A qualitative investigation of major urinary proteins in relation to the onset of aggressive behavior and dispersive motivation in male wild house mice (<Emphasis Type="Italic">Mus musculus domesticus</Emphasis>)

The physiological basis for population differentiation of dispersal timing during individual development in male wild house mice is still unknown. As major urinary proteins (MUPs) are known to convey information about competitive ability in male mice, we examined individual MUP profiles defined by isoelectric-focusing (IEF) patterns in relation to developmental timing of dispersive motivation. As an experimental paradigm marking the development of the dispersal propensity, we used agonistic onset between litter mate brothers when kept in pairs under laboratory conditions. Agonistic onset is known to reflect the initiation of dispersive motivation. Hence, we compared individual MUP IEF patterns between fraternal pairs that did or did not develop agonistic relationships before the age of 2 months. Urine was collected on the day of weaning and at the beginning of adulthood. We investigated whether there was a significant co-occurrence of particular MUP IEF patterns with the agonistic onset in male mice. We assumed that, based on this co-occurrence, particular MUP IEF patterns and/or a particular dynamic of MUP IEF expression from weaning to adulthood may be considered a physiological predictor of a specific behavioral strategy in male mice (i.e. submissive-philopatric or agonistic-dispersive strategy). We found that agonistic males expressed more MUP IEF bands than amicable ones at weaning, but these differences disappeared later on. The presence of two particular IEF bands at weaning was significantly associated with early agonistic onset. Our study suggests that MUPs could have a predictive value for the onset of aggressive behavior and dispersal tendency in male wild house mice.     

Female House Mice do not Differ in Their Exploratory Behaviour from Males

Dispersal is an important ecological trait affecting genetic variation and dynamics of populations. Hence, the exploratory behaviour prior to actual dispersal may be crucial for potentially dispersing individuals. In mammals, females are traditionally seen as the more philopatric sex and dispersal as male‐biased behaviour, and so behavioural strategies related to the exploration of novel resources should be differentially expressed in males and females. In addition, due to sexual selection exploratory strategies may be expected to vary according to females’ reproductive phase. We employed a standard open‐field test as an approximation of the first phase of dispersal, using adult house mice representing two subspecies, M. m. musculus and M. m. domesticus. We tested the prediction that exploration of neutral area varies in females during different phases of the oestrus cycle and is different between both sexes and subspecies. We expected to find higher exploration in males, as the more dispersing sex and less pronounced subspecies‐specific differences in females than in males. We found no significant effect of the oestrous phase on any of the parameters of the exploratory behaviour measured. Sexual dimorphism was found only in latency to enter the arena in M. m. domesticus where females hesitated longer to enter a new area than males. Significant subspecies‐specific differences were found in three of four tested exploration parameters, so we conclude that females of both subspecies follow similar strategies to those displayed by males. Musculus mice show shorter latency to enter a new area, but once inside, domesticus mice explore the arena significantly longer, with less frequent retreats to a shelter. Our results thus highlight that the role of female dispersal in interdemic gene flow should not be neglected.     

Repeatability and heritability of exploratory behaviour in great tits from the wild

We investigated whether individual great tits, Parus major, vary consistently in their exploratory behaviour in a novel environment and measured the repeatability and heritability of this trait. Wild birds were caught in their natural habitat, tested in the laboratory in an open field test on the following morning, then released at the capture site. We measured individual consistency of exploratory behaviour for recaptured individuals (repeatability) and estimated the heritability with parent-offspring regressions and sibling analyses. Measures of exploratory behaviour of individuals at repeated captures were consistent in both sexes and study areas (repeatabilities ranged from 0.27 to 0.48). Exploration scores did not differ between the sexes, and were unrelated to age, condition at fledging or condition during measurement. Heritability estimates were 0.22-0.41 (parent-offspring regressions) and 0.37-0.40 (sibling analyses). We conclude that (1) consistent individual variation in open field behaviour exists in individuals from the wild, and (2) this behavioural variation is heritable. This is one of the first studies showing heritable variation in a behavioural trait in animals from the wild, and poses the question of how this variation is maintained under natural conditions. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Fine‐scale genetic structure reflects sex‐specific dispersal strategies in a population of sociable weavers (Philetairus socius)

Dispersal is a critical driver of gene flow, with important consequences for population genetic structure, social interactions and other biological processes. Limited dispersal may result in kin‐structured populations in which kin selection may operate, but it may also increase the risk of kin competition and inbreeding. Here, we use a combination of long‐term field data and molecular genetics to examine dispersal patterns and their consequences for the population genetics of a highly social bird, the sociable weaver (Philetairus socius), which exhibits cooperation at various levels of sociality from nuclear family groups to its unique communal nests. Using 20 years of data, involving capture of 6508 birds and 3151 recaptures at 48 colonies, we found that both sexes exhibit philopatry and that any dispersal occurs over relatively short distances. Dispersal is female‐biased, with females dispersing earlier, further, and to less closely related destination colonies than males. Genotyping data from 30 colonies showed that this pattern of dispersal is reflected by fine‐scale genetic structure for both sexes, revealed by isolation by distance in terms of genetic relatedness and significant genetic variance among colonies. Both relationships were stronger among males than females. Crucially, significant relatedness extended beyond the level of the colony for both sexes. Such fine‐scale population genetic structure may have played an important role in the evolution of cooperative behaviour in this species, but it may also result in a significant inbreeding risk, against which female‐biased dispersal alone is unlikely to be an effective strategy.     

Immune gene variability influences roe deer natal dispersal

Dispersal is a key life‐history trait governing the response of individuals, populations and species to changing environmental conditions. In the context of global change, it is therefore essential to better understand the respective role of condition‐, phenotype‐ and genetic‐dependent drivers of dispersal behaviour. Although the importance of immune function and pathogen infestation in determining patterns of dispersal is increasingly recognised, no study to our knowledge has yet investigated the influence of immune gene variability on dispersal behaviour. Here, we filled this knowledge gap by assessing whether individual heterozygosity at five immune gene loci (one from the Major histocompatibility complex and four from encoding Toll‐like receptors) influences roe deer natal dispersal. We found that dispersal propensity was affected by immune gene diversity, suggesting potential pathogen‐mediated selection through over‐dominance. However, the direction of this effect differed between high and low quality individuals, suggesting that dispersal propensity is driven by two different mechanisms. In support of the condition‐dependent dispersal hypothesis, dispersal propensity increased with increasing body mass and, among high quality individuals only (standardized body mass > 18 kg), with increasing immune gene diversity. However, among poor quality individuals, we observed the opposite pattern such that dispersal propensity was higher for individuals with lower immune gene diversity. We suggest that these poor quality individuals expressed an emergency dispersal tactic in an attempt to escape a heavily infested environment associated with poor fitness prospects. Our results have potentially important consequences in terms of population genetics and demography, as well as host–pathogen evolution.     

Increased propensity for aerial dispersal in disturbed habitats due to intraspecific variation and species turnover

Animal dispersal depends on multiple factors, such as habitat features and life‐history traits of the species. We studied the propensity for ballooning dispersal in spiders under standardized laboratory conditions. The 1269 tested individuals belonged to 124 species and originated from 16 sites with wide variation in habitat type. Spiders from disturbed habitats ballooned 5.5 times more than spiders from stable habitats. In Meioneta rurestris , for which we had enough data for a single‐species analysis, individuals were most dispersive if they originated from highly disturbed habitats. While the data for the other species were not sufficient for single‐species analyses, a hierarchical model that included the data simultaneously on all species suggested that dispersal propensity generally increases within species with the level of habitat disturbance. Dispersal probability showed a trend to increase with niche width, but the higher commonness of species with wide niches provides an alternative explanation for this pattern. As the prevalence of especially dispersive species was highest in disturbed habitats, variation in dispersal propensity was influenced by both inter‐ and intraspecific factors. We conclude that the positive correlation between niche width and dispersal propensity enables generalist species to utilize highly disturbed habitats, whereas the persistence of specialist species with restricted dispersal ability requires the conservation of stable habitats.     

Dispersal mechanisms in a captive wild house mouse population (Mus domesticus Rutty)

The ecological implications of dispersal have been discussed in many studies of wild animals in the field but little is known about the social mechanisms leading to the emigration of certain members of a group. To study the social background of dispersal in wild house mice ( Mus domesticus Rutty) a population cage system was evaluated that allowed permanent observation of individually marked animals. It consisted of ten cages connected to a central cage by transparent plastic tubes. Two of these cages were defined as 'dispersal cages' and could be reached only by swimming through a water basin. Dispersal was defined as a permanent stay in one of these cages for at least 4 days. At the beginning of the experiment one pair of house mice with their litter was placed into the cage system. Each of six experiments lasted for 6 months during which data on spacing, social interactions, body condition, reproduction, mortality and dispersal were collected by daily observations. Results regarding this study could be summarized as follows: (1) dispersal in house mice is male-biased; (2) there are interfamiliar differences in dispersal age, dispersal rate, and in the development of the population structure; (3) after reaching sexual maturity subdominant males are evicted by the dominant one; (4) reproductive rate among females drops with increasing birth order, thus only the oldest females within a group reproduce; (5) females born under high population density conditions can only reproduce after dispersal.     

Aggressive behaviour in related and unrelated wild house mice (Mus musculus L.)

Aggressive behaviour was observed to be rare in small family groups of confined wild house mice, Mus musculus L. Unrelated mice were attacked when they were introduced to a family group and in their presence intra-family aggressive behaviour increased. When two family groups of mice were allowed to meet there were frequent aggressive encounters between unrelated animals and the two groups remained separate. Resident mice were found to be aggressive towards males and females individually isolated and returned to their own family after 2 or 3 weeks absence but not after 1 week. The possibility is discussed that in wild mice odour discrimination influences the dispersal and build-up of free-living populations.     

First evidence for heritable variation in cooperative breeding behaviour

Understanding the evolution of complex social behaviours, such as cooperative breeding, is a fundamental problem in evolutionary biology, which has attracted much theoretical and empirical interest. Variation within and between species in the frequency of helping behaviour has been typically associated with variation in direct costs and benefits due to ecological constraints, or with indirect fitness payoffs (i.e. kin selection). Here, we provide the first evidence that individual variation in cooperative behaviour within a natural population also has a heritable component. Using a seven-generation pedigree in a wild population of western bluebirds (Sialia mexicana), we show significant heritable variation for the propensity to help rather than breed, as well as for the probability of having a helper at the nest. We also document a strong positive relationship between a bird's lifespan and its prospect of receiving help when breeding, in accordance with earlier comparative studies across species. These findings provide useful insights into the possible mechanisms which have led to the evolution of cooperative breeding and other social systems.     

Artificial selection for increased dispersal results in lower fitness

Dispersal often covaries with other traits, and this covariation was shown to have a genetic basis. Here, we wanted to explore to what extent genetic constraints and correlational selection can explain patterns of covariation between dispersal and key life‐history traits—lifespan and reproduction. A prediction from the fitness‐associated dispersal hypothesis was that lower genetic quality is associated with higher dispersal propensity as driven by the benefits of genetic mixing. We wanted to contrast it with a prediction from a different model that individuals putting more emphasis on current rather than future reproduction disperse more, as they are expected to be more risk‐prone and exploratory. However, if dispersal has inherent costs, this will also result in a negative genetic correlation between higher rates of dispersal and some aspects of performance. To explore this issue, we used the dioecious nematode Caenorhabditis remanei and selected for increased and decreased dispersal propensity for 10 generations, followed by five generations of relaxed selection. Dispersal propensity responded to selection, and females from high‐dispersal lines dispersed more than females from low‐dispersal lines. Females selected for increased dispersal propensity produced fewer offspring and were more likely to die from matricide, which is associated with a low physiological condition in Caenorhabditis nematodes. There was no evidence for differences in age‐specific reproductive effort between high‐ and low‐dispersal females. Rather, reproductive output of high‐dispersal females was consistently reduced. We argue that our data provide support for the fitness‐associated dispersal hypothesis.     

Dispersal in house mice

This review evaluates direct (live-trapping) and indirect (genetic) methods to study dispersal in wild house mice ( Mus musculus ) and summarizes field and experimental data to examine the causes and consequences of dispersal. Commensal house mice (associated with human habitations, farms, food stores and other anthropogenic habitats) typically show lower rates of dispersal than feral house mice (living in crops, natural and semi-natural habitats). However, early claims of long-term fine-scale genetic structure in commensal house mice (due to low rates of dispersal) are not supported by recent data. Dispersal becomes obligatory when habitat conditions deteriorate, but most dispersal occurs below the local environmental carrying capacity and is due to social interactions with conspecifics. Excursions are relatively frequent and probably allow mice to assess the quality of habitats before dispersing. Young males have the greatest tendency to disperse, apparently prompted mainly by aggressive interactions with dominant males. If they do disperse, females integrate into new groups more easily than do males. Dispersing house mice risk loss of condition or death, but may gain reproductive opportunities on arrival in a new location. House mice can be transported passively as stowaways with humans; this contributes to population persistence and genetic structure at regional scales and has allowed house mice to spread world-wide.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 565–583.     

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 for divergence in male harmfulness or female resistance in response to changes in the opportunity for dispersal

The outcome of sexual conflict can depend on the social environment, as males respond to changes in the inclusive fitness payoffs of harmfulness and harm females less when they compete with familiar relatives. Theoretical models also predict that if limited male dispersal predictably enhances local relatedness while maintaining global competition, kin selection can produce evolutionary divergences in male harmfulness among populations. Experimental tests of these predictions, however, are rare. We assessed rates of dispersal in female and male seed beetles Callosobruchus maculatus, a model species for studies of sexual conflict, in an experimental setting. Females dispersed significantly more often than males, but dispersing males travelled just as far as dispersing females. Next, we used experimental evolution to test whether limiting dispersal allowed the action of kin selection to affect divergence in male harmfulness and female resistance. Populations of C. maculatus were evolved for 20 and 25 generations under one of three dispersal regimens: completely free dispersal, limited dispersal and no dispersal. There was no divergence among treatments in female reproductive tract scarring, ejaculate size, mating behaviour, fitness of experimental females mated to stock males or fitness of stock females mated to experimental males. We suggest that this is likely due to insufficient strength of kin selection rather than a lack of genetic variation or time for selection. Limited dispersal alone is therefore not sufficient for kin selection to reduce male harmfulness in this species, consistent with general predictions that limited dispersal will only allow kin selection if local relatedness is independent of the intensity of competition among kin.     

OF MICE AND KIN: THE FUNCTIONAL SIGNIFICANCE OF KIN BIAS IN SOCIAL BEHAVIOUR

1. Sharing recent ancestry (kinship) increases the degree of genetic similarity between individuals, where genetic similarity could mean anything from sharing a particular allele to sharing an entire genome. 2. Genetic similarity can influence behavioural and other responses between individuals in a number of ways, discriminatory and non-discriminatory. All are likely to result in kin bias, because of the correlation between genetic similarity and kinship, but only some should be regarded as involving kin discrimination. 3. Non-discriminatory kin bias could arise through close relatives sharing, for instance, physical characteristics (such as those influencing competitive ability), thresholds of behavioural response or requirements for particular resources. 4. Discriminatory kin bias could arise through the direct perception of genetic similarity between individuals (direct similarity discrimination) or the use of cues likely to correlate with genetic similarity (indirect similarity discrimination--of which kin discrimination is one form). Alternatively, it could arise incidentally through mistaken identity or discrimination at some other level, such as species identification. 5. Experiments with laboratory and wild house mice have revealed kin bias in a number of contexts, including (a) parental and infanticidal behaviour, (b) sexual development and behaviour and (c) investigatory behaviour and passive body contact among juveniles and adults. 6. While kin bias in mice has been interpreted as evidence for kin discrimination, there are several problems with such an interpretation. These include (a) pronounced and complex effects of familiarity on discrimination, (b) a high risk of error-proneness in the indirect cues used in apparent kin discrimination and (c) weak and easily disrupted kin bias effects in certain contexts. 7. Consideration of social structure and discriminatory responses within populations of wild house mice leads to an alternative explanation for some kin bias in terms of incidental discrimination based on social group membership. 8. Several results from laboratory experiments suggest incidental discrimination is a more parsimonious explanation than kin discrimination for some intrasexual kin bias in behaviour. However, kin or direct similarity discrimination appears to be the most likely explanation for other aspects of intrasexual kin bias and for intersexual kin bias.     

Oxytocin and Social Preference in Female House Mice (Mus musculus domesticus)

In social species, same‐sex individuals may form social bonds behaviourally expressed as individual preferences, resulting in fitness benefits such as increased offspring survival, longevity and group cohesion. As a result of individual preferences, female house mice (Mus musculus domesticus) form social affiliations while communally nursing and may do so with kin or non‐kin. However, the mechanisms behind the formation of such preferences are unknown. Oxytocin has been linked to a range of social behaviours including bond facilitation, social memory and parental care. Here, we experimentally increased oxytocin in pairs of unfamiliar, unrelated females and predicted that females with elevated oxytocin would demonstrate increased affiliative behaviours compared against a control. Subsequently, we tested for the formation of a social preference, using a preference test with the previous partner and a new unfamiliar female. Our results indicated no significant effect of treatment on positive and negative behaviours between females during the three initial cohabitation days. In both treatments, females demonstrated increased socio‐positive behaviours and cohabitation time with their partner and decreased socio‐negative behaviours and latency to meet, over the 3‐d period. During the partner preference test, control but not oxytocin females demonstrated a significant preference for their cohabitation partner, and oxytocin females spent similar amounts of time with both stimulus females. Therefore, increasing peripheral oxytocin appears not to be involved in the facilitation of initial encounters with a stranger but may hinder the formation of a preference for this new partner.     

Dispersal behaviour of African wild dogs in Kenya

Dispersal behaviour plays a key role in social organisation, demography and population genetics. We describe dispersal behaviour in a population of African wild dogs (Lycaon pictus) in Kenya. Almost all individuals, of both sexes, left their natal packs, with 45 of 46 reproductively active “alpha” individuals acquiring their status through dispersal. Dispersal age, group size and distance did not differ between males and females. However, only females embarked on secondary dispersal, probably reflecting stronger reproductive competition among females than males. When dispersing, GPS-collared wild dogs travelled further than when resident, both in daylight and by night, following routes an order of magnitude longer than the straight-line distance covered. Dispersers experienced a daily mortality risk three times that experienced by adults in resident packs. The detailed movement data provided by GPS-collars helped to reconcile differences between dispersal patterns reported previously from other wild dog populations. However, the dispersal patterns observed at this and other sites contrast with those assumed in published demographic models for this endangered species. Given the central role of dispersal in demography, models of wild dog population dynamics need to be updated to account for improved understanding of dispersal processes.     

Affinity towards maternal odour and offspring dispersal in the common lizard

Kin competition is known to strongly influence the spacing behaviour of juveniles in the common lizard, a species which is only weakly social and has no parental care. We examined whether variation in offspring dispersal propensity was associated with variation in their ability to discriminate odours of their mothers versus unknown females. We caught pregnant females in two populations and reared them in the laboratory until parturition. Just after birth, we recorded the juvenile response to the odour of either their mother or an unrelated female. Mothers and offspring were then released at the mother's last capture point, and juvenile movement was monitored by recapture for 2 years. Philopatric individuals were more attracted to the odour of their mother and more repulsed by the odour of an unrelated female than dispersers. In this species, ability to discriminate maternal cues appears to be implied in the spacing behaviour of juveniles. Since juvenile males and females reacted in the same way, the main role of mother discrimination is likely to be the avoidance of kin competition by promoting juvenile dispersal. This does not necessary require mother discrimination to imply mother recognition, or not to be implied in other functions such as inbreeding avoidance. Further research is needed to clarify these points. We suggest that context-dependent kin discrimination may be involved in the spacing behaviour of a large number of species, even for those species where other social functions for kin discrimination have been proposed.     

Cooperation‐mediated plasticity in dispersal and colonization

Kin selection theory predicts that costly cooperative behaviors evolve most readily when directed toward kin. Dispersal plays a controversial role in the evolution of cooperation: dispersal decreases local population relatedness and thus opposes the evolution of cooperation, but limited dispersal increases kin competition and can negate the benefits of cooperation. Theoretical work has suggested that plasticity of dispersal, where individuals can adjust their dispersal decisions according to the social context, might help resolve this paradox and promote the evolution of cooperation. Here, we experimentally tested the hypothesis that conditional dispersal decisions are mediated by a cooperative strategy: we quantified the density‐dependent dispersal decisions and subsequent colonization efficiency from single cells or groups of cells among six genetic strains of the unicellular Tetrahymena thermophila that differ in their aggregation level (high, medium, and low), a behavior associated with cooperation strategy. We found that the plastic reaction norms of dispersal rate relative to density differed according to aggregation level: highly aggregative genotypes showed negative density‐dependent dispersal, whereas low‐aggregation genotypes showed maximum dispersal rates at intermediate density, and medium‐aggregation genotypes showed density‐independent dispersal with intermediate dispersal rate. Dispersers from highly aggregative genotypes had specialized long‐distance dispersal phenotypes, contrary to low‐aggregation genotypes; medium‐aggregation genotypes showing intermediate dispersal phenotype. Moreover, highly aggregation genotypes showed evidence for beneficial kin‐cooperation during dispersal. Our experimental results should help to resolve the evolutionary conflict between cooperation and dispersal: cooperative individuals are expected to avoid kin‐competition by dispersing long distances, but maintain the benefits of cooperation by dispersing in small groups.     

Behavioural synchronization of large‐scale animal movements – disperse alone,but migrate together?

Dispersal and migration are superficially similar large‐scale movements, but which appear to differ in terms of inter‐individual behavioural synchronization. Seasonal migration is a striking example of coordinated behaviour, enabling animal populations to track spatio‐temporal variation in ecological conditions. By contrast, for dispersal, while social context may influence an individual's emigration and settlement decisions, transience is believed to be mostly a solitary behaviour. Here, we review differences in drivers that may explain why migration appears to be more synchronized than dispersal. We derive the prediction that the contrast in the importance of behavioural synchronization between dispersal and migration is linked to differences in the selection pressures that drive their respective evolution. Although documented examples of collective dispersal are rare, this behaviour may be more common than currently believed, with important consequences for eco‐evolutionary dynamics. Crucially, to date, there is little available theory for predicting when we should expect collective dispersal to evolve, and we also lack empirical data to test predictions across species. By reviewing the state of the art in research on migration and collective movements, we identify how we can harness these advances, both in terms of theory and data collection, to broaden our understanding of synchronized dispersal and its importance in the context of global change.     

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.