Dispersal, interpatch movements, and survival in a shrubland breeding bird community

ABSTRACT Dispersal events can affect the distribution, abundance, population structure, and gene flow of animal populations, but little is known about long‐distance movements due to the difficulty of tracking individuals across space. We documented the natal and breeding dispersal of shrubland birds among 13 study sites in a 1000 km² area in southeastern Ohio. In addition, we radio‐marked and tracked 37 adult males of one shrubland specialist, the Yellow‐breasted Chat (Icteria virens). We banded 1925 juveniles and 2112 adults of nine shrubland species from 2002 to 2005. Of these, 33 (1.7%) juveniles were encountered in subsequent years (2003–2006) as adults (natal dispersal) and 442 (20.9%) birds initially banded as breeding adults were re‐encountered in subsequent years (breeding dispersal). Apparent survival of juvenile shrubland birds on their natal patches was 0.024 (95% CI 0.016–0.036). After accounting for the probability of detection, we found that 21% of birds banded as juveniles and recaptured as adults returned to their natal patches, whereas 78% of adult birds showed fidelity to the patch where they were originally captured. Moreover, natal dispersers tended to move farther than breeding dispersers (corrected natal median = 1.7 km ± 0.37; corrected breeding median = 0.23 km ± 0.10). We used our estimates of natal dispersal and annual apparent survival to estimate true survival at 0.11 (95% CI 0.07–0.18) for juveniles in their first year. However, this estimate was only applicable for birds dispersing within 7 km of their natal patches. Interpatch movements of radio‐marked Yellow‐breasted Chats were not uncommon, with 13 of 37 males located in more than one habitat patch. Overall, we observed low natal philopatry, but high adult site fidelity for shrubland birds in our study area. Considering the frequency of short‐distance movements observed (median = 531 m, range = 88–1045 m), clustering of patches within 1 km might facilitate use of shrubland habitat.     

Affinity for natal environments by dispersers impacts reproduction and explains geographical structure of a highly mobile bird

Understanding dispersal and habitat selection behaviours is central to many problems in ecology, evolution and conservation. One factor often hypothesized to influence habitat selection by dispersers is the natal environment experienced by juveniles. Nonetheless, evidence for the effect of natal environment on dispersing, wild vertebrates remains limited. Using 18 years of nesting and mark–resight data across an entire North American geographical range of an endangered bird, the snail kite (Rostrhamus sociabilis), we tested for natal effects on breeding-site selection by dispersers and its consequences for reproductive success and population structure. Dispersing snail kites were more likely to nest in wetlands of the same habitat type (lacustrine or palustrine) as their natal wetland, independent of dispersal distance, but this preference declined with age and if individuals were born during droughts. Importantly, dispersing kites that bred in natal-like habitats had lower nest success and productivity than kites that did not. These behaviours help explain recently described population connectivity and spatial structure across their geographical range and reveal that assortative breeding is occurring, where birds are more likely to breed with individuals born in the same wetland type as their natal habitat. Natal environments can thus have long-term and large-scale effects on populations in nature, even in highly mobile animals.     

Philopatry and natal dispersal in a sedentary population of western marsh harrier

Dispersal is a key life-history trait because it influences population dynamics and population genetic structure. From a behavioural perspective, the study of natal dispersal requires some understanding of the mechanisms that affect individual movements, because movements of an animal form a path that is continuous throughout its life. Our aim was to investigate juvenile dispersal strategies in the western marsh harrier Circus aeruginosus , between fledging and first breeding attempt (from 1 to 4 years later, depending on the sex and individual). Using radio tracking, we monitored dispersing juvenile harriers and their home-range size variations within a sedentary population in central western France from 2001 to 2007. Juvenile dispersal strategy was mainly characterized by a very high natal philopatry (i.e. birds that remained within the study area) and short-distance dispersal. All but one bird (out of 39) remained within the study area between their first winter and their first spring, and 96.0% during their first spring. The distance moved at 2 years of age was significantly larger for males than for females (3800± sd 3593 m and 935± sd 481 m for seven males and six females, respectively), in contrast to most bird species studied so far. Home-range size was not sex biased and significantly decreased with age. In addition, non-breeding birds had larger home ranges (1603± sd 2128 ha) than breeders (349± sd 185 ha). Using data obtained from other populations, juvenile marsh harrier dispersal strategies appeared to be determined by migratory status (migratory birds dispersing farther) and demographic parameters (juvenile survival or fecundity).     

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.     

Female-biased natal dispersal in the Siberian flying squirrel

Natal dispersal is usually sex biased in birds and mammals.Female-biased natal dispersal is the prevailing pattern in birdsbut is rare among mammals. Hypotheses explaining sex bias indispersal include the mate-defense mating hypothesis, whichpredicts male-biased dispersal, the resource-defense hypothesispredicting female-biased dispersal, and the competition hypothesis,which predicts that if dispersal is caused by competition forresources between sexes, then the subdominant sex will disperse.We studied natal dispersal of Siberian flying squirrels Pteromysvolans using radio telemetry in Southern Finland in 1996–2004.Of 86 juveniles that survived over the dispersal period, almostall young females dispersed from the natal site, whereas almost40% of males were philopatric. Dispersal was farther for femalesthan males. Females began dispersal on average 2 weeks earlierthan males and were lighter in mass at the onset of dispersalthan later dispersing males. No mate- or resource-defense matingsystem could be found among males, but females seemed to defendnest and apparently food resources, in contrast to the expectationof dispersal bias in resource-defense systems. Competition forresources between sexes does not explain female bias either:in the flying squirrel, the female seems to be the dominantsex. We propose that young females are subordinate to theirmothers and have to disperse to find a vacant, suitable sitefor reproduction.     

Lifetime philopatry in the blue-footed booby: a longitudinal study

Philopatry over the lifetime and its relationship with reproductivesuccess were examined using longitudinal records of nest locationand reproduction of individual blue-footed boobies. Males showedshorter natal dispersal than females, and natal dispersal distanceof both sexes were unrelated to either first reproductive successor lifetime reproductive success. Throughout the early lifetime,males and females nested closer to their first breeding sitesthan to their natal sites, and comparison with a simulationof successive breeding dispersals in random directions showedthat male and female blue-footed boobies are philopatric tothe first breeding site. Therefore, throughout the early lifetime,the first breeding site seems to function as a point of referencefor breeding site use together with the previous season's site.Males and females with shorter natal dispersal distances showedstronger lifetime philopatry to their first breeding sites,suggesting stable individual variation in competitive abilityor dispersal phenotype. However, early lifetime philopatry tofirst breeding sites was unrelated to annual breeding success.Compared with simple fidelity to previous breeding sites, lifetimephilopatry to first breeding sites should result in increasedkin interactions and greater selection for kin recognition,altruism and inbreeding avoidance, as well as long-term familiaritywith neighbors.     

Dispersal polymorphisms from natal phenotype–environment interactions have carry‐over effects on lifetime reproductive success of a tropical parrot

We examined how interactions between an individual's phenotype and its environment affect natal dispersal at multiple scales and the effects on lifetime reproductive success using a 22‐year study of green‐rumped parrotlets (Forpus passerinus). Dispersal increased or decreased lifetime reproductive success depending upon an individual's natal environment and phenotype. Many of the phenotypic traits and environmental conditions that influenced lifetime reproductive success also influenced dispersal, such as clutch size and competition, and this differed with scale. By examining phenotype–environment interactions, we observed both positive and negative effects of rainfall, habitat quality and competition on dispersal depending upon phenotype. The dispersal behaviours of juveniles typically resulted in higher lifetime reproductive success. Thus, individuals commonly exhibit ideal free behaviour and results provide support for the occurrence and maintenance of dispersal polymorphisms. This study highlights the long‐term, carry‐over effects of natal environment, natal phenotype and dispersal tactic on lifetime reproductive success.     

Seasonal variation of reproductive success under female philopatry and male-biased dispersal in a common vole population

Variation of reproductive success, an important determinant of the opportunity for sexual selection, is an outcome of competition within one sex for mating with members of the other sex. In promiscuous species, males typically compete for access to females, and their reproductive strategies are strongly related to the spatial distribution of females. I used 10 microsatellite loci and the mtDNA control region to determine seasonal differences in the reproductive success of males and females of the common vole (Microtus arvalis), one of the most numerous mammals in Europe. The sex-related spatial structure and bias in dispersal between genders were also assessed. Standardized variance of the reproductive success of females did not vary seasonally due to the continuity of female philopatry throughout the breeding season and to the constancy of the number of females reproducing successfully in each season. The males are the dispersing sex, undergoing both natal and breeding dispersal. Their standardized variance of reproductive success was significantly higher than that for females in July, when only two males monopolized 80% of the females in the population and when variance of male reproductive success was highest (I_m = 7.70). The seasonally varying and high standardized variance of male reproductive success may be explained by male-male competition for matings, coupled with seasonal changes in the age structure of the population.     

Mate and territory availability affect breeding dispersal and divorce in a resident Southern House Wren Troglodytes aedon musculus population

Breeding dispersal is the movement of an individual between breeding attempts and is usually associated with the disruption of the social pair bond, although mates may disperse together as a social unit. In monogamous territorial species, the decision to disperse may be affected by individual attributes such as sex, age and condition of the disperser. However, environmental and social contexts may also play a crucial role in the decision to disperse. We analysed capture‐resighting data collected over 9 years to study breeding dispersal and divorce rates of a Southern House Wren Troglodytes aedon musculus population in South Temperate Argentina. Between‐season dispersal was more frequent than within‐season dispersal, with females dispersing more often than males, both between and within seasons. Both within‐season and between‐season breeding dispersal probability was affected by territory availability, but not by previous breeding success. When the adult sex ratio (ASR) was more skewed towards males, male between‐season dispersal was also affected by mating status, with widowed and single males dispersing more often than paired males. Within‐season divorce increased the reproductive success of females but not males, and was affected by the availability of social partners (with increasingly male‐skewed ASR). Our results suggest that territorial vacancies and mating opportunities affect dispersal and divorce rates in resident Southern House Wrens, highlighting the importance of social and environmental contexts for dispersal behaviour and the stability of social pair bonds.     

Natal dispersal patterns are not associated with inbreeding avoidance in the Seychelles warbler

In this study, we test whether patterns of territory inheritance, social mate choice and female-biased natal dispersal act as inbreeding avoidance mechanisms in the cooperatively breeding Seychelles warbler. Our results show that Seychelles warblers do not reduce the likelihood of inbreeding by avoiding related individuals as mates. The occurrence of natural and experimentally induced territory inheritance did not depend on whether the remaining breeder was a parent of the potential inheritor or an unrelated breeder. Furthermore, dispersing individuals were no less related to their eventual mates than expected given the pool of candidates they could mate with. The female bias in natal dispersal distance observed in the Seychelles warbler does not facilitate inbreeding avoidance because, contrary to our prediction, there was no sex difference in the clustering of related opposite sex breeders around the natal territories of dispersers. As a result, the chance of females mating with relatives was not reduced by their greater dispersal distance compared with that of males.     

Male Seychelles warblers use territory budding to maximize lifetime fitness in a saturated environment

In cooperatively breeding species, helping at the nest and buddingoff part of the natal territory have been advanced as strategiesto increase fitness in an environment that is saturated withterritories. The importance of helping or territory buddingas a determinant of lifetime reproductive success (LRS) hasbeen debated because the potential benefits of both strategiescould not be separated. Here we test the causes and the immediateand future fitness consequences of single dispersal decisionstaken by male Seychelles warblers (Acrocephalus sechellensis).Males breeding in high-quality territories (high food abundance)have significantly higher LRS than similar-aged males buddingoff part of the parental territory. Initially, budders havea low reproductive success (because of limited food resourcesor absence of a breeding partner). However, they have a longlife span and inherit high-quality territories through sitedominance, by which they gain higher LRS than breeders on low-qualityterritories, helpers, or floaters. Experimental creation ofmale breeding territory vacancies showed that most young malesbecame budders because of intense competition for high-quality territories. The translocation of warblers to the previouslyunoccupied Aride Island shows that males behave according tothe expected fitness benefits of each dispersal strategy. Inthe absence of competition for territories on Aride, all youngmales bred in high-quality territories. However, after saturationof high-quality habitat with territories, most males becamebudders rather than breeders on low-quality habitat, helpers,or floaters.     

Immigration and gene flow in a northern willow tit (Parus montanus) population

To study quantitatively the relationship between immigration rate and gene flow we used 9-year data from a willow tit Parus montanus population in a continuous forest habitat. We compared components of lifetime reproductive success, and parental survival rate between immigrant and resident (‘status’) birds by taking individual age into consideration also. Of the fitness components, survival was independent of status and sex, averaging 0.60 annually. Of the male and female breeders, on average 63.1% and 75.6%, respectively, originated from unknown natal areas, implying extensive immigration. Pair formation was nonassortative with respect to the origin of partners. The effect of status on reproductive success was significant only for females: immigrants produced larger clutches and hatched more young in all age classes studied. However, the difference between the groups diminished until fledging and the offspring produced by the immigrant females showed lower local survival rate (4.6% of the young) than did those by the residents (5.9%). Therefore, the contribution of immigrant females to the local gene pool was lower than expected on the basis of immigration rate. However, we propose that the result implies differential propensities to long-distance dispersal rather than overall survival prospects, since we detected no quality differences suggesting reduced survival chances among descendants of immigrant females. Therefore, gene flow into the population was slightly lower than immigration rate. It is possible that immigrant females are more prone to invest in progeny that effectively disperse further to search for new vacancies. The conditions prevailing during nestling growth may be crucial in determining whether an individual will later become a resident or leave the natal area, but the genetic component of this trait should also be considered. We suggest that widespread propagules is a better strategy than philopatry for a short-lived species to minimize the risk of losing all descendants in temporally and spatially varying conditions.     

Costs and benefits of female-biased natal philopatry in the common goldeneye

Sex-biased natal dispersal in long-lived species may resultin interactions between parents and mature young of the philopatricsex. To investigate the evolutionary basis of natal philopatryin a noncooperative species, the common goldeneye Bucephalaclangula, we studied possible costs and benefits of simultaneousbreeding of females and philopatric daughters. We did not find any fitness consequences of a daughter's breeding on their mother'sbreeding in terms of nest-site selection, body weight, clutchsize, hatching date, or hatching success. Our results, therefore,did not support the assumption of the local resource competitionhypothesis, that the natally philopatric sex should be morecostly to a breeding parent. As possible benefits for daughters returning to their natal area, we tested inheritance of nestsites from mothers and explored whether daughters utilize thepresence of their mother by parasitically sneaking into hermother's nest. Daughters' nest-site selection was not associatedwith the presence of their mothers. A comparison between daughtersand control females revealed that daughters chose their nestsite closer to their natal nest than expected by nest-siteavailability alone. Daughters could not expect to inherit anest site from their mother, and we did not find other indicationsof cooperation between relatives either. The mother's clutchsize did not increase in the year breeding with the daughter, indicating daughters do not parasitize their mother's nest.We suggest that benefits such as decreased nest predation riskassociated with nesting close to the natal nest site may beimportant in the natal philopatric behavior of the species.     

Reproductive skew among males in a female-dominated mammalian society

The purpose of this study was to document patterns of reproductiveskew among male spotted hyenas (Crocuta crocuta), a speciesin which many normal mammalian sex roles are reversed. We usedpaternity determined from 12 microsatellite markers togetherwith demographic and behavioral data collected over 10 yearsfrom a free-living population to document relationships among reproductive success (RS), social rank, and dispersal statusof male hyenas. Our data suggest that dispersal status andlength of residence are the strongest determinants of RS. Natalmales comprise over 20% of the adult male population, yet theysire only 3% of cubs, whereas immigrants sire 97%. This reproductiveadvantage to immigrants accrues despite the fact that immigrants are socially subordinate to all adult natal males, and it providesa compelling ultimate explanation for primary dispersal inthis species. High-ranking immigrants do not monopolize reproduction,and tenure accounts for more of the variance in male reproductivesuccess than does social rank. Immigrant male hyenas rarelyfight among themselves, so combat between rivals may be a relativelyineffectual mode of sexual selection in this species. Instead,female choice of mates appears to play an important role in determining patterns of paternity in Crocuta. Our data supporta "limited control" model of reproductive skew in this species,in which female choice may play a more important role in limitingcontrol by dominant males than do power struggles among males.     

Costs and benefits of natal dispersal in yearling mallards Anas platyrhynchos

Costs and benefits of natal dispersal have not been fully evaluated in birds. We compared timing of breeding and nesting success for yearling female mallards Anas platyrhynchos returning to or dispersing from their natal areas. Information about natal origins was discerned with feather‐isotopes and combined with detailed reproductive histories for 503 radio‐marked females monitored at 16 study sites across the Canadian Aspen Parklands, during 1993–2000. A natal origin assignment model based on feather‐ δ³⁴S, δD, δ¹⁵N, and δ¹³C values correctly assigned 81% (112 of 138) of known‐source yearlings to region of origin; region‐specific rates ranged from 70–90%. Timing of breeding and nesting success was not related to whether or not a female had dispersed from its region of natal origin in Aspen Parkland (i.e. short‐distance dispersal) versus the southern prairies or boreal forest regions (i.e. long‐distance dispersal). Rather, nesting success was best modeled to include effects of site‐specific wetland and breeding pair abundances and an interaction between local breeding pair and wetland densities. Nest success performance relative to dispersal distance varied among study sites but was unrelated to local upland nest cover, wetland habitat conditions, or conspecific density. Thus, we detected no strong costs of dispersal but some evidence that long‐distance dispersers presumably benefitted when they were able to acquire better nest sites.     

Banded mongooses avoid inbreeding when mating with members of the same natal group

Inbreeding and inbreeding avoidance are key factors in the evolution of animal societies, influencing dispersal and reproductive strategies which can affect relatedness structure and helping behaviours. In cooperative breeding systems, individuals typically avoid inbreeding through reproductive restraint and/or dispersing to breed outside their natal group. However, where groups contain multiple potential mates of varying relatedness, strategies of kin recognition and mate choice may be favoured. Here, we investigate male mate choice and female control of paternity in the banded mongoose (Mungos mungo), a cooperatively breeding mammal where both sexes are often philopatric and mating between relatives is known to occur. We find evidence suggestive of inbreeding depression in banded mongooses, indicating a benefit to avoiding breeding with relatives. Successfully breeding pairs were less related than expected under random mating, which appeared to be driven by both male choice and female control of paternity. Male banded mongooses actively guard females to gain access to mating opportunities, and this guarding behaviour is preferentially directed towards less closely related females. Guard–female relatedness did not affect the guard's probability of gaining reproductive success. However, where mate‐guards are unsuccessful, they lose paternity to males that are less related to the females than themselves. Together, our results suggest that both sexes of banded mongoose use kin discrimination to avoid inbreeding. Although this strategy appears to be rare among cooperative breeders, it may be more prominent in species where relatedness to potential mates is variable, and/or where opportunities for dispersal and mating outside of the group are limited.     

Influence of helping and breeding experience on reproductive performance in the Seychelles warbler: a translocation experiment

Reproductive success of the cooperative breeding Seychelleswarbler (Acrocephalus sechellensis) increases with age. Thisage effect is not due to differential survival or increasedreproductive effort, but to accumulated helping and breedingexperience. In their first year of breeding, reproductive performanceof inexperienced warblers with neither helping nor breedingexperience was significandy lower than that of warblers of thesame age with either previous helping or breeding experience.Reproductive performance was the same for primiparae with helpingexperience and for birds with breeding experience. Female primiparaewith helping experience or breeding experience built betternests and spent more time incubating than inexperienced females,which led to increased hatching success. Male primiparae withhelping experience or males with breeding experience guardedthe clutch better than inexperienced males, which led to reducedegg predation. Even-aged warblers with different previous experienceswere transferred to unoccupied islands, where birds startedbreeding immediately in high-quality territories. The experimentshowed that birds with helping experience produced their firstfledgling as fast as experienced breeders, and significandyfaster than inexperienced birds. Breeding performance did notimprove further with experience after the first successful breedingattempt. Only birds with previous breeding experience who pairedwith inexperienced birds, were likely to change mate. The otherpair combinations remained stable. Thus, primiparous birds withhelping experience have greater lifetime reproductive successthan inexperienced primiparae of the same age. This experimentshows that helping behavior has not only been selected for inthe context of promoting an individual's indirect fitness, butalso in the context of gaining helping experience which translatesinto improved reproductive success when a helper becomes a breeder.[Behav Ecol 7: 326-333 (1996)]     

Early age at first breeding and high natal philopatry in the Red‐necked Nightjar Caprimulgus ruficollis

Recruitment rates, natal philopatry and the onset of breeding activity are documented for the first time for the order Caprimulgiformes. Of 171 Red‐necked Nightjars Caprimulgus ruficollis ringed as fledglings in southwestern Spain between 2008 and 2011, 31 (18%) were later recovered. Females tended to disperse slightly further (680 m) than males (570 m), and no individual from the study site or nearby locations was ever recovered outside the area where it hatched, suggesting high natal philopatry. Most males (94%) and females (73%) recruited into the breeding population in their first year of life, and only a few individuals were not recovered until their second (10%) or third (7%) year of life. In contrast to most long‐lived birds, nearly all (95.5%) Red‐necked Nightjars started to reproduce in their first potential breeding season.     

The disperser dilemma in cooperatively breeding birds

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

Parasitism, host immune defence and dispersal

Host-parasite interactions have been hypothesized to affect the evolution of dispersal by providing a possibility for hosts to escape debilitating parasites, and by influencing the level of local adaptation. We used a comparative approach to investigate the relationship between a component of host immune function (which reflects the evolutionary history of parasite-induced natural selection) and dispersal in birds. We used a sample of 46 species of birds for which we had obtained field estimates of T-cell response for nestlings, mainly from our own field studies in Denmark and Spain. Bird species with longer natal, but not with longer breeding dispersal distances had a stronger mean T-cell-mediated immune response in nestlings than species with short dispersal distances. That was also the case when controlling for the potentially confounding effect of migration from breeding to wintering area, which is known from previous studies to be positively associated with dispersal distance. These relationships held even when controlling for similarity among species because of common ancestry. Avian hosts with a larger number of different breeding habitats had weaker mean T-cell-mediated immune responses than habitat specialists. This relationship held even when controlling for similarity among species because of common ancestry. Therefore, T-cell-mediated immunity is an important predictor of evolutionary changes in dispersal ability among common European birds.