greater male turnover within groups. This last result indicates that within the natal group, female offspring have more opportunities than males to mate with nonrelatives, which might help to explain the unusual pattern of female-biased philopatry and male-biased dispersal in this system. We suggest that the novel approach adopted here is likely to be particularly useful for short-term studies or those conducted on rare or difficult-to-observe species, as it allows one to establish general patterns of philopatry and genetic structure without the need for long-term monitoring of identifiable individuals.     

Dispersal Status Influences Hormones and Behavior in the Male Spotted Hyena

Male spotted hyenas (Crocuta crocuta) reach puberty at 24 months of age and then invariably emigrate from their natal clans 1 to 38 months later. Thus there are two classes of reproductively mature males in everyCrocutaclan: adult natal males born in the clan and adult immigrant males born elsewhere. In one free-living hyena population in Kenya, these two groups of males were compared with respect to measures of aggression, social dominance, sexual behavior, and circulating hormone levels. Adult natal males engaged in higher hourly rates of aggression than did immigrants, won all fights with immigrants, and were socially dominant to immigrants. In addition, adult natal males engaged in far lower hourly rates of sexual behavior with resident females than did immigrants, and natal males were never observed to copulate with natal females. Mean basal plasma cortisol values did not differ between the two groups of adult males, but cortisol concentrations in immigrants were positively correlated with tenure in the clan and with immigrant male social rank. Adult natal males had plasma testosterone levels significantly lower than those of immigrants. Social rank and plasma testosterone values were positively correlated among immigrant males. Thus two different relationships appear to exist between circulating testosterone and social rank in maleCrocuta:one apparent in immigrants and the other in natal adult males. Our results suggest that dispersal might disinhibit testosterone secretion in postpubertal male hyenas.     

Dispersal patterns in a medium‐density Irish badger population: Implications for understanding the dynamics of tuberculosis transmission

European badgers (Meles meles) are group‐living mustelids implicated in the spread of bovine tuberculosis (TB) to cattle and act as a wildlife reservoir for the disease. In badgers, only a minority of individuals disperse from their natal social group. However, dispersal may be extremely important for the spread of TB, as dispersers could act as hubs for disease transmission. We monitored a population of 139 wild badgers over 7 years in a medium‐density population (1.8 individuals/km²). GPS tracking collars were applied to 80 different individuals. Of these, we identified 25 dispersers, 14 of which were wearing collars as they dispersed. This allowed us to record the process of dispersal in much greater detail than ever before. We show that dispersal is an extremely complex process, and measurements of straight‐line distance between old and new social groups can severely underestimate how far dispersers travel. Assumptions of straight‐line travel can also underestimate direct and indirect interactions and the potential for disease transmission. For example, one female disperser which eventually settled 1.5 km from her natal territory traveled 308 km and passed through 22 different territories during dispersal. Knowledge of badgers' ranging behavior during dispersal is crucial to understanding the dynamics of TB transmission, and for designing appropriate interventions, such as vaccination.     

Dispersal strategies in Tasmanian native hens (Gallinula mortierii)

Individuals in cooperatively breeding species face a complexset of decisions when they reach reproductive maturity. Duringan 8-year study, we examined the histories of 214 Tasmaniannative hens (Gallinula mortierii) from hatching to examinethe strategies they used to acquire breeding positions andthe reproductive success they experienced in those breedingpositions. Two-thirds of young delayed dispersal from theirnatal groups for at least a year. Ecological constraints werea partial cause of delayed dispersal; high-quality territorieswere rare and remained occupied due to high adult survivorship.There were also clear benefits of philopatry. Individuals thatinherited breeding positions on their natal territories gainedbetter quality positions and experienced higher reproductivesuccess in their first breeding attempts than did individualswho dispersed to other groups. Multivariate analyses showedthat the method of acquisition of breeding positions was theonly factor significantly related to the quality of the breedingpositions attained. Males were more likely to inherit breeding positions in their natal groups than were females. The compositionsof individuals' natal groups had no effect on whether theyinherited breeding positions or dispersed. In contrast, thecompositions of groups did appear to affect whether other birdsdispersed into them, with birds rarely moving into groups thatcontained breeders or nonbreeders of the same sex as the potential dispersers. Short-term removals of breeders confirmed this finding.These results suggest that both ecological constraints andbenefits of philopatry explain delayed dispersal in this species.     

Behaviour and breeding biology of the cooperatively breeding Grey-backed Fiscal Shrike Lanius excubitorius in Kenya

Grey-backed Fiscal Shrikes Lanius excubitorius were studied over a 2j year period near Lake Naivasha, Kenya. Grey-backs are cooperative breeders, with group sizes ranging from two to II. Only one pair breeds per group, with all other group members aiding in the rearing of young. The study population ranged from 64 to 79 individuals that occurred in from 13 to 16 groups. Non-breeding helpers made up to 66% of the population, with male helpers being more numerous overall than females. The annual survival rate was 65%, with no differences detected between the survival of males and females, or of breeders and helpers. Only male helpers were observed to acquire breeding status within the natal territory. Some female helpers acquired breeding positions in territories adjacent to their natal territories. Group territorial displays occurred throughout the year but were most pronounced prior to breeding during rainy periods. Reproductive success was very low, with only 14.5% of the recorded breeding attempts leading to fledged young. Large groups (four or more birds) had greater reproductive success than small groups, but many factors other than, or in addition to, group size may have influenced this pattern. The breeding male contributed the most food to the incubating female and to the nestlings. Male helpers and the breeding female contributed more to nestlings than did female helpers. Observations on the post-fledging period indicate that socialization and establishment of dominance may be of importance in group-living species living in a restricted ecological and social setting.