Melanin-based colouration predicts natal dispersal in the barn owl, Tyto alba

Searching for a suitable breeding site is an important decision in the life of most animals. The decisions where to settle and how far to travel before doing so depend on many factors. Individual differences in dispersal distance could result from different strategies (e.g. specialists versus generalists), which might result in similar reproductive success in different habitats, or different competitive abilities to acquire a territory close to the natal site. The barn owl is polymorphic in melanic coloration, which is associated with many physiological and behavioural traits such as habitat choice, stress response and docility, raising the possibility that the coloration is also related to dispersal. We studied natal dispersal (from rearing site to site of first breeding attempt) and breeding dispersal (from one breeding site to the next) in barn owls using a long-term data set. Darker reddish individuals moved further than paler individuals during natal dispersal, but not during breeding dispersal. A cross-fostering experiment showed that the colour of the biological and foster parents had no influence on dispersal distance. The distance dispersed by parents and same-sex offspring was correlated, whereas natal and breeding dispersal were not repeatable within individuals, indicating that they are two different processes. Given that the distance travelled in natal dispersal appears to be heritable, the underlying genes might be coupled to those related to coloration. We discuss hypotheses to explain the potential adaptive function of the link between coloration and natal dispersal.

The short- and long-term fitness consequences of natal dispersal in a wild bird population

Dispersal is a key process in population and evolutionary ecology. Individual decisions are affected by fitness consequences of dispersal, but these are difficult to measure in wild populations. A long-term dataset on a geographically closed bird population, the Mauritius kestrel, offers a rare opportunity to explore fitness consequences. Females dispersed further when the availability of local breeding sites was limited, whereas male dispersal correlated with phenotypic traits. Female but not male fitness was lower when they dispersed longer distances compared to settling close to home. These results suggest a cost of dispersal in females. We found evidence of both short- and long-term fitness consequences of natal dispersal in females, including reduced fecundity in early life and more rapid aging in later life. Taken together, our results indicate that dispersal in early life might shape life history strategies in wild populations.

Maternal modulation of natal dispersal in a passerine bird: An adaptive strategy to cope with parasitism?

The decision of how far to disperse from the natal territory has profound and long-lasting consequences for young animals, yet the optimal dispersal behavior often depends on environmental factors that are difficult or impossible to assess by inexperienced juveniles. Natural selection thus favors mechanisms that allow the adaptive and flexible adjustment of the offspring's dispersal behavior by their parents via either paternal or maternal effects. Here we show that different dispersal strategies maximize the reproductive success of young great tits (Parus major) originating from a parasite-infested or a parasite-free nest and demonstrate that differential transfer of maternal yolk androgens in response to parasitism can result in a modification of the offspring's dispersal behavior that appears adaptive. It demonstrates that prenatal maternal effects are an important yet so far neglected determinant of natal dispersal and highlights the potential importance of maternal effects in mediating coevolutionary processes in host-parasite systems.

Bright moonlight triggers natal dispersal departures

Upon leaving their natal area, dispersers are confronted with unknown terrains. Species-specific perceptual ranges (i.e. the maximum distance from which an individual can perceive landscape features) play a crucial role in spatial movement decisions during such wanderings. In nocturnal animals that rely on vision, perceptual range is dramatically enhanced during moonlight, compared to moonless conditions. This increase of the perceptual range is an overlooked element that may be responsible for the successful crossing of unfamiliar areas during dispersal. The information gathered from 143 radio-tagged eagle owl Bubo bubo juveniles in Spain, Finland and Switzerland shows that, although the decision to initiate dispersal is mainly an endogenous phenomenon determined by the attainment of a given age (∼6 months), dispersers leave their birthplace primarily under the best light conditions at night, i.e. when most of the lunar disc is illuminated. This sheds new light into the mechanisms that may trigger dispersal from parental territory.

Modelling dispersal behaviour on a fractal landscape

We use a spatially explicit population model to explore the population consequences of different habitat selection mechanisms on landscapes with fractal variation in habitat quality. We consider dispersal strategies ranging from random walks to perfect habitat selectors for two species of arboreal marsupial, the greater glider (Petauroides volans) and the mountain brushtail possum (Trichosurus caninus). In this model increasing habitat selection means individuals obtain higher quality territories, but experience increased mortality during dispersal. The net effect is that population sizes are smaller when individuals actively select habitat. We find positive relationships between habitat quality and population size can occur when individuals do not use information about the entire landscape when habitat quality is spatially autocorrelated. We also find that individual behaviour can mitigate the negative effects of spatial variation on population average survival and fecundity. (C) 1998 Elsevier Science Ltd. All rights reserved.

Social structures, genetic structures and dispersal strategies in Australian rabbit (Oryctolagus cuniculus) populations

In this study, the pattern of movement of young male and female rabbits and the genetic structures present in adult male and female populations in four habitats was examined. The level of philopatry in young animals was found to vary between 18-90% for males and 32-95% for females in different populations. It was skewed, with more males dispersing than females in some but not all populations. Analysis of allozyme data using spatial autocorrelation showed that adult females from the same social group, unlike males, were significantly related in four of the five populations studied. Changes in genetic structure and rate of dispersal were measured before and during the recovery of a population that was artificially reduced in size. There were changes in the rate and distance of dispersal with density and sex. Subadults of both sexes moved further in the first year post crash (low density) than in the following years. While the level of dispersal for females was lower than that of the males for the first 3 years, thereafter (high density) both sexes showed similar, low levels of dispersal (20%). The density at which young animals switch behaviour between dispersal and philopatry differed for males and females. The level of genetic structuring in adult females was high in the precrash population, reduced in the first year post crash and undetectable in the second year. Dispersal behaviour of rabbits both affects the genetic structure of the population and changes with conditions. Over a wide range of levels of philopatry, genetic structuring is present in the adult female, but not the male population. Consequently, though genetic structuring is present, it does not lead to inbreeding. More long-distance movements are found in low-density populations, even though vacant warrens are available near birth warrens. The distances moved decreased as density increased. Calculation of the effective population size (N-e) shows that changes in dispersal distance offset changes in density, so that N-e remains constant.

Divorce, dispersal and incest avoidance in the cooperatively breeding superb fairy-wren Malurus cyaneus

1. Between 1988 and 2001, we studied social relationships in the superb fairy-wren Malurus cyaneus (Latham), a cooperative breeder with male helpers in which extra-group fertilizations are more common than within-pair fertilizations. 2. Unlike other fairy-wren species, females never bred on their natal territory. First-year females dispersed either directly from their natal territory to a breeding vacancy or to a foreign 'staging-post' territory where they spent their first winter as a subordinate. Females dispersing to a foreign territory settled in larger groups. Females on foreign territories inherited the territory if the dominant female died, and were sometimes able to split the territory into two by pairing with a helper male. However, most dispersed again to obtain a vacancy. 3. Females dispersing from a staging post usually gained a neighbouring vacancy, but females gaining a vacancy directly from their natal territory travelled further, perhaps to avoid pairing or mating with related males. 4. Females frequently divorced their partner, although the majority of relationships were terminated by the death of one of the pair. If death did not intervene, one-third of pairings were terminated by female-initiated divorce within 1000 days. 5. Three divorce syndromes were recognized. First, females that failed to obtain a preferred territory moved to territories with more helpers. Secondly, females that became paired to their sons when their partner died usually divorced away from them. Thirdly, females that have been in a long relationship divorce once a son has gained the senior helper position. 6. Dispersal to avoid pairing with sons is consistent with incest avoidance. However, there may be two additional benefits. Mothers do not mate with their sons, so dispersal by the mother liberates her sons to compete for within-group matings. Further, divorcing once their son has become a breeder or a senior helper allows the female to start sons in a queue for dominance on another territory. Females that do not take this option face constraints on their ability to recruit more sons into the local neighbourhood.

Local Competition, Inbreeding, and the Evolution of Sex-Biased Dispersal.

Using game theory, we developed a kin-selection model to investigate the consequences of local competition and inbreeding depression on the evolution of natal dispersal. Mating systems have the potential to favor strong sex biases in dispersal because sex differences in potential reproductive success affect the balance between local resource competition and local mate competition. No bias is expected when local competition equally affects males and females, as happens in monogamous systems and also in polygynous or promiscuous ones as long as female fitness is limited by extrinsic factors (breeding resources). In contrast, a male-biased dispersal is predicted when local mate competition exceeds local resource competition, as happens under polygyny/promiscuity when female fitness is limited by intrinsic factors (maximal rate of processing resources rather than resources themselves). This bias is reinforced by among-sex interactions: female philopatry enhances breeding opportunities for related males, while male dispersal decreases the chances that related females will inbreed. These results meet empirical patterns in mammals: polygynous/promiscuous species usually display a male-biased dispersal, while both sexes disperse in monogamous species. A parallel is drawn with sex-ratio theory, which also predicts biases toward the sex that suffers less from local competition. Optimal sex ratios and optimal sex-specific dispersal show mutual dependence, which argues for the development of coevolution models.

Female-biased dispersal in the monogamous mammal Crocidura russula: evidence from field data and microsatellite patterns.

We investigated dispersal patterns in the monogamous Crocidura russula, based both on direct field observations (mark-recapture data) and on genetic analyses (microsatellite loci). Natal dispersal was found to be low. Most juveniles settled within their natal territory or one immediately adjacent. Migration rate was estimated to two individuals per year and per population. The correlation between genetic and geographical distances over a 16 km transect implies that migration occurs over short ranges. Natal dispersal was restricted to first-litter juveniles weaned in early May; this result suggests a direct dependence of dispersal on reproductive opportunities. Natal dispersal was highly female biased, a pattern unusual among mammals. Its association with monogamy provides support for the resource-competition model of dispersal. Our results demonstrate that a state-biased dispersal can be directly inferred from microsatellite genotype distributions, which opens new perspectives for empirical studies in this area.

Weather to disperse? Evidence that climatic conditions influence vertebrate dispersal

1. Dispersal is regarded as critical to the stability of existing populations and the spread of invading species, but empirical data on the effect of travelling conditions during the transfer phase are rare. We present evidence that both timing and distance of ex-natal dispersal in buzzards (Buteo buteo) are strongly affected by weather. 2. Dispersal was recorded more often when the wind changed to a more southerly direction from the more common westerly winds, and when minimum temperatures were lower. The effect of wind direction was greatest in the winter and minimum temperature was most important in the autumn. Poor weather did not appear to initiate dispersal. 3. Dispersal distance was most strongly correlated with maximum temperature during dispersal and wind direction in the following 5-day period. Combined with the sex of the buzzard these three variables accounted for 60% of the variation in dispersal distance. 4. These results are important for conservationists who manage species recovery programs and wildlife managers who model biological invasions.

Post-fledging care, philopatry and recruitment in brown thornbills

1. We describe patterns of post-fledging care, dispersal and recruitment in four cohorts of brown thornbills Acanthiza pusilla. We examine what factors influence post-fledging survival and determine how post-hedging care and the timing of dispersal influence the probability of recruitment in this small, pair breeding, Australian passerine. 2. Fledgling thornbills were dependent on their parents for approximately 6 weeks. Male fledglings were more likely than female fledglings to survive until independence. For both sexes, the probability of reaching independence increased as nestling weight increased and was higher for nestlings that fledged later in the season. 3. The timing of dispersal by juvenile thornbills was bimodal. Juveniles either dispersed by the end of the breeding season or remained on their natal territory into the autumn and winter. Juveniles that delayed dispersal were four times more likely to recruit into the local breeding population than juveniles that dispersed early. 4. Delayed dispersal was advantageous because individuals that remained on their natal territory suffered little mortality and tended to disperse only when a local vacancy was available. Consequently, the risk of mortality associated with obtaining a breeding vacancy using this dispersal strategy was low. 5. Males, the more philopatric sex, were far more likely than females to delay dispersal. Despite the apparent advantages of prolonged natal philopatry, however, only 54% of pairs that raised male fledglings to independence had sons that postponed dispersal, and most of these philopatric sons gained vacancies before their parents bred again. Consequently, few sons have the opportunity to help their parents. Constraints on delayed dispersal therefore appear to play a major role in the evolution of pair-breeding in the brown thornbill.

Personality and melanin-based colouration traits in barn owls, tawny owl and kestrels

Individuals need to adapt to their local environment in order to survive. When selection pressures differ in local populations, polymorphism can evolve. Colour polymorphism is one of the most obvious polymorphisms since it is readily observable. Different sources of colouration exist, but melanin-based colouration is one of the most common in birds. The melanocortin system produces this colouration and because the melanocortin system has pleiotropic effects on behavioural and physiological traits, it is a good candidate to be an underlying mechanism to explain the maintenance of colour polymorphism. In this thesis I studied three different raptors which all display melanin-based colouration; barn owls (Tyto alba), tawny owls (Strix aluco) and Eurasian kestrels (Falco tinnunculus). The main question was if there was a relationship between melanin-based colouration and individual behavioural differences. The underlying hypothesis is that colour could be a signal of certain adaptive traits. Our goal was to find evolutionary explanations for the persistence of colour polymorphism. I found that nestling kestrels and barn owls differ in anti-predatory behaviour, with respect to their melanic colouration (chapters 1 and 2). Darker individuals show less reaction to human handling, but in kestrels aggression and colouration are related in opposite ways than in barn owls. More reddish barn owls travel greater distances in natal dispersal and this behaviour is repeatable between parents and same sex offspring (chapter 3). Dark reddish tawny owls defend their nests more intensely against intruders and appear to suffer less from nest predation (chapter 4). Finally I show that polymorphism in the Melanocortin 1 receptor gene (MC1R), which is strongly correlated with reddish colouration in the barn owl, is related to natal dispersal distance, providing a first indication for a genetic basis of the relation between this behaviour and colouration (chapter 5). My results demonstrate a clear link between melanin-based colouration and animal personality traits. I demonstrated this relation in three different species, which shows there is most likely a general underlying mechanism responsible. Different predation pressures might have shaped the reactions to predation, but also differences in sex-related colouration. Male-like and female-like colouration might signal more or less aggressive behaviour. Fluctuating environmental conditions might cause different individual strategies to produce equal reproductive success. The melanocortin system with its pleiotropic effects might be an underlying mechanism, as suggested by the results from the genetic polymorphism, the similar results found in these three species and by the similar relations reported in other species. This thesis demonstrates that colouration and individual differences are correlated and it provides the first glimpse of an underlying system. We can now conduct a more directed search for underlying mechanisms and evolutionary explanations with the use of quantitative genetic methods.

9

Using analytical tools from game theory, we investigate the relevance of a series of hypotheses concerning natal dispersal, focusing in particular on the interaction between inbreeding and kin competition, as well as on the components of mating and social systems that are likely to interfere with these phenomena. A null model of pure kin competition avoidance predicts a balanced equilibrium in wich both sexes disperse equally. Inbreeding costs have the potential to destabilize the equilibrium, resulting in strongly sex-biased dispersal. This effect is mostly evident when the peculiarities of the mating system induce asymmetries in dispersal and/or inbreeding costs, or when kin cooperation counteracts kin competition. Inbreeding depression, however, is not the only possible cause for sex biases. The relevance of our results to empirical findings is dicussed and suggestions are made for further empirical or modelling work.

High connectivity in a long-lived High-Arctic seabird, the ivory gull Pagophila eburnea

Species may cope with rapid habitat changes by distribution shifts or adaptation to new conditions. A common feature of these responses is that they depend on how the process of dispersal connects populations, both demographically and genetically. We analyzed the genetic structure of a near-threatened high-Arctic seabird, the ivory gull (Pagophila eburnea) in order to infer the connectivity among gull colonies. We analyzed 343 individuals sampled from 16 localities across the circumpolar breeding range of ivory gulls, from northern Russia to the Canadian Arctic. To explore the roles of natal and breeding dispersal, we developed a population genetic model to relate dispersal behavior to the observed genetic structure of worldwide ivory gull populations. Our key finding is the striking genetic homogeneity of ivory gulls across their entire distribution range. The lack of population genetic structure found among colonies, in tandem with independent evidence of movement among colonies, suggests that ongoing effective dispersal is occurring across the Arctic Region. Our results contradict the dispersal patterns generally observed in seabirds where species movement capabilities are often not indicative of dispersal patterns. Model predictions show how natal and breeding dispersal may combine to shape the genetic homogeneity among ivory gull colonies separated by up to 2800 km. Although field data will be key to determine the role of dispersal for the demography of local colonies and refine the respective impacts of natal versus breeding dispersal, conservation planning needs to consider ivory gulls as a genetically homogeneous, Arctic-wide metapopulation effectively connected through dispersal.