Niches in the Anthropocene: passerine assemblages show niche expansion from natural to urban habitats

Human‐mediated habitat transformation is increasingly evident around the world. Yet, how this transformation influences species’ niche width and overlap remains unclear. On the one hand, human‐mediated habitat transformation promotes increased species similarity through trait‐based filtering, and an increased prevalence of generalist species with broad niches, resulting in functional homogenization. On the other hand, species that colonize transformed habitats could use empty niches, resulting in decreased species similarity and an expansion of assemblage‐level niche space. Here we explore these two alternatives in eight highly diverse passerine assembles in natural, rural and urban habitats in south and southwest China, a rapidly developing region of the world. Based on stable isotopes, we found that species’ niche width increased from natural to human‐made habitats, but there were no differences in niche overlap among habitats. Therefore, we found evidence for niche expansion, with generalists appearing to use empty niches created by human habitat modification, and with assemblages being comprised of complementary species. Further research is needed to determine whether increased between‐ or within‐individual niche variation is the main driver of niche expansion in transformed habitats.     

Reduced dispersal propensity in the wingless waterstrider Aquarius najas in a highly fragmented landscape

Dispersal behaviour of animals in fragmented habitats has generated intensive theoretical attention but empirical data on the evolution of dispersal are still relatively scarce. Theory predicts reduced dispersal propensity in small and isolated habitat patches. We tested these predictions in the waterstrider Aquarius najas, a wingless species with special habitat demands. Flightlessness constrains insect dispersal and as a stream specialist A. najas cannot survive on still water. Lakes therefore represent a dispersal barrier for this species. We measured dispersal propensity of the waterstrider A. najas which originated from ten fragmented populations. In the experiment, we transplanted laboratory-grown individuals to the field. We did not find differences between sex in dispersal propensity. However, we found that waterstriders that originated from small and isolated patches moved less than individuals from large and more continuous habitats. This suggests that the cost of dispersal over hostile surrounding habitats may be high. We conclude that a low likelihood of dispersal in A. najas is an adaptation to small and isolated stream habitats.     

Understanding the impact of flooding on trait-displacements and shifts in assemblage structure of predatory arthropods on river banks

1. Species assemblages of naturally disturbed habitats are governed by the prevailing disturbance regime. Consequently, stochastic flood events affect river banks and the inhabiting biota. Predatory arthropods occupy predominantly river banks in relation to specific habitat conditions. Therefore, species sorting and stochastic processes as induced by flooding are supposed to play important roles in structuring riparian arthropod assemblages in relation to their habitat preference and dispersal ability. 2. To ascertain whether assemblages of spiders and carabid beetles from disturbed river banks are structured by stochastic or sorting mechanisms, diversity patterns and assemblage-wide trait-displacements were assessed based on pitfall sampling data. We tested if flooding disturbance within a lowland river reach affects diversity patterns and trait distribution in both groups. 3. Whereas the number of riparian spider species decreased considerably with increased flooding, carabid beetle diversity benefited from intermediate degrees of flooding. Moreover, regression analyses revealed trait-displacements, reflecting sorting mechanisms particularly for spiders. Increased flooding disturbance was associated with assemblage-wide increases of niche breadth, shading and hygrophilic preference and ballooning propensity for spider (sub)families. Trait patterns were comparable for Bembidiini carabids, but were less univocal for Pterostichini species. Body size decreased for lycosid spiders and Bembidiini carabids with increased flooding, but increased in linyphiid spiders and Pterostichini carabids. 4. Our results indicate that mainly riparian species are disfavoured by either too high or too low degrees of disturbance, whereas eurytopic species benefit from increased flooding. Anthropogenic alterations of flooding disturbance constrain the distribution of common hygrophilous species and/or species with high dispersal ability, inducing shifts towards less specialized arthropod assemblages. River banks with divergent degrees of flooding impact should be maintained throughout dynamic lowland river reaches in order to preserve typical riparian arthropod assemblages.     

Ballooning propensity of canopy and understorey spiders in a mature temperate hardwood forest

1. Spiders frequently disperse and colonise habitats through ballooning, a passive aerial dispersal process. Ballooning is pre‐eminent in open habitat spider communities and its propensity can be modulated by habitat conditions and availability, and by life‐history traits such as body size, degree of specialisation, and feeding behaviour. 2. Using spiders from the canopy and understorey of a north‐temperate hardwood forest as a model system, our main objectives were to detect if foliage spiders of a mature forest disperse through ballooning, and identify life‐history traits that influence ballooning propensity. 3. Our results demonstrate that foliage spiders living in the canopy and understorey of a mature forest do balloon, and in some cases have very high ballooning propensities similarly to open field spiders. Species level models showed that small body size had a strong positive effect on ballooning for juveniles of species with large‐bodied adults, while individuals of small‐bodied species initiated ballooning regardless of size, habitat or development stage. A generalised linear mixed model indicated that small size web‐building spiders from the Retro Tibial Apophysis (RTA) and Orbicularia clades had the highest propensity for foliage spiders of this north‐temperate hardwood forest. 4. In conclusion, we provide the first demonstration that forest spiders can have high ballooning propensities and that ballooning propensity is negatively affected by body size and positively affected by the prominent use of silk to catch prey. However, spiders originating from the canopy and understorey of a north‐temperate hardwood forest did not differ in their ballooning propensities.     

Aerial dispersal ability does not drive spider success in a crop landscape

1. Although spiders can colonise ecosystems by air, dispersal capabilities differ among spider species. Web‐building spiders are thought to balloon at higher rates than hunting spiders. Spider success in agricultural systems may also depend on habitat preferences. Few studies have examined the success of aerially dispersing spiders in crop systems, and information about the dispersal capabilities of spiders in putative source habitats is limited. 2. Spiders were monitored in the air and on the foliage of vineyards and adjacent oak woodland in order to compare the aerial spider faunas between these disparate habitats and to determine whether highly dispersive species contributed disproportionately to the spider community in vineyards. 3. The results show that most aerially dispersing spiders in both habitats were web‐building dwarf spiders, Erigone spp. (Linyphiidae), although hunting spiders were also well represented in the air, especially in oak woodland. Most woodland spiders in the air appeared to be residents of oak woodland and probably dispersed only short distances. 4. Conversely, only a subset of the aerial spider fauna established in vineyards in high numbers. Spiders that dominated the aerial fauna were under‐represented on vineyard foliage, whereas several hunting spiders dispersed aerially at low rates but dominated vineyard spider composition. 5. These results suggest that aerial dispersal ability may allow spiders to reach crop systems, but that establishment depends on habitat preferences and/or competitive ability.     

Metacommunities of spiders in grassland habitat fragments of an agricultural landscape

Arthropod communities in fragmented agricultural landscapes depend on local processes and the interactions between communities in the habitat islands. We aimed to study metacommunity structure of spiders, a group that is known for high dispersal power, local niche partitioning and for engaging in species interactions. While living in fragmented habitats could lead to nestedness, other ecological traits of spiders might equally lead to patterns dominated either by species interactions or habitat filtering. We asked, which community pattern will prevail in a typical agricultural landscape with isolated patches of semi-natural habitats. Such a situation was studied by sampling spiders in 28 grassland locations in a Hungarian agricultural landscape. We used the elements of metacommunity structure (EMS) framework to distinguish between alternative patterns that reveal community organization. The EMS analysis indicated coherent species ranges, high turnover and boundary clumping, suggesting Clementsian community organization. The greatest variation in species composition was explained by local habitat characteristics, indicating habitat filtering. The influence of dispersal could be detected by the significant effect of landscape composition, which was strongest at 500 m. We conclude that dispersal allows spiders to respond coherently to the environment, creating similar communities in similar habitats. Consistent habitat differences, such as species rich versus species poor vegetation, lead to recognisably different, recurrent communities. These characteristics make spiders a predictable and diverse source of natural enemies in agricultural landscapes. Sensitivity to habitat composition at medium distances warns us that landscape homogenization may alter these metacommunity processes.     

Central European plant species from more productive habitats are more invasive at a global scale

Aim Accumulating evidence indicates that species may be pre‐adapted for invasion success in new ranges. In the light of increasing global nutrient accumulation, an important candidate pre‐adaptation for invasiveness is the ability to grow in nutrient‐rich habitats. Therefore we tested whether globally invasive species originating from Central Europe have come from more productive rather than less productive habitats. A further important candidate pre‐adaptation for invasiveness is large niche width. Therefore, we also tested whether species able to grow across habitats with a wider range of productivity are more invasive. Location Global with respect to invasiveness, and Central European with respect to origin of study species. Methods We examined whether average habitat productivity and its width across habitats are significant predictors of the success of Central European species as aliens and as weeds elsewhere in the world based on data in the Global Compendium of Weeds. The two habitat productivity measures were derived from nutrient indicator values (after Ellenberg) of accompanying species present in vegetation records of the comprehensive Czech National Phytosociological Database. In the analyses, we accounted for phylogenetic relatedness among species and for size of the native distribution ranges. Results Species from more productive habitats and with a wider native habitat‐productivity niche in Central Europe have higher alien success elsewhere in the world. Weediness of species increased with mean habitat productivity. Niche width was also an important determinant of weediness for species with their main occurrence in nutrient‐poor habitats, but not for those from nutrient‐rich habitats. Main conclusions Our results indicate that Central European plant species from productive habitats and those species from nutrient‐poor habitat with wide productivity‐niche are pre‐adapted to become invasive. These results suggest that the world‐wide invasion success of many Central European species is likely to have been promoted by the global increase of resource‐rich habitats.     

Landscape genetics of an early successional specialist in a disturbance‐prone environment

Species that specialize in disturbed habitats may have considerably different dispersal strategies than those adapted to more stable environments. However, little is known of the dispersal patterns and population structure of such species. This information is important for conservation because many postfire specialists are at risk from anthropogenic changes to natural disturbance regimes. We used microsatellite markers to assess the effect of landscape variation and recent disturbance history on dispersal by a small mammal species that occupies the early seral stage of vegetation regeneration in burnt environments. We predicted that a postfire specialist would be able to disperse over multiple habitat types (generalist) and not exhibit sex‐biased dispersal, as such strategies should enable effective colonization of spatially and temporally variable habitat. We found significant differentiation between sites that fitted an isolation‐by‐distance pattern and spatial autocorrelation of multilocus genotypes to a distance of 2–3 km. There was no consistent genetic evidence for sex‐biased dispersal. We tested the influence of different habitat‐ and fire‐specific landscape resistance scenarios on genetic distance between individuals and found a significant effect of fire. Our genetic data supported recently burned vegetation having greater conductance for gene flow than unburnt habitat, but variation in habitat quality between vegetation types and occupied patches had no effect on gene flow. Postfire specialists must evolve an effective dispersal ability to move over distances that would ensure access to early successional stage vegetation. Natural disturbance and natural heterogeneity may therefore not influence population genetic structure as negatively as expected.     

Contrasting responses of arable spiders to the landscape matrix at different spatial scales

Aim Animal communities can be influenced by the composition of the surrounding landscape through immigration. Depending on habitat preferences, however, the effect of the landscape matrix can be positive or negative and can vary with scale. We tested this idea with arable spiders and tried to infer dispersal distances from relationships between local density and landscape composition at different spatial scales. Location Thirty‐eight landscapes around the cities of Göttingen and Giessen, Germany. Methods Spiders were captured with pitfall traps in one field of winter wheat in each landscape. Landscape composition around the fields was characterized at 11 scales from 95 m to 3 km radius by land‐use mapping and subsequent GIS analysis. Correlation tests were performed between landscape composition and local densities or species richness. Results In both study regions, local species richness was enhanced by non‐crop habitats on a landscape scale. The overall densities of wolf spiders (Lycosidae), long‐jawed spiders (Tetragnathidae), crab spiders (Thomisidae), and dwarf sheet spiders (Hahniidae) increased significantly in landscapes with high percentages of non‐crop habitats. Out of the 40 species tested, 19 responded positively to the percentage of non‐crop habitats in the surrounding landscape, and five responded negatively. Depending on the species, the spatial scales with the highest explanatory power ranged from 95 m to 3 km radius around the study fields, potentially reflecting dispersal distances. Main conclusions Arable spider species showed contrasting responses to the landscape context with respect both to the direction and to the spatial scale of the relationship. The variation in landscape requirements among species ensures high spider densities in a wide range of situations, which contributes to ecosystem resilience. However, species richness of arable spiders depends on heterogeneous landscapes with high percentages of non‐crop habitats.     

Niche properties of Central European spiders: shading, moisture and the evolution of the habitat niche

Aim  Niche theory emphasizes the importance of environmental conditions for the distribution and abundance of species. Using a macroecological approach our study aimed at identifying the important environmental gradients for spiders. We generated numerical values of niche position and niche width. We also investigated relationships between these niche properties as well as the degree of phylogenetic conservatism in order to draw conclusions about the evolution of the habitat niche.
Location  Central Europe: lowlands of Austria, Belgium, Germany, Luxembourg, the Netherlands and Switzerland.
Methods  We analysed 244 published spider communities from 70 habitat types by correspondence analysis. The resulting community scores were used to test for correlations with habitat characteristics. Species scores were used to derive niche position (mean scores) and niche width (standard deviation of scores). To test for niche conservatism we estimated variance components across the taxonomic hierarchy.
Results  The first two axes of the correspondence analysis were correlated with shading and moisture, respectively. Niche width had a hump-shaped relationship to both environmental gradients. β-diversity was strikingly higher in open habitats than in forests. Habitat niche conservatism was lower than phylogenetic conservatism in body size.
Main conclusions  Environmental factors are important drivers for the β-diversity of spiders, especially across open habitats. This underlines the importance of preserving the whole range of moisture conditions in open habitats. Narrow niches of species occurring at the ends of both environmental gradients indicate that adaptations to extreme habitats lead to constraints in ecological flexibility. Nevertheless, the habitat niche of species seems to evolve much faster than morphological or physiological traits.     

Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types

Aim   We analysed the variation of species richness in the European freshwater fauna across latitude. In particular, we compared latitudinal patterns in species richness and β-diversity among species adapted to different habitat types.
Location   Europe.
Methods   We compiled data on occurrence for 14,020 animal species across 25 pre-defined biogeographical regions of European freshwaters from the Limnofauna Europaea . Furthermore, we extracted information on the habitat preferences of species. We assigned species to three habitat types: species adapted to groundwater, lotic (running water) and lentic (standing water) habitats. We analysed latitudinal patterns of species richness, the proportion of lentic species and β-diversity.
Results   Only lentic species showed a significant species–area relationship. We found a monotonic decline of species richness with latitude for groundwater and lotic habitats, but a hump-shaped relationship for lentic habitats. The proportion of lentic species increased from southern to northern latitudes. β-Diversity declined from groundwater to lentic habitats and from southern to northern latitudes.
Main conclusions   The differences in the latitudinal variation of species richness among species adapted to different habitat types are in part due to differences in the propensity for dispersal. Since lentic habitats are less persistent than lotic or groundwater habitats, lentic species evolved more efficient strategies for dispersal. The dispersal propensity of lentic species facilitated the recolonization of central Europe after the last glaciation. Overall, we stress the importance of considering the history of regions and lineages as well as the ecological traits of species for understanding patterns of biodiversity.     

Spatial sorting may explain evolutionary dynamics of wing polymorphism in pygmy grasshoppers

Wing polymorphism in insects provides a good model system for investigating evolutionary dynamics and population divergence in dispersal‐enhancing traits. This study investigates the contribution of divergent selection, trade‐offs, behaviour and spatial sorting to the evolutionary dynamics of wing polymorphism in the pygmy grasshopper Tetrix subulata (Tetrigidae: Orthoptera). We use data for > 2800 wild‐caught individuals from 13 populations and demonstrate that the incidence of the long‐winged (macropterous) morph is higher and changes faster between years in disturbed habitats characterized by succession than in stable habitats. Common garden and mother‐offspring resemblance studies indicate that variation among populations and families is genetically determined and not influenced to any important degree by developmental plasticity in response to maternal condition, rearing density or individual growth rate. Performance trials show that only the macropterous morph is capable of flight and that propensity to fly differs according to environment. Mark–recapture data reveal no difference in the distance moved between free‐ranging long‐ and short‐winged individuals. There is no consistent difference across populations and years in number of hatchlings produced by long‐ and shorter‐winged females. Our findings suggest that the variable frequency of the long‐winged morph among and within pygmy grasshopper populations may reflect evolutionary modifications driven by spatial sorting due to phenotype‐ and habitat type–dependent emigration and immigration.     

HOW DOES POLLEN VERSUS SEED DISPERSAL AFFECT NICHE EVOLUTION?

In heterogeneous landscapes, the genetic and demographic consequences of dispersal influence the evolution of niche width. Unless pollen is limiting, pollen dispersal does not contribute directly to population growth. However, by disrupting local adaptation, it indirectly affects population dynamics. We compare the effect of pollen versus seed dispersal on the evolution of niche width in heterogeneous habitats, explicitly considering the feedback between maladaptation and demography. We consider two scenarios: the secondary contact of two subpopulations, in distinct, formerly isolated habitats, and the colonization of an empty habitat with dispersal between the new and ancestral habitat. With an analytical model, we identify critical levels of genetic variance leading to niche contraction (secondary contact scenario), or expansion (new habitat scenario). We confront these predictions with simulations where the genetic variance freely evolves. Niche contraction occurs when habitats are very different. It is faster as total gene flow increases or as pollen predominates in overall gene flow. Niche expansion occurs when habitat heterogeneity is not too high. Seed dispersal accelerates it, whereas pollen dispersal tends to retard it. In both scenarios very high seed dispersal leads to extinction. Overall, our results predict a wider niche for species dispersing seeds more than pollen.     

On humans and wildlife in Mediterranean islands

Aim To investigate the effects of human‐induced landscape changes in Mediterranean islands on the ecological and evolutionary responses of bird communities and populations. The combination of mass extinction of large mammals and massive deforestation by humans was hypothesized to produce new selection regimes to which organisms were likely to respond. Habitat selection and niche breadth have been investigated at the scale of species, and phenotypic variation at the scale of local populations. Location The study was carried out along habitat gradients and in habitat mosaics at different spatial scales on the island of Corsica and in areas of similar size and structure in continental France. Methods Two sets of gradients have been used for investigating habitat selection and niche breadth: gradients of altitude, and gradients of vegetation structure. Population studies focused on the blue tit, Cyanistes caeruleus. Large samples of breeding attempts by this species in 10 habitats provided detailed data on phenotypic variation of fitness‐related traits both on Corsica and on the mainland. Results The extent of niche space used by birds differed substantially depending on which habitat gradient was considered. Many species have been found to contract their habitat niche along the elevation gradient on Corsica compared with the mainland, whereas all species in the vegetation gradient broadened their niche on the island. Breeding patterns of the blue tit differed considerably depending on whether they settle in deciduous oaks (Quercus humilis) or in evergreen sclerophyllous oaks (Quercus ilex). Phenotypic variation of breeding traits was much higher on the island, where more populations were correctly timed for the best breeding period than on the mainland, a pattern that is likely to result from lower dispersal of organisms on the island. Main conclusions The differences in observed niche breadth between the two series of habitat gradients is explained both by the species‐specific ecology of the species and the human‐induced environmental history of Corsica. Large‐scale landscape changes provided new opportunities for island colonization by non‐forest species, which are isolated as small, ‘fugitive’ local populations. In both gradients, forest species that are typical components of the Corsican bird fauna definitely expanded their niche and occupied a wider range of habitats on Corsica than on the mainland. At the population scale, landscapes included habitat patches with contrasted selection regimes, which resulted in high phenotypic variation for many fitness‐related traits. Reduced dispersal of birds on the island resulted in a much higher degree of local differentiation on Corsica than on the mainland.     

Response of ants to human‐altered habitats with reference to seed dispersal of the myrmecochore Corydalis giraldii Fedde (Papaveraceae)

Anthropogenic habitat disturbance has potential consequences for ant communities. However, there is limited information on the effects of ant responses on associated ecological processes such as seed dispersal. We investigated the effect of disturbance on the abundance, richness, and composition of ant communities and the resulting seed‐dispersal services for a herbaceous myrmecochore, Corydalis giraldii (Papaveraceae), in an undisturbed habitat (forest understory), moderately disturbed habitat (abandoned arable field), and highly disturbed habitat (road verge) on Qinling Mountains, China. In total, we recorded 13 ant species, and five out of these were observed to transport seeds. The community composition of dispersers was significantly different amongst habitats. The richness of the dispersers did not differ among the habitats, but their total abundance varied significantly across habitats and was 21% lower in the road verge than in the abandoned arable fields. The major seed‐dispersing ant species in both the forest understory and the abandoned arable field were large‐bodied (Myrmica sp. and Formica fusca, respectively), whereas the major seed‐dispersing ants found in the road verge were the small‐bodied Lasius alienus. This difference resulted in lower seed removal rates and dispersal distances in the road verge than in the other two habitats. The different dispersal patterns were attributed primarily to differences in dispersing ant abundance and identity, most likely in response to habitats with different degree of anthropogenic disturbance. The possible influence of disturbance on the ecological specialization of ant‐seed dispersal interaction is also discussed.     

The impact of hedge-forest connectivity and microhabitat conditions on spider and carabid beetle assemblages in agricultural landscapes

Agricultural intensification in terms of decreasing landscape complexity and connectivity has negatively affected biodiversity. Linear landscape elements composed of woody vegetation like hedges may counteract this negative trend by providing habitats and enhancing habitat connectivity for different organisms. Here, we tested the impacts of habitat type (forest edges vs. hedges) and hedges’ isolation (connected vs. isolated hedges) from forests as well as microhabitat conditions (percentage of bare ground and width) on trait-specific occurrence of ground-dwelling arthropods, namely spiders and carabids. Arthropods were grouped by habitat specialisation (forest vs. open-habitat species vs. generalists), hunting strategy (web-building or hunting spiders) and dispersal ability (wing morphology of carabids). Spider and carabid assemblage composition was strongly influenced by habitat type and isolation, but not by microhabitat conditions. Activity density of forest species and brachypterous carabids was higher in forest edges compared to hedges, whereas open-habitat species and macropterous carabids showed reverse patterns, with no effects of isolation. Occurrence of generalist carabids, but not spiders, was higher in hedges compared to forest edges. Habitat type and isolation did not affect spiders with different hunting strategy. Microhabitat conditions were less important for spider and carabid occurrence. Our study concludes that on a landscape scale, type of linear woody habitat is more important for arthropod occurrence than isolation effects and microhabitat conditions, depending on traits. Hedges provide refuges for species specialised to open habitats and species with high dispersal ability, such as macropterous carabids. Forest edges enhance persistence of species specialised to forests and species with low dispersal ability, such as brachypterous carabids.     

Overlap and partitioning of the ecological and isotopic niches

Recently, it was proposed that stable isotope patterns can be used to quantify the width of the ecological niche of animals. However, the potential effects of habitat use on isotopic patterns of consumers have not been fully explored and consequently isotopic patterns may yield deceptive estimates of niche width. Here, we simulated four different scenarios of a consumer foraging across an isotopically heterogeneous landscape to test the combined effects of habitat and diet selection on the widths of the isotopic niche. We then modeled the actions of a naïve researcher who randomly sampled consumers from the simulated populations, and used these results to assess the overlap and partitioning of the isotopic and the ecological niches when habitat‐derived differences among isotope signatures are not considered. Our results suggest that populations of dietary specialists exhibited broader isotopic niches than populations composed of dietary generalists, and habitat generalists exhibited narrower isotopic niche widths compared with populations of individuals that foraged in specific habitats. The conversion of isotopic niches to ecological niches without knowledge of foraging behavior and habitat‐derived isotopic differences transformed an informative δ‐space into ‘a blurry p‐space’. Therefore, knowledge of habitat‐derived differences in stable isotope values and understanding of habitat use and individual foraging behavior are critical for the correct quantification of the ecological niche.     

A spatially explicit model of sex ratio evolution in response to sex-biased dispersal

Sex-biased dispersal occurs in all seed plants and many animal species. Theoretical models have shown that sex-biased dispersal can lead to evolutionarily stable biased sex ratios. Here, we use a spatially explicit chessboard model to simulate the evolution of sex ratio in response to sex-biased dispersal range and sex-biased dispersal rate. Two life cycles are represented in the model: one in which both sexes disperse before mating (DDM), the other in which males disperse before mating and mated females or zygotes disperse after mating (DMD). Model parameters include factors like dispersal rate, dispersal range, number of individuals per patch, and habitat heterogeneity.When dispersal range is sex biased, we find that, in a homogeneous environment, the sex ratio is generally biased towards the sex that disperses more widely (sex ratio range: 0.47–0.52). In a heterogeneous environment, the sex ratio is generally biased towards the more dispersive sex in good habitats, and towards the less dispersive sex in poor habitats (sex ratio range: 0–1). This is opposite to the effect of sex-biased dispersal rate, which favours the production of the more dispersive sex in poor habitats and the less dispersive sex in good habitats (sex ratio range: 0–1). To allow for a comparison with theoretical predictions, data concerning sex-biased dispersal and habitat-dependent sex ratios should thus incorporate information about the spatial scale of both dispersal and environmental heterogeneity.     

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.     

Habitat type predicts genetic population differentiation in freshwater invertebrates

A basic challenge in evolutionary biology is to establish links between ecology and evolution of species. One important link is the habitat template. It has been hypothesized, that the spatial and temporal settings of a habitat strongly influence the evolution of species dispersal propensity. Here, we evaluate the importance of the habitat type on genetic population differentiation of species using freshwater habitats as a model system. Freshwater habitats are either lentic (standing) or lotic (running). On average, lotic habitats are more stable and predictable over space and time than lentic habitats. Therefore, lentic habitats should favour the evolution of higher dispersal propensity which ensures population survival of lentic species. To test for such a relationship, we used extensive data on species' genetic population differentiation of lentic and lotic freshwater invertebrates retrieved from published allozyme studies. Overall, we analysed more than 150 species from all over the world. Controlling for several experimental, biological and geographical confounding effects, we always found that lentic invertebrates exhibit, on average, lower genetic population differentiation than lotic species. This pattern was consistent across insects, crustaceans and molluscs. Our results imply fundamental differences in genetic population differentiation among species adapted to either lentic or lotic habitats. We propose that such differences should occur in a number of other habitat types that differ in spatio-temporal stability. Furthermore, our results highlight the important role of lotic habitats as reservoirs for evolutionary processes and the potential for rapid speciation.