Breeding dispersal strategies following reproductive failure explain low apparent survival of immigrant Temminck's stints

In some animal populations, immigrants have lower survival than philopatric individuals. Costs of dispersal or low phenotypic quality of dispersers may explain the pattern. However, apparent adult survival estimates, which describe real survival combined with site fidelity cannot be separated from permanent emigration. Thus, heterogeneity in breeding dispersal propensities of immigrants and philopatrics can bias fitness correlates of dispersal. Differences in breeding dispersal propensities may be caused by different strategies in response to environmental cues inducing dispersal, such as reproductive success. In such cases, the reported differences between immigrants and philopatric individuals may not reflect true variation in survival. We studied whether dispersal status specific apparent adult survival is associated with reproductive success in a Temminck's stint Calidris temminckii population. We analysed two long term capture–recapture datasets characterised by low and high nest predation levels. Philopatric individuals had higher apparent adult survival than immigrants in both datasets and the difference was highlighted during the high nest predation period. By contrasting return rates between successful and unsuccessful breeders as a proxy for dispersal, we found that unsuccessful immigrants breeding for the first time dispersed more likely than successful immigrants, but such a pattern was not found among philopatric individuals. Our results support the hypothesis that immigrant and philopatric individuals have different breeding dispersal strategies following reproductive failure and that their apparent adult survival differences are at least partly explained by different breeding dispersal propensities. Our results also suggest that the recent decline of the study population reflects a multiple response to increased nest predation through decreased local recruitment and increased emigration.     

The evolution of dispersal in a two-patch system: some consequences of differences between migrants and residents

We investigate how age-structure and differences in certain demographic traits between residents and immigrants of a single species act to determine the evolutionarily stable dispersal strategy in a two-patch environment that is heterogeneous in space but constant in time. These two factors have been neglected in previous models of the evolution of dispersal, which generally consider organisms with very simple life-cycles and assume that, whatever their origin, individuals in a given habitat have the same bio-demographic characteristics. However, there is increasing empirical evidence that dispersing individuals have different demographic properties from phylopatric ones. We develop a matrix model in which recruitment depends on local population densities. We assume that dispersal entails a proportional cost to immigrant fecundity, which can be compensated by differences in survival rates between immigrants and residents. The evolutionarily stable strategies (ESS) for dispersal are identified using a combination of analytical expressions and numerical simulations. Our results show that philopatry is selected (1) when dispersal rates do not vary in space, (2) when the metapopulation is a source-sink system and (3) when dispersal rates vary in space (asymmetric dispersal) and immigrants do not compensate for their reduced fecundity. We observe that non-zero asymmetric dispersal rates may be evolutionarily stable when (1) immigrants and residents are demographically alike and (2) immigrants compensate totally for their reduced fecundity through an increase in adult survival. Under these conditions, we find that the ESS occurs when the fitnesses at equilibrium in the two habitats, measured in our model by the realized reproductive rates, are each equal to unity. A comparison with previous studies suggests a unifying rule for the evolution of dispersal: the dispersal rates which permit the spatial homogenization of fitnesses are ESSs. This condition provides new insight into the evolutionary stability of source-sink systems. It also supports the hypothesis that immigrants have adapted demographic strategies, rather than the hypothesis that dispersal is costly and immigrants are at a disavantage compared with residents.     

On the evolutionary stability of sink populations

The evolution of adaptive behaviours can influence population dynamics. Conversely, population dynamics can affect both the rate and direction of adaptive evolution. This paper examines reasons why sink populations – populations maintained by immigration, preventing local extinction – might persist in the habitat repertoire of a species over evolutionary time-scales. Two such reasons correspond to standard explanations for deviations from an ideal free habitat distribution: organisms may not be free to settle in whichever habitat has the highest potential fitness, and may be constrained by costs, perceptual limitations, or mode of dispersal in the acuity of their habitat selectivity. Here, I argue that a third general reason for persistent sink populations is provided by unstable population dynamics in source habitats. I present a simple model illustrating how use of a sink habitat may be selectively advantageous, when a source population has unstable dynamics (which necessarily reflects temporal variation in local fitnesses). Species with unstable local dynamics in high-quality habitats should be selected to utilize a broader range of habitats than species with stable local dynamics, and in particular in some circumstances should utilize sink habitats. This observation has implications for the direction of niche evolution, and the likelihood of niche conservatism.     

Lifetime fitness correlates of natal dispersal distance in a colonial bird

1. Obtaining empirical evidence of the consequences of dispersal distance on fitness is challenging in wild animals because long-term, unbiased data on reproduction, survival and movement are notoriously difficult to obtain. 2. Lifetime fitness correlates of natal dispersal distance were studied in an isolated population of the facultatively colonial lesser kestrel Falco naumanni (Fleischer) monitored during 8 years at north-eastern Spain, where most birds (83%) dispersed from their natal colony to settle at distances ranging from 112 m to 136.5 km. 3. Neither annual breeding success nor age at recruitment was affected by natal dispersal distance. However, a capture-mark-recapture analysis revealed that survival during the year following recruitment decreased exponentially with dispersal distance, with differences of up to 15% between philopatrics and long-distance dispersers. In subsequent years, it remained similar irrespective of the natal dispersal distance moved. These results did not seem to be biased by long-distance dispersers settling differentially in the periphery of the population (which could emigrate permanently and be considered dead in future occasions) or within-individual consistency in successive dispersal distances, so our results appear to reflect genuine survival differences between dispersal tactics. 4. Average lifetime fledgling production, average lifetime recruitment success and rate-sensitive individual fitness (λ(ind)) also decreased with the distance from the natal to the first-breeding colony, indicating that dispersal decisions early in life affecting immediate survival prospects may translate into long-term fitness costs. 5. Both survival and lifetime fitness models including continuous dispersal distances significantly improved the characterization of the effect on fitness compared with models considering dispersal as a discrete process (i.e. dispersal vs. philopatry at a colony level). 6. Long-distance dispersers were more likely to establish new colonies regardless of whether they recruited in the centre or the periphery of the population, revealing their important role in the colonization of unoccupied patches. Individuals experienced a higher probability of mortality in small and newly funded colonies, so lifetime fitness costs of dispersal seem to be explained by recruitment in sites where average quality is low because of high uncertainty in survival prospects.     

Condition-dependent dispersal of a patchily distributed riparian ground beetle in response to disturbance

In common with many habitat elements of riverine landscapes, exposed riverine sediments (ERS) are highly disturbed, naturally patchy and regularly distributed, whose specialists are strongly adapted to flood disturbance and loss of habitat due to succession. Investigations of dispersal in ERS habitats therefore provide an important contrast to the unnaturally fragmented, stable systems usually studied. The present investigation analysed the three interdependent stages of dispersal: (1) emigration, (2) inter-patch movement and (3) immigration of a common ERS specialised beetle, Bembidion atrocaeruleum (Stephens 1828) (Coleoptera, Carabidae), in a relatively unmodified section of river, using mark–resight methods. Dispersal was correlated with estimates of local population size and density, water level and patch quality in order to test for condition-dependent dispersal cues. Flood inundation of habitat was found to increase strongly the overall rate of dispersal, and the rate of emigration was significantly higher from patches that were heavily trampled by cattle. Strongly declining numbers of dispersers with distance suggested low dispersal rates during periods of low water level. Dispersal in response to habitat degradation by cattle trampling would likely lead to a higher overall population fitness than a random dispersal strategy. Dispersal distances were probably adapted to the underlying habitat landscape distribution, high-flow dispersal cues and ready means of long-distance dispersal through hydrochory. Species whose dispersal is adapted to the natural habitat distribution of riverine landscapes are likely to be strongly negatively affected by reduced flood frequency and intensity and habitat fragmentation through flow regulation or channelisation.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

The new kid on the block: immigrant males win big whereas females pay fitness cost after dispersal

Dispersal is nearly universal; yet, which sex tends to disperse more and their success thereafter depends on the fitness consequences of dispersal. We asked if lifetime fitness differed between residents and immigrants (successful between‐population dispersers) and their offspring using 29 years of monitoring from North American red squirrels (Tamiasciurus hudsonicus) in Canada. Compared to residents, immigrant females had 23% lower lifetime breeding success (LBS), while immigrant males had 29% higher LBS. Male immigration and female residency were favoured. Offspring born to immigrants had 15–43% lower LBS than offspring born to residents. We conclude that immigration benefitted males, but not females, which appeared to be making the best of a bad lot. Our results are in line with male‐biased dispersal being driven by local mate competition and local resource enhancement, while the intergenerational cost to immigration is a new complication in explaining the drivers of sex‐biased dispersal.     

Annual Survival and Turnover Rates of an Afrotropical Robin in a Fragmented Forest

Whether or not subdivided populations persist in fragmented landscapes primarily depends on how well individuals can survive within discrete habitat patches. Using data from six capture–recapture sessions, survival probabilities of the white-starred robin were estimated in seven indigenous forest patches in the highly fragmented Taita Hills forests, SE Kenya. We found no significant differences in survival probability either among fragment-size categories (large 135 ha, medium 95 ha, and small 2–8 ha) or between adult and first-year birds. However, males had a higher probability of survival from one year to the next than females. Turnover rates of adult birds were higher for females than males, but also higher in the medium and small patches than in the large one within each sex. That survival probability was similar among fragments, but turnover rates differed denoted that different processes caused extirpation from the patches. We suggest that mortality associated with dispersal was probably a more important cause of extirpation than within-patch mortality in the largest habitat, which had the lowest turnover rates. Conversely, high within-patch mortality, for instance due to predation during incubation, could have been more important in the smaller, more disturbed habitats. These results lend support to the proposition that avian conservation efforts should be focussed both at the landscape level to improve connectivity between fragments and reduce mortality during dispersal, and at the patch level to exclude other mortality sources such as nest predation.     

Dispersal and habitat cuing of Eurasian red squirrels in fragmented habitats

Animal dispersal and subsequent settlement is a key process in the life history of many organisms, when individuals use demographic and environmental cues to target post-dispersal habitats where fitness will be highest. To investigate the hypothesis that environmental disturbance (habitat fragmentation) may alter these cues, we compared dispersal patterns of 60 red squirrels (Sciurus vulgaris) in three study sites that differ in habitat composition and fragmentation. We determined dispersal distances, pre- and post-dispersal habitat types and survival using a combination of capture–mark–recapture, radio-tracking and genetic parentage assignment. Most (75%) squirrels emigrated from the natal home range with mean dispersal distance of 1,014 ± 925 m (range 51–4,118 m). There were no sex-related differences in dispersal patterns and no differences in average dispersal distance, and the proportion of dispersers did not differ between sites. In one of the sites, dispersers settled in patches where density was lower than in the natal patch. In the least fragmented site, 90% of animals settled in the natal habitat type (habitat cuing) against 44–54% in the more strongly fragmented sites. Overall, more squirrels settled in the natal habitat type than expected based on habitat availability, but this was mainly due to individuals remaining within the natal wood. In the highly fragmented landscape, habitat cuing among emigrants did not occur more frequently than expected. We concluded that increased habitat fragmentation seemed to reduce reliable cues for habitat choice, but that dispersing squirrels settled in patches with lower densities of same-sex animals than at the natal home range or patch, independent of degree of fragmentation.     

Lifetime fitness of short- and long-distance dispersing great reed warblers

Dispersal is of prime importance for many evolutionary processes and has been studied for decades. The reproductive consequences of dispersal have proven difficult to study, simply because it is difficult to keep track of dispersing individuals. In most previous studies evaluating the fitness effects of dispersal, immigrants at a study locality have been lumped into one category and compared to philopatric individuals. This is unfortunate, because there are reasons to believe that immigrants with long and short dispersal distances may differ substantially in reproductive success. In the present study, we used a combination of capture-recapturing and multilocus microsatellite genotyping to categorize great reed warblers at our Swedish study site as philopatric individuals or short- or long-distance dispersing immigrants. We then performed novel comparisons of lifetime reproductive success (LRS) and survival rates of these three dispersal categories. The birds belonged to cohorts 1987-1996, and data for their LRS were gathered between 1988 and 2003. The analyses showed that philopatric males attracted more females, produced more fledglings and recruits throughout their lives, and survived better than immigrants. Among the immigrant males, those categorized as long-distance dispersers had lowest LRS and survival probability. Models that included covariates of potential importance showed that the difference in LRS between dispersal categories was partly caused by corresponding variation in number of breeding years at our study site. These results indicate that short- and, in particular, long-distance dispersers were of poor phenotypic quality, but it may also be proposed that immigrants attracted few females because they were poorly adapted to the local social environment. In females, the number of local recruits corrected for the number of breeding years (as well as for number of fledglings) differed between dispersal categories in a pattern that suggests an intermediate optimal dispersal distance. Short-distance dispersers recruited more offspring per year (and per fledgling) than both philopatric individuals and long-distance dispersers. Data suggest that the low LRS of philopatric females was related to costs of inbreeding. The low LRS of long-distance dispersing females may have resulted from their offspring being especially prone to disperse outside the study area, but also other potential explanations exist, such as local maladaptation. Our study highlights the importance of separating immigrant birds on the basis of their genetic similarity to the local study population when analyzing variation in LRS and inferring realized gene flow.     

Density‐dependent fitness,not dispersal movements,drives temporal variation in spatial genetic structure in dark‐eyed juncos (Junco hyemalis)

Some studies have found that dispersal rates and distances increase with density, indicating that density‐dependent dispersal likely affects spatial genetic structure. In an 11‐year mark–recapture study on a passerine, the dark‐eyed junco, we tested whether density affected dispersal distance and/or fine‐scale spatial genetic structure. Contrary to expectations, we found no effect of predispersal density on dispersal distance or the proportion of locally produced juveniles returning to the population from which they hatched. However, even though density did not affect dispersal distance or natal return rates, we found that density still did affect spatial genetic structure. We found significant positive spatial genetic structure at low densities of (postdispersal) adults but not at high densities. In years with high postdispersal (adult) densities that also had high predispersal (juvenile) densities in the previous year, we found negative spatial genetic structure, indicating high levels of dispersal. We found that density also affected fitness of recruits, and fitness of immigrants, potentially linking these population parameters with the spatial genetic structure detected. Immigrants and recruits rarely nested in low postdispersal density years. In contrast, in years with high postdispersal density, recruits were common and immigrants had equal success to local birds, so novel genotypes diluted the gene pool and effectively eliminated positive spatial genetic structure. In relation to fine‐scale spatial genetic structure, fitness of immigrants and new recruits is poorly understood compared to dispersal movements, but we conclude that it can have implications for the spatial distribution of genotypes in populations.     

Genomics meets applied ecology: Characterizing habitat quality for sloths in a tropical agroecosystem

Understanding how habitat quality in heterogeneous landscapes governs the distribution and fitness of individuals is a fundamental aspect of ecology. While mean individual fitness is generally considered a key to assessing habitat quality, a comprehensive understanding of habitat quality in heterogeneous landscapes requires estimates of dispersal rates among habitat types. The increasing accessibility of genomic approaches, combined with field‐based demographic methods, provides novel opportunities for incorporating dispersal estimation into assessments of habitat quality. In this study, we integrated genomic kinship approaches with field‐based estimates of fitness components and approximate Bayesian computation (ABC) procedures to estimate habitat‐specific dispersal rates and characterize habitat quality in two‐toed sloths (Choloepus hoffmanni) occurring in a Costa Rican agricultural ecosystem. Field‐based observations indicated that birth and survival rates were similar in a sparsely shaded cacao farm and adjacent cattle pasture–forest mosaic. Sloth density was threefold higher in pasture compared with cacao, whereas home range size and overlap were greater in cacao compared with pasture. Dispersal rates were similar between the two habitats, as estimated using ABC procedures applied to the spatial distribution of pairs of related individuals identified using 3,431 single nucleotide polymorphism and 11 microsatellite locus genotypes. Our results indicate that crops produced under a sparse overstorey can, in some cases, constitute lower‐quality habitat than pasture–forest mosaics for sloths, perhaps because of differences in food resources or predator communities. Finally, our study demonstrates that integrating field‐based demographic approaches with genomic methods can provide a powerful means for characterizing habitat quality for animal populations occurring in heterogeneous landscapes.     

Population dynamics of an expanding passerine at the distribution margin

Individuals may be maladapted to novel environments at the species’ distribution margin. We investigated population dynamics in a marginal habitat where reproduction has been proven poor. Survival, population growth rate (λ) and its components, breeding and natal dispersal were studied in great tits Parus major breeding at the northern margin of its distribution in northern Finland. We used long term capture–mark–recapture data sets. Study area size and population density were used to explain adult survival rates. The average annual estimates of adult survival rose from 0.371 to 0.388 between the periods of 1971–1984 and 1999–2009. The estimates are slightly lower than estimates of small passerines in Europe. Low local survival rate of fledglings (0.050–0.055) probably reflects intensified emigration from this low quality area. Temporal variation in λ was large (0.498–1.856). Despite of low adult survival and recruitment rates, the mean estimates of λ (1.008 and 1.033) indicate an overall stability in the population size. Indeed, our results suggest that the immigration has an important role in the population dynamics of northern great tits. Thus the population is demographically and genetically dependent on core habitats which may cause adaptive problems due to intensive gene flow. Given those limitations, options for evolution of local adaptations in northern distribution margins are discussed.     

Population dynamics in two-patch environments: Some anomalous consequences of an optimal habitat distribution

The effect of dispersal on population size and stability is explored for a population that disperses passively between two discrete habitat patches. Two basic models are considered. In the first model, a single population experiences density-dependent growth in both patches. A graphical construction is presented which allows one to determine the spatial pattern of abundance at equilibrium for most reasonable growth models and rates of dispersal. It is shown under rather general conditions that this equilibrium is unique and globally stable. In the second model, the dispersing population is a food-limited predator that occurs in both a source habitat (which contains a prey population) and a sink habitat (which does not). Passive dispersal between source and sink habitats can stabilize an otherwise unstable predator-prey interaction. The conditions allowing this are explored in some detail. The theory of optimal habitat selection predicts the evolutionarily stable distribution of a population, given that individuals can freely move among habitats so as to maximize individual fitness. This theory is used to develop a heuristic argument for why passive dispersal should always be selectively disadvantageous (ignoring kin effects) in a spatially heterogeneous but temporally constant environment. For both the models considered here, passive dispersal may lead to a greater number of individuals in both habitats combined than if there were no dispersal. This implies that the evolution of an optimal habitat distribution may lead to a reduction in population size; in the case of the predator-prey model, it may have the additional effect of destabilizing the interaction. The paper concludes with a discussion of the disparate effects habitat selection might have on the geographical range occupied by a species.     

Evolutionary consequences of asymmetric dispersal rates

We study the consequences of asymmetric dispersal rates (e.g., due to wind or current) for adaptive evolution in a system of two habitat patches. Asymmetric dispersal rates can lead to overcrowding of the "downstream" habitat, resulting in a source-sink population structure in the absence of intrinsic quality differences between habitats or can even cause an intrinsically better habitat to function as a sink. Source-sink population structure due to asymmetric dispersal rates has similar consequences for adaptive evolution as a source-sink structure due to habitat quality differences: natural selection tends to be biased toward the source habitat. We demonstrate this for two models of adaptive evolution: invasion of a rare allele that improves fitness in one habitat but reduces it in the other and antagonistic selection on a quantitative trait determined by five additive loci. If a habitat can sustain a population without immigration, the conditions for adaptation to that habitat are most favorable if there is little or no immigration from the other habitat; the influence of emigration depends on the magnitude of the allelic effects involved and other parameters. If, however, the population is initially unable to persist in a given habitat without immigration, our model predicts that the population will be most likely to adapt to that habitat if the dispersal rates in both directions are high. Our results highlight the general message that the effect of gene flow upon local adaptation should depend profoundly on the demographic context of selection.     

Habitat patch size alters the importance of dispersal for species diversity in an experimental freshwater community

Increased dispersal of individuals among discrete habitat patches should increase the average number of species present in each local habitat patch. However, experimental studies have found variable effects of dispersal on local species richness. Priority effects, predators, and habitat heterogeneity have been proposed as mechanisms that limit the effect of dispersal on species richness. However, the size of a habitat patch could affect how dispersal regulates the number of species able to persist. We investigated whether habitat size interacted with dispersal rate to affect the number of species present in local habitats. We hypothesized that increased dispersal rates would positively affect local species richness more in small habitats than in large habitats, because rare species would be protected from demographic extinction. To test the interaction between dispersal rate and habitat size, we factorially manipulated the size of experimental ponds and dispersal rates, using a model community of freshwater zooplankton. We found that high‐dispersal rates enhanced local species richness in small experimental ponds, but had no effect in large experimental ponds. Our results suggest that there is a trade‐off between patch connectivity (a mediator of dispersal rates) and patch size, providing context for understanding the variability observed in dispersal effects among natural communities, as well as for developing conservation and management plans in an increasingly fragmented world.     

Emigration to new habitats by voles: the cost of dispersal paradox

We tested the hypothesis that dispersal and philopatry are components of a mixed evolutionarily stable strategy (ESS). The hypothesis predicts that fitness of dispersers should be equal to that of philopatric individuals. Alternatively, fitness of dispersers could be lower (the resident fitness hypothesis) or greater (the cost of dispersal hypothesis) than that of philopatric individuals. We compared fitness of individuals that moved to new habitats (emigrants) and those that remained within habitat boundaries (residents) in populations of the prairie vole, Microtus ochrogaster, and the meadow vole, M. pennsylvanicus. We established vole populations in four enclosures (). Within each enclosure, voles were free to move between four types of habitats that varied in the availability of supplemental food and the amount of vegetative cover. We analysed two fitness components: the survival rates of all individuals, and pregnancy rates of females. Our study showed that emigrants generally had greater fitness than residents and that the difference in fitness was habitat dependent (i.e. was greater when individuals were emigrating from low-quality habitats than from high-quality habitats). High-food, high-cover habitats were the only habitat types for which fitness of emigrants was lower than that of residents. Similar patterns occurred in both prairie voles and meadow voles. Our results support the cost of dispersal hypothesis.     

Source–sink dynamics: how sinks affect demography of sources

Models of source–sink population dynamics have to make assumptions about whether, and eventually how, demographic parameters in source habitats are dependent on the demography in sink habitats. However, the empirical basis for making such assumptions has been weak. Here we report a study on experimental root vole populations, where estimates of demographic parameters were contrasted between source patches in source–sink (treatment) and source–source systems (control). In the presence of a sink patch (simulated by a pulsed removal of immigrants), source‐patch populations failed to increase over the breeding season, mainly due to a high spatially density‐dependent dispersal rate from source to sink patches. The per capita recruitment rate was almost two times higher in source–sink than in the source–source systems, but this did not compensate for the loss rate due to dispersal from source to sink patches. Sex ratio in the source–sink systems became less female biased, probably as a result of an enhanced frequency of dispersal movements in females. Good knowledge of the degree of density‐and habitat‐dependent dispersal is critical for predicting the dynamics of source–sink populations.     

Interactions between habitat quality and connectivity affect immigration but not abundance or population growth of the butterfly, Parnassius smintheus

Habitat geometry has been a primary focus in studies of spatially structured systems. Recent studies have indicated that a more comprehensive approach including habitat quality may be needed, however most previous studies have neglected potential interactions between quality and geometry. We investigated the effects of habitat quality for the butterfly Parnassius smintheus among a series of 17 sub‐populations. Specifically, we examined how habitat connectivity and local nectar flower density affect dispersal, and local population abundance and growth. We first determined which flower species were potentially important by examining nectar flower electivity and then quantified nectar flower density in meadows over a five year period (2003–2007). These data along with meadow connectivity were compared to local population statistics derived from mark–recapture over the same time period. The number of immigrants to a meadow increased as meadow connectivity increased, but showed no direct relationship with nectar flower density; however, there was a significant interaction between meadow connectivity and nectar flower density such that meadows with high connectivity and a high density of nectar flowers received the greatest number of immigrants. The number of emigrants from a meadow increased with increasing habitat quality and connectivity, but showed no interactive effect. The abundance of butterflies increased with meadow connectivity, but showed no relationship with habitat quality or any interactive effect. Separate experiments showed that access to nectar flowers significantly increased female reproductive output, but not lifespan. Despite the effects on immigration and reproductive output, local population growth rates also showed no relationship to nectar flower density. Our results indicate that habitat quality can be important for immigration in spatially structured populations; however, effects of habitat quality may not necessarily translate into higher abundance or population growth. Additionally, habitat quality should not be considered independently from habitat isolation, particularly if it directly affects dispersal. Preserving or augmenting habitat quality will do little to bolster immigration or colonization without adequate connectivity.     

Sex ratio evolution when fitness and dispersal vary

In a heterogeneous environment, when the fitness of males and females are differently influenced by habitat quality, habitat-dependent sex ratios may evolve to favor the production of the sex that benefits more (or loses less) from the local habitat. Similarly, sex-biased dispersal favors the evolution of habitat-dependent sex ratios. The present study documents the convergence stable sex ratios expected in the presence of sex-specific fitness gains when dispersal is partial, sex-biased or costly, using a simple model with patches of two qualities. Results show that partial dispersal reduces the sex ratio bias expected with sex-specific fitness gains. The direction of the sex ratio bias can be reversed by sex-biased dispersal or the existence of sex-specific dispersal costs, provided that fitness gains for the two sexes are not too different. The reversal of the sex ratio bias is more readily observed when sex-specific dispersal rates are opposite and extreme. Both dispersal and fitness gains, especially when they are sex-specific, should thus be considered when making predictions about sex ratio evolution in a heterogeneous environment.     

Factors affecting the local abundance of two anemonefishes (Amphiprion frenatus and A. perideraion) around a semi-closed bay in Puerto Galera,the Philippines

Many marine organisms disperse or migrate among habitats, which affects their abundance patterns at individual local habitats. To clarify the factors affecting the distribution patterns of two anemonefishes (Amphiprion frenatus and A. perideraion), we measured the habitat patch size (anemone size), patch isolation (mean distance from other anemones), presence/absence of other anemonefish species, depth, and abundance of the two anemonefishes at each anemone around a semi-closed bay (up to 3.7 km) in Puerto Galera, the Philippines. We assumed that local abundance increases with habitat size and decreases with patch isolation because of greater resource availability and reduced rates of recruitment from other patches. Local abundance of A. frenatus was related to habitat size and the presence of other anemonefish species, whereas that of A. perideraion was affected by the presence of other anemonefish species and water depth. Interspecific competition and/or niche differentiation of habitat can explain the negative relationship between the local abundance of the target species and other anemonefish. Patch isolation was not significant for both species probably because the dispersal rate was not directly proportional to the geographic distance between patches at our study site.