Integrating across life-history stages: consequences of natal habitat effects on dispersal

Ecological and evolutionary processes are affected by forces acting at both local and regional scales, yet our understanding of how these scales interact has remained limited. These processes are fundamentally linked through individuals that develop as juveniles in one environment and then either remain in the natal habitat or disperse to new environments. Empirical studies in a diverse range of organisms have demonstrated that the conditions experienced in the natal habitat can have profound effects on the adult phenotype. This environmentally induced phenotypic variation can in turn affect the probability that an individual will disperse to a new environment and the ecological and evolutionary impact of that individual in the new environment. We synthesize the literature on this process and propose a framework for exploring the linkage between local developmental environment and dispersal. We then discuss the ecological and evolutionary implications of dispersal asymmetries generated by the effects of natal habitat conditions on individual phenotypes. Our review indicates that the influence of natal habitat conditions on adult phenotypes may be a highly general mechanism affecting the flow of individuals between populations. The wealth of information already gathered on how local conditions affect adult phenotype can and should be integrated into the study of dispersal as a critical force in ecology and evolution.     

Natal habitat effects drive density—dependent scaling of dispersal decisions

The dispersal behavior of a species is critical for the stability and persistence of its populations across a landscape. How population density affects dispersal decisions is important for predicting these dynamics, as the form of density‐dependent dispersal influences the stability and persistence of populations. Natal habitat experience often has strong impacts on individual dispersal behavior as well, but its influence on density‐dependent dispersal behaviors remains unexplored. Here we address this conceptual gap in two experiments separately examining habitat selection and emigration from recently colonized patches for two species of flour beetle Tribolium sp. We found that interactions between the quality of habitat experienced during natal development and current habitat for dispersal capable adults can strongly affect the form of density dependence, including reversing the direction of nonlinearities (accelerating to decelerating), or even negating the influence of population density for individual dispersal decisions. Across heterogeneous landscapes, where individuals from different populations may experience different natal habitats, this altering of density‐dependent relationships is predicted by theory to fundamentally influence regional population dynamics. Our results indicate that species which occur across heterogeneous environments, such as during conservation reintroductions, or as invasive species spread, have much potential for natal experience to interact with density dependence and influence local and regional population dynamics.     

Genetic consequences of natal dispersal in the colonial lesser kestrel

Dispersal is a life-history trait that plays a fundamental role in population dynamics, influencing evolution, species distribution, and the genetics and structure of populations. In spite of the fact that dispersal has been hypothesized to be an efficient behavioural mechanism to avoid inbreeding, the expected relationship between dispersal and mate relatedness still remains controversial. Here, we examine the genetic consequences of natal dispersal, namely the higher chance of obtaining genetically less similar mates as a result of moving from natal to breeding sites, in a lesser kestrel (Falco naumanni) population. Relatedness between individuals tended to decrease with distance between their breeding colonies, indicating that the study population follows an 'isolation-by-distance' pattern of spatial genetic structure. Such a fine-scale genetic structure generates a scenario in which individuals can potentially increase the chance of obtaining genetically less similar mates by dispersing over larger distances from their natal colony. Using dispersal information and genotypic data, we showed that mate relatedness decreased with natal dispersal distance, an effect that remained significant both while including and excluding philopatric individuals from the data set. These results, together with the well known detrimental consequences of reduced genetic diversity in the study population, suggest that dispersal may have evolved, at least in part, to avoid the negative fitness consequences of mating with genetically similar individuals.     

Annual variation in long‐distance dispersal driven by breeding and non‐breeding season climatic conditions in a migratory bird

Long‐distance dispersal is a fundamental process in ecology and evolution but the factors that influence these movements remain poorly understood in most species. We used stable hydrogen isotopes to quantify the rate and direction of long‐distance immigration in a breeding population of American redstarts and to test whether the settlement decisions that result in long‐distance dispersal are driven by habitat saturation or by the phenology of breeding‐season resources. Our results provide evidence that both natal dispersal and breeding dispersal were influenced by the timing of breeding‐season phenology, with both age classes more likely to disperse north in years when the onset of breeding‐season phenology occurs earlier than normal. Yearlings were also more likely to disperse north following winters with poor habitat quality on their non‐breeding grounds, demonstrating that carry‐over effects from the non‐breeding season influence natal dispersal in this species. Collectively, these results are consistent with the hypothesis that American redstarts use the phenology of breeding season resources as a cue to select breeding sites. Our results suggest that long‐distance dispersal may allow individuals to rapidly respond to advancing phenology caused by global climate change, though their ability to do so may be constrained by long‐term decline in habitat quality predicted for their tropical non‐breeding grounds.     

The evolution of an 'intelligent' dispersal strategy: biased, correlated random walks in patchy landscapes

Theoretical work exploring dispersal evolution focuses on the emigration rate of individuals and typically assumes that movement occurs either at random to any other patch or to one of the nearest‐neighbour patches. There is a lack of work exploring the process by which individuals move between patches, and how this process evolves. This is of concern because any organism that can exert control over dispersal direction can potentially evolve efficiencies in locating patches, and the process by which individuals find new patches will potentially have major effects on metapopulation dynamics and gene flow. Here, we take an initial step towards filling this knowledge gap. To do this we constructed a continuous space population model, in which individuals each carry heritable trait values that specify the characteristics of the biased correlated random walk they use to disperse from their natal patch. We explore how the evolution of the random walk depends upon the cost of dispersal, the density of patches in the landscape, and the emigration rate. The clearest result is that highly correlated walks always evolved (individuals tended to disperse in relatively straight lines from their natal patch), reflecting the efficiency of straight‐line movement. In our models, more costly dispersal resulted in walks with higher correlation between successive steps. However, the exact walk that evolved also depended upon the density of suitable habitat patches, with low density habitat evolving more biased walks (individuals which orient towards suitable habitat at quite large distances from that habitat). Thus, low density habitat will tend to develop individuals which disperse efficiently between adjacent habitat patches but which only rarely disperse to more distant patches; a result that has clear implications for metapopulation theory. Hence, an understanding of the movement behaviour of dispersing individuals is critical for robust long‐term predictions of population dynamics in fragmented landscapes.     

Natal dispersal and personalities in great tits (Parus major)

Dispersal is a major determinant of the dynamics and genetic structure of populations, and its consequences depend not only on average dispersal rates and distances, but also on the characteristics of dispersing and philopatric individuals. We investigated whether natal dispersal correlated with a predisposed behavioural trait: exploratory behaviour in novel environments. Wild great tits were caught in their natural habitat, tested the following morning in the laboratory using an open field test and released at the capture site. Natal dispersal correlated positively with parental and individual exploratory behaviour, using three independent datasets. First, fast-exploring parents had offspring that dispersed furthest. Second, immigrants were faster explorers than locally born birds. Third, post-fledging movements, comprising a major proportion of the variation in natal dispersal distances, were greater for fast females than for slow females. These findings suggest that parental behaviour influenced offspring natal dispersal either via parental behaviour per se (e.g. via post-fledging care) or by affecting the phenotype of their offspring (e.g. via their genes). Because this personality trait has a genetic basis, our results imply that genotypes differ in their dispersal distances. Therefore, the described patterns have profound consequences for the genetic composition of populations.     

No evidence of sex‐biased dispersal in an island population of Common Blackbirds Turdus merula

Natal dispersal has major consequences for the dynamics and genetic structure of populations. Female‐biased natal dispersal, otherwise the norm in birds, is overridden when the place to move is limited, as on isolated islands. This effect was confirmed for the fist time in a European study system, the Common Blackbird breeding on Heligoland Island. Spatially restricted and sexually uniform natal dispersal may be a prerequisite for successful establishment of populations on remote islands or isolated habitat fragments, and this could play a major role in speciation processes.     

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.     

鸟类的扩散行为研究进展

扩散是生物个体之间相互远离的单线性运动,是生物的基本特征之一,对种群的分布、动态及遗传结构等方面均有重要影响.扩散有出生扩散和繁殖扩散等主要形式.动物发生扩散的主要原因包括:避免近亲繁殖、减少竞争、改变繁殖地点等.近年来,扩散已经成为鸟类学研究的前沿领域.评述了鸟类扩散行为的性别差异、体质对于扩散的影响,阐述了扩散的基本过程及栖息地选择、长距离扩散等内容,同时介绍了环志标记、无线电遥测、分子生物学等研究鸟类扩散的主要方法.展望了鸟类扩散研究的发展趋势,认为新技术和新方法的应用将成为扩散生态学家关注的重要问题,未来研究将更加重视对鸟类扩散理论问题的探讨,而对鸟类扩散行为的研究成果也会更广泛地应用于濒危物种及其栖息地的保护工作中.     

Density, social information, and space use in the common lizard (Lacerta vivipara)

Socially acquired information is widespread in the animal kingdom.Many individuals make behavioral decisions based on such socialinformation. In particular, individuals may decide to leaveor select their habitat based on social information. Few studieshave investigated the role of density-related information, apotential social cue about habitat quality in dispersal. Here,we tested for the possibility that the phenotype of intrudercommon lizards (Lacerta vivipara) may inadvertently carry informationabout their natal population density. We found that such informationuse is likely. The behavior of focal lizard was influenced bythe natal population density of the intruder it was interactingwith. This suggests that individuals may use the behavior ofothers to acquire appropriate information about surroundingsand to base spatial decisions on this information. Density-relatedinformation may then affect individual movement decisions andthus metapopulation dynamics.     

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.     

Patterns and processes of dispersal behaviour in arvicoline rodents

A good understanding of mammalian societies requires measuring patterns and comprehending processes of dispersal in each sex. We investigated dispersal behaviour in arvicoline rodents, a subfamily of mammals widespread in northern temperate environments and characterized by a multivoltine life cycle. In arvicoline rodents, variation in life history strategies occurs along a continuum from precocial to delayed maturation that reflects seasonal and ecological fluctuations. We compared dispersal across and within species focusing on the effects of external (condition-dependent) and internal (phenotype-dependent) factors. Our data revealed substantial, unexplained variation between species for dispersal distances and a strong variation within species for both dispersal distance and fraction. Some methodological aspects explained variation across studies, which cautions against comparisons that do not control for them. Overall, the species under consideration display frequent short-distance dispersal events and extremely flexible dispersal strategies, but they also have hitherto unexpected capacity to disperse long distances. Female arvicolines are predominantly philopatric relative to males, but we found no clear association between the mating system and the degree of sex bias in dispersal across species. Dispersal is a response to both various proximate and ultimate factors, including competition, inbreeding avoidance, mate searching and habitat quality. In particular, our review suggests that costs and benefits experienced during transience and settlement are prime determinants of condition dependence. Patterns of phenotype-dependent dispersal are idiosyncratic, except for a widespread association between an exploration/activity syndrome and natal dispersal. Consequences for population dynamics and genetic structures are discussed.     

Context-dependent dispersal,public information,and heterospecific attraction in newts

Dispersal is one of the main processes that determine community structure. Individuals make dispersal decisions according to environmental and/or social cues that reflect the fitness prospects in a given patch. The presence and abundance of heterospecifics within the same ecological guild, and/or their breeding success, may act as public information that influences movement decisions. To date, most studies investigating the role of heterospecific attraction have focused on habitat choice, using both experimental and correlational approaches. The present study is the first to examine how long-term variation in heterospecific density in breeding patches may affect dispersal patterns in spatially structured populations. We investigate how the dispersal decisions of the great crested newt (Triturus cristatus) are related to the variable density of two other newt species, the alpine newt (Ichthyosaura alpestris) and the palmate newt (Lissotriton helveticus). To examine this issue, we used capture–recapture data collected in an experimental pond network over a 20-year period. The results revealed that the great crested newt’s dispersal is context dependent and is affected by variation in heterospecific density: individuals were less likely to emigrate from ponds with high heterospecific density and were more likely to immigrate to ponds with high heterospecific density. These findings suggest that individuals adjust their dispersal decisions at least partly based on public information provided by heterospecifics. This mechanism may play a critical role in the dynamics of spatially structured populations and community functioning.

     

Strategic reduction of help before dispersal in a cooperative breeder

In cooperative breeders, sexually mature subordinates can either queue for chances to inherit the breeding position in their natal group, or disperse to reproduce independently. The choice of one or the other option may be flexible, as when individuals respond to attractive dispersal options, or they may reflect fixed life-history trajectories. Here, we show in a permanently marked, natural population of the cooperatively breeding cichlid fish Neolamprologus pulcher that subordinate helpers reduce investment in territory defence shortly before dispersing. Such reduction of effort is not shown by subordinates who stay and inherit the breeding position. This difference suggests that subordinates ready to leave reduce their investment in the natal territory strategically in favour of future life-history perspectives. It seems to be part of a conditional choice of the dispersal tactic, as this reduction in effort appears only shortly before dispersal, whereas philopatric and dispersing helpers do not differ in defence effort earlier in life. Hence, cooperative territory defence is state-dependent and plastic rather than a consistent part of a fixed life-history trajectory.     

Habitat choice stabilizes metapopulation dynamics by enabling ecological specialization

Dispersal is a key trait responsible for the spread of individuals and genes among local populations, thereby generating eco‐evolutionary interactions. Especially in heterogeneous metapopulations, a tight coupling between dispersal, population dynamics and the evolution of local adaptation is expected. In this respect, dispersal should counteract ecological specialization by redistributing locally selected phenotypes (i.e. migration load). Habitat choice following an informed dispersal decision, however, can facilitate the evolution of ecological specialization. How such informed decisions influence metapopulation size and variability is yet to be determined. By means of individual‐based modelling, we demonstrate that informed decisions about both departure and settlement decouple the evolution of dispersal and that of generalism, selecting for highly dispersive specialists. Choice at settlement is based on information from the entire dispersal range, and therefore decouples dispersal from ecological specialization more effectively than choice at departure, which is only based on local information. Additionally, habitat choice at departure and settlement reduces local and metapopulation variability because of the maintenance of ecological specialization at all levels of dispersal propensity. Our study illustrates the important role of habitat choice for dynamics of spatially structured populations and thus emphasizes the importance of considering that dispersal is often informed.     

Describing dispersal under habitat constraints: A randomization approach in lesser kestrels

Animal dispersal is usually studied with capture-mark-reencounter data, which provide information on realized dispersal but rarely on underlying processes. In this context, the unreliable assumption of all habitat being available is usually made when describing and analysing dispersal patterns. However, actual settlement options may be constrained by the spatial distribution of appropriate patches, so an important task to understand movement patterns is to adequately describe dispersal when the dispersers’ options are constrained by the sites that are available to them. Using a long-term monitored population of the migratory lesser kestrel, we show how randomization procedures can be used to describe dispersal strategies in such situations. This species breeds colonially in discrete patches, most individuals (83%) disperse from their natal colony, and dispersers tend to move short distances (median=7.2 km). Observed patterns (natal dispersal rates and median dispersal distances of birds emigrating from their natal colony) were compared with those expected from two null models of random settlement of individuals: in any colony available in the whole population, or within the subpopulation (cluster of colonies) of origin. Our simulations indicate that philopatry to the natal colony was much higher than expected under both null expectations, and observed distances were much lower than expected in the whole population. When individuals were constrained to settle within their natal subpopulation in the simulations, dispersal distances were longer than expected in females, but were higher or lower in males depending on year. Dispersal was not only constrained by the spatial distribution of settlement options, but specific hypotheses arise that can be helpful to design and conduct further research. These results challenge previous interpretations of observed dispersal patterns, which may not reflect free decisions of individuals but environmental or social constraints. We suggest using simulation procedures as a routine to advance in the understanding of dispersal ecology and evolution.     

Estimation by capture-recapture of recruitment and dispersal over several sites

Dispersal in animal populations is intimately linked with accession to reproduction, i.e. recruitment, and population regulation. Dispersal processes are thus a key component of population dynamics to the same extent as reproduction or mortality processes. Despite the growing interest in spatial aspects of population dynamics, the methodology for estimating dispersal, in particular in relation with recruitment, is limited. In many animal populations, in particular vertebrates, the impossibility of following individuals over space and time in an exhaustive way leads to the need to frame the estimation of dispersal in the context of capture-recapture methodology. We present here a class of age-dependent multistate capture-recapture models for the simultaneous estimation of natal dispersal, breeding dispersal, and age-dependent recruitment. These models are suitable for populations in which individuals are marked at birth and then recaptured over several sites. Under simple constraints, they can be used in populations where non-breeders are not observed, as is often the case with colonial waterbirds monitored on their breeding grounds. Biological questions can be addressed by comparing models differing in structure, according to the generalized linear model philosophy broadly used in capture-recapture methodology. We illustrate the potential of this approach by an analysis of recruitment and dispersal in the roseate tern Sterna dougallii .     

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.     

Natal dispersal in great bustards: the effect of sex, local population size and spatial isolation

1. We investigated the causes of natal dispersal in four Spanish areas where 35 breeding groups of the polygynous great bustard Otis tarda were monitored intensively. A total of 392 juveniles were radio-tracked between 1991 and 2006 by ground and via aeroplane to avoid potential biases derived from the non-detection of long-distance dispersers. 2. We explored 10 explanatory variables that were related to individual phenotypic features, habitat and conspecific traits in terms of group size and breeding performance, and spatial distribution of available breeding groups. Probability of group change and natal dispersal distances were investigated separately through multifactorial analyses. 3. Natal dispersal occurred in 47.8% of the birds and median natal dispersal distance of dispersers was 18.1 km (range 4.97-178.42 km). Sex largely determined the dispersal probability, with 75.6% of males being dispersers and 80.0% of females being philopatric, in contrast to the general pattern of female-biased dispersal found in most avian species. 4. Both the frequency of natal dispersal and dispersal distances were affected by the spatial distribution of breeding groups. More isolated groups showed a higher proportion of philopatric individuals, the effect being more evident in males than in females. This implies a reduction in gene flow in fragmented populations, as most genetic exchange is achieved through male dispersal. Additionally, dispersers hatched in more isolated groups tended to exhibit longer dispersal distances, which increases the associated energetic costs and mortality risks. 5. The dispersal decision was influenced by the number of conspecifics in the natal group. The individual probability of natal dispersal was related inversely to the size of the natal group, which supports the balanced dispersal model and the conspecific attraction hypothesis. 6. Overall, our results provide a good example of phenotypic plasticity and reinforce the current view that dispersal is an evolutionary complex trait conditioned by the interaction of individual, social and environmental causes that vary between individuals and populations.