Phenotypic and genetic differentiation among yellow monkeyflower populations from thermal and non-thermal soils in Yellowstone National Park

In flowering plants, soil heterogeneity can generate divergent natural selection over fine spatial scales, and thus promote local adaptation in the absence of geographic barriers to gene flow. Here, we investigate phenotypic and genetic differentiation in one of the few flowering plants that thrives in both geothermal and non-thermal soils in Yellowstone National Park (YNP). Yellow monkeyflowers (Mimulus guttatus) growing at two geothermal ("thermal") sites in YNP were distinct in growth form and phenology from paired populations growing nearby (<500?m distant) in non-thermal soils. In simulated thermal and non-thermal environments, thermal plants remained significantly divergent from non-thermal plants in vegetative, floral, mating system, and phenological traits. Plants from both thermal populations flowered closer to the ground, allocated relatively more to sexual reproduction, were more likely to initiate flowering under short daylengths, and made smaller flowers that could efficiently self-fertilize without pollinators. These shared differences are consistent with local adaptation to life in the ephemeral window for growth and reproduction created by winter and spring snowmelt on hot soils. In contrast, habitat type (thermal vs. non-thermal) explained little of the genetic variation at neutral markers. Instead, we found that one thermal population (Agrostis Headquarters; AHQ-T) was strongly differentiated from all other populations (all F (ST)?>?0.34), which were only weakly differentiated from each other (all F (ST)?相似文献   

Local genetic differentiation among populations of the mass-flowering tropical shrub Erythroxylum havanense (Erythroxylaceae)

This study is aimed to understand the role of life-history traits in determining the genetic structure of populations. We used Erythroxylum havanense, a distylous shrub with synchronous and massive patterns of flowering and fruiting. We suggest that the high concentration of ephemeral resources produced by mass flowering satiates both pollinators and frugivores, restricting gene flow and leading to genetic differentiation among populations. Using random amplified polymorphic DNAs as genetic markers, we estimated genetic diversity and structure statistics to quantify the amount and distribution of genetic variation within and among five populations from the Pacific coast of Mexico. High levels of genetic variation within populations and significant differentiation among populations located very near to each other were found. Furthermore, spatial autocorrelation analysis indicated the presence of significant genetic structure at short spatial distances. We suggest that by influencing the foraging behavior of pollinators and frugivores, mass flowering may produce the observed patterns of genetic structure, while small differences in flowering or fruiting phenology could further reinforce the isolation of nearby populations.     

Life-history variation in agricultural and wild populations of Erucastrum nasturtiifolium (Brassicaceae)

In the present study we relate the variability in life-history traits (such as flowering time and lifespan) of the herbaceous biennial–perennial Erucastrum nasturtiifolium (Brassicaceae) to habitat type. We studied plant populations from arable fields and from eroded mountain habitats, such as badlands and rocky slopes. Collection sites ranged from low basin to sub-alpine regions in the NE Iberian Peninsula. Plants were grown under common garden conditions to evaluate genetic variation among and within populations. Plants were also subjected to a resource gradient to detect genetic variation in phenotypic plasticity. The populations exhibited differentiation across a number of life-history traits (mainly flowering time and lifespan) and morphological traits related to growth (basal stem diameter, plant height and number of branches). This suggests that life-history differences among populations are genetically based. Moreover, our results show that variation in flowering time and lifespan are affected by habitat type independent of other abiotic factors such as altitude or continentality. Thus, populations from arable fields started flowering in their first year and displayed annual cycles, whereas those from wild habitats generally flowered in their second year and showed biennial or even perennial cycles. Intra-population differences in flowering time were observed in only one population, and were related to nutrient availability. We suggest that early-flowering and shorter lifespan populations of E. nasturtiifolium may have been selected from late-flowering and longer lifespan populations as part of a selective process ensuring survival and future offspring amidst unpredictable and frequently disturbed environments such as exist in many agricultural habitats.     

Divergent selection on flowering time contributes to local adaptation in Mimulus guttatus populations

The timing of when to initiate reproduction is an important transition in any organism's life cycle. There is much variation in flowering time among populations, but we do not know to what degree this variation contributes to local adaptation. Here we use a reciprocal transplant experiment to examine the presence of divergent natural selection for flowering time and local adaptation between two distinct populations of Mimulus guttatus. We plant both parents and hybrids (to tease apart differences in suites of associated parental traits) between these two populations into each of the two native environments and measure floral, vegetative, life-history, and fitness characters to assess which traits are under selection at each site. Analysis of fitness components indicates that each of these plant populations is locally adapted. We obtain striking evidence for divergent natural selection on date of first flower production at these two sites. Early flowering is favored at the montane site, which is inhabited by annual plants and characterized by dry soils in midsummer, whereas intermediate (though later) flowering dates are selectively favored at the temperate coastal site, which is inhabited by perennial plants and is almost continually moist. Divergent selection on flowering time contributes to local adaptation between these two populations of M. guttatus, suggesting that genetic differentiation in the timing of reproduction may also serve as a partial reproductive isolating barrier to gene flow among populations.     

Spatial autocorrelation analysis offers new insights into gene flow in the Australian bush rat,Rattus fuscipes

Dispersal is a fundamental process that influences the response of species to landscape change and habitat fragmentation. In an attempt to better understand dispersal in the Australian bush rat, Rattus fuscipes, we have combined a new multilocus autocorrelation method with hypervariable microsatellite genetic markers to investigate fine-scale (< or = 1 km) patterns of spatial distribution and spatial genetic structure. The study was conducted across eight trapping transects at four sites, with a total of 270 animals sampled. Spatial autocorrelation analysis of bush rat distribution revealed that, in general, animals occurred in groups or clusters of higher density (< or = 200 m across), with intervening gaps or lower density areas. Spatial genetic autocorrelation analysis, based on seven hypervariable microsatellite loci (He = 0.8) with a total of 80 alleles, revealed a consistent pattern of significant positive local genetic structure. This genetic pattern was consistent for all transects, and for adults and sub-adults, males and females. By testing for autocorrelation at multiple scales from 10 to 800 m we found that the extent of detectable positive spatial genetic structure exceeded 500 m. Further analyses detected significantly weaker spatial genetic structure in males compared with females, but no significant differences were detected between adults and sub adults. Results from Mantel tests and hierarchical AMOVA further support the conclusion that the distribution of bush rat genotypes is not random at the scale of our study. Instead, proximate bush rats are more genetically alike than more distant animals. We conclude that in bush rats, gene flow per generation is sufficiently restricted to generate the strong positive signal of local spatial genetic structure. Although our results are consistent with field data on animal movement, including the reported tendency for males to move further than females, we provide the first evidence for restricted gene flow in bush rats. Our study appears to be the first microsatellite-based study of fine-scale genetic variation in small mammals and the first to report consistent positive local genetic structure across sites, age-classes, and sexes. The combination of new forms of autocorrelation analyses, hypervariable genetic markers and fine-scale analysis (< 1 km) may thus offer new evolutionary insights that are overlooked by more traditional larger scaled (> 10 km) population genetic studies.     

Glacial survival and local adaptation in an alpine leaf beetle

The challenge in defining conservation units so that they represent evolutionary entities has been to combine both genetic properties and ecological significance. Here we make use of the complexity of the European Alps, with their genetic landscape shaped by geographical barriers and postglacial colonization, to examine the correlation between ecological and genetic divergence. Montane species, because of the fragmentation of their present habitat, constitute extreme cases in which to test if genetically distinct subgroups based on neutral markers are also ecologically differentiated and show local adaptation. In the leaf beetle Oreina elongata, populations show variation in host plant use and a patchy distribution throughout the Alps and Apennines. We demonstrate that despite very strong genetic isolation (F(ST) = 0.381), variation in host plant use has led to differences in larval life-history traits between populations only as a secondary effect of host defence chemistry, and not through physiological adaptation to plant nutritional value. We also establish that populations that are more ecologically different in terms of larval performance are also more genetically divergent. In addition, morphological variation used to define subspecies appears to be mirrored in the population genetics of this species, resulting in almost perfect clustering based on microsatellite data. Finally, we argue from their strong genetic structure and congruent distribution that the subspecies of O. elongata were divided among the same glacial refugia within the Alps that have been proposed for alpine plants.     

Phenotypic plasticity allows the Mediterranean parsley frog Pelodytes punctatus to exploit two temporal niches under continuous gene flow

Environmental changes, such as climate change, lead to the opening of new niches. In such situations, species that adapt to new niches can survive and/or expand their ranges. However, gene flow can hamper genetic adaptation to new environments. Alternatively, recent models have highlighted the importance of phenotypic plasticity in tracking environmental change. Here, we investigate whether phenotypic plasticity or genetic evolution (or both) allows an amphibian species to exploit two divergent climatic niches. In the Mediterranean region, the parsley frog Pelodytes punctatus breeds both in spring, as do most other species, and in autumn, a temporal niche not exploited by most other species, but which may become increasingly important with global warming. Conditions of development are dramatically different between the two seasons and deeply impact tadpole life-history traits. To determine whether these temporal niches are exploited by two genetically differentiated subpopulations, or whether the bimodal phenology arises in a panmictic population displaying plastic life-history traits, we use two complementary approaches. We measure both molecular genetic differentiation and quantitative-trait differentiation between spring and autumn cohorts, using microsatellites and common garden experiments, respectively. Seasonal cohorts were not genetically differentiated and differences in tadpole life history between cohorts were not maintained in laboratory conditions. We conclude that phenotypic plasticity, rather than genetic adaptation, allows Parsley frog to exploit two contrasting temporal niches.     

POPULATION STRUCTURE ALONG A STEEP ENVIRONMENTAL GRADIENT: CONSEQUENCES OF FLOWERING TIME AND HABITAT VARIATION IN THE SNOW BUTTERCUP,RANUNCULUS ADONEUS

Few studies have determined how gene flow and selection interact to generate population genetic structure in heterogeneous environments. One way to identify the potential role played by natural selection is to compare patterns of spatial genetic structure between different life cycle stages and among microenvironments. We examined patterns of spatial structure in a population of the snow buttercup (Ranunculus adoneus), using both adult plants and newly emerged seedlings. The study population spans a steep environmental gradient caused by gradual melting of snow within a permanent snowbed. Early-melting sites are characterized by denser vegetation, more fertile soils, and a longer growing season than late-melting sites tens of meters away. The flowering time of R. adoneus is controlled entirely by time of snowmelt, so the contiguous population is phenologically substructured into a series of successively flowering cohorts, reducing the opportunity for direct pollen transfer between early- and late-melting sites. For four highly polymorphic enzyme loci in this tetraploid species, there was subtle, but statistically significant, genetic differentiation between early, middle, and late-melting cohorts; adults usually showed greater differentiation among snowmelt zones than did seedlings. At two loci in adults and one locus in seedlings, homozygotes were more common than predicted at Hardy-Weinberg equilibrium, even when assuming maximum levels of double reduction during meiosis. This pattern suggests the occurrence of self-fertilization and/or population substructure. To determine how spatial isolation and phenological separation each contribute to genetic substructure, we used bivariate regression models to predict the numbers of allele differences between randomly paired individuals as a function of meters separation in space and days separation in flowering time. For newly emerged seedlings, we found that spatial separation was positively associated with genetic difference, but that the additional contribution of phenological separation to genetic difference was not significant. This implies that seeds and/or pollen move effectively across the snowmelt gradient, despite differences in flowering time. As was true for seedlings, spatial separation between paired adults contributed to greater genetic difference, but for a given spatial separation, the genetic difference between adult plants was reduced by phenological separation. This result implies that postemergence selection is favoring at least some seeds that migrate across the snowmelt gradient. Directional gene flow across the snowmelt gradient probably results from a genetic source-sink interaction, that is, the colonization of ecologically marginal late-melting sites by high quality seeds produced by the larger subpopulation in early-melting sites. Effective gene flow from high to low quality microenvironments is likely to impede adaptation to late-melting locations.     

Plant reproductive systems and evolution during biological invasion

Recent biological invasions provide opportunities to investigate microevolution during contemporary timescales. The tempo and scope of local adaptation will be determined by the intensity of natural selection and the amounts and kinds of genetic variation within populations. In flowering plants, genetic diversity is strongly affected by interactions between reproductive systems and stochastic forces associated with immigration history and range expansion. Here, we explore the significance of reproductive system diversity for contemporary evolution during plant invasion. We focus in particular on how reproductive modes influence the genetic consequences of long-distance colonization and determine the likelihood of adaptive responses during invasion. In many clonal invaders, strong founder effects and restrictions on sexual reproduction limit opportunities for local adaptation. In contrast, adaptive changes to life-history traits should be a general expectation in both outbreeding and inbreeding species. We provide evidence that evolutionary modifications to reproductive systems promote the colonizing ability of invading populations and that reproductive timing is an important target of selection during range expansion. Knowledge of the likelihood and speed at which local adaptation evolves in invasive plants will be particularly important for management practices when evolutionary changes enhance ecological opportunities and invasive spread.     

Natural selection drives clinal life history patterns in the perennial sunflower species, Helianthus maximiliani

In plants, ecologically important life history traits often display clinal patterns of population divergence. Such patterns can provide strong evidence for spatially varying selection across environmental gradients but also may result from nonselective processes, such as genetic drift, population bottlenecks and spatially restricted gene flow. Comparison of population differentiation in quantitative traits (measured as Q(ST) ) with neutral molecular markers (measured as F(ST) ) provides a useful tool for understanding the relative importance of adaptive and nonadaptive processes in the formation and maintenance of clinal variation. Here, we demonstrate the existence of geographic variation in key life history traits in the diploid perennial sunflower species Helianthus maximiliani across a broad latitudinal transect in North America. Strong population differentiation was found for days to flowering, growth rate and multiple size-related traits. Differentiation in these traits greatly exceeds neutral predictions, as determined both by partial Mantel tests and by comparisons of global Q(ST) values with theoretical F(ST) distributions. These findings indicate that clinal variation in these life history traits likely results from local adaptation driven by spatially heterogeneous environments.     

Evidence for Fine-Scale Natal Homing Among Island Beach Spawning Sockeye Salmon, Oncorhynchus nerka

Salmonid fishes aggregate for breeding at spatially defined, suitable habitats. These aggregations may evolve into discrete populations when precise natal homing leads to reproductive isolation, and local regimes of selection lead to adaptation. Population structure is often defined by persistent differences in selectively neutral genetic markers and in mean values of morphological and life-history traits between locations. This approach is limited by the spatial scale at which traits diverge; low levels of reproductively successful straying, combined with similar selective pressures on life-history traits resulting from similar habitat features and environmental conditions, can significantly reduce the power of these discriminatory methods. We compared data on three life-history traits and polymorphism of DNA microsatellites for evidence of population subdivision among sockeye salmon spawning on spatially discrete but physically similar beaches on islands in Iliamna Lake, Alaska. We found small but significant differences in average body length, body depth and age composition between sites as well as significant interactions between site and year. These interactions, reflecting random variation in growth or recruitment among sites, are a powerful tool for discriminating populations with similar mean trait values. These results suggest fine-scale homing to natal sites, but the microsatellite data revealed no evidence of restricted gene flow among sites. There seems to be enough straying among the populations to prevent differentiation at neutral traits but enough homing for them to be functionally distinct.     

On the evolution of clonal plant life histories

Clonal plant life histories are special in at least four respects: (1) Clonal plants can also reproduce vegetatively, (2) vegetative reproduction can be realised with short or long spacers, (3) and it may allow to plastically place vegetative offspring in benign patches. (4) Moreover, ramets of clonal plants may remain physically and physiologically integrated. Because of the apparent utility of such traits and because ecological patterns of distribution of clonal and non-clonal plants differ, adaptation is a tempting explanation of observed clonal life-history variation. However, adaptive evolution requires (1) heritable genetic variation and (2) a trait effect on fitness, and (3) it may be constrained if other evolutionary forces are overriding selection or by constraints, costs and trade-offs. (1) The few studies undertaken so far reported broad-sense heritability for clonal traits. Variation in selectively neutral genetic markers appears as pronounced in populations of clonal as non-clonal plants. However, neutral markers may not reflect heritable variation of life-history traits. Moreover, clonal plants may have been sampled at larger spatial scales. Empirical information on the contribution of somatic mutations to heritable variation is lacking. (2) Clonal life-history traits were found to affect fitness. However, much of this evidence stems from artificial rather than natural environments. (3) The relative importance of gene flow, inbreeding, and genetic drift, compared with selection, in the evolution of clonal life histories is hardly explored. Benefits of clonal life-history traits were frequently studied and found. However, there is also evidence for constraints, trade-offs, and costs. In conclusion, though it is very likely, that clonal life-history traits are adaptive, it is neither clear to which degree this is the case, nor which clonal life-history traits constitute adaptations to which environmental factors. Moreover, evolutionary interactions among clonal life-history traits and between clonal and non-clonal ones, such as the mating system, are not well explored. There remains much interesting work to be done in this field – which will be particularly interesting if it is done in the field.     

Glacial History Affected Phenotypic Differentiation in the Alpine Plant,Campanula thyrsoides

Numerous widespread Alpine plant species show molecular differentiation among populations from distinct regions. This has been explained as the result of genetic drift during glacial survival in isolated refugia along the border of the European Alps. Since genetic drift may affect molecular markers and phenotypic traits alike, we asked whether phenotypic differentiation mirrors molecular patterns among Alpine plant populations from different regions. Phenotypic traits can be under selection, so we additionally investigated whether part of the phenotypic differentiation can be explained by past selection and/or current adaptation. Using the monocarpic Campanula thyrsoides as our study species, a common garden experiment with plants from 21 populations from four phylogeographic groups located in regions across the Alps and the Jura Mountains was performed to test for differentiation in morphological and phenological traits. Past selection was investigated by comparing phenotypic differentiation among and within regions with molecular differentiation among and within regions. The common garden results indicated regional differentiation among populations for all investigated phenotypic traits, particularly in phenology. Delayed flowering in plants from the South-eastern Alps suggested adaptation to long sub-mediterranean summers and contrasted with earlier flowering of plants experiencing shorter growing seasons in regions with higher elevation to the West. Comparisons between molecular and phenotypic differentiation revealed diversifying selection among regions in height and biomass, which is consistent with adaptation to environmental conditions in glacial refugia. Within regions, past selection acted against strong diversification for most phenotypic traits, causing restricted postglacial adaptation. Evidence consistent with post-glacial adaptation was also given by negative correlation coefficients between several phenotypic traits and elevation of the population''s origin. In conclusion, our study suggests that, irrespective of adaptation of plants to their current environment, glacial history can have a strong and long-lasting influence on the phenotypic evolution of Alpine plants.     

Do surface plant and soil seed bank populations differ genetically? a multipopulation study of the desert mustard Lesquerella fendleri (Brassicaceae)

Seed banks are an important component of many plant populations, but few empirical studies have investigated the genetic relationship between soil seeds and surface plants. We compared the genetic structure of soil seeds and surface plants of the desert mustard Lesquerella fendleri within and among five ecologically diverse populations at the Sevilleta National Wildlife Refuge in Central New Mexico. At each site, 40 Lesquerella surface plants and 40 samples of soil seeds were mapped and genetically analyzed using starch gel electrophoresis. Overall allele frequencies of soil seeds and surface plants showed significant differences across the five populations and within three of the five individual populations. Surface plants had significantly greater amounts of single and multilocus heterozygosity, and mean surface plant heterozygosity was also greater at the total population level and in four of the five individual populations. Overall soil seed (bot not surface plant) homozygosity was significantly greater than predicted by Hardy-Weinberg expectations at the total and individual population levels. Although F-alpha estimates revealed similarly small but significant genetic divergence within each life-history stage, estimates of coancestry showed that fine-scale (0.5-2 m) genetic correlations among the surface plant genotypes were roughly twice those of soil seed genotypes. An unweighted pair group method with arithrnetic mean cluster analysis indicated that in the two geographically closest sites, the surface plants were slightly more genetically similar to each other than to their own respective seed banks. We also found weak and/or negative demographic associations between Lesquerella soil seed and surface plant densities within each of the five sites. We discuss the difficulties involved with sampling and genetically comparing these two life-history stages.     

Climate change shifts natural selection and the adaptive potential of the perennial forb Boechera stricta in the Rocky Mountains

Heritable genetic variation is necessary for populations to evolve in response to anthropogenic climate change. However, antagonistic genetic correlations among traits may constrain the rate of adaptation, even if substantial genetic variation exists. We examine potential genetic responses to selection by comparing multivariate genetic variance–covariances of traits and fitness (multivariate Robertson–Price identities) across different environments in a reciprocal transplant experiment of the forb Boechera stricta in the Rocky Mountains. By transplanting populations into four common gardens arrayed along an elevational gradient, and exposing populations to control and snow removal treatments, we simulated future and current climates and snowmelt regimes. Genetic variation in flowering and germination phenology declined in plants moved downslope to warmer, drier sites, suggesting that these traits may have a limited ability to evolve under future climates. Simulated climate change via snow removal altered the strength of selection on flowering traits, but we found little evidence that genetic correlations among traits are likely to affect the rate of adaptation to climate change. Overall, our results suggest that climate change may alter the evolutionary potential of B. stricta, but reduced expression of genetic variation may be a larger impediment to adaptation than constraints imposed by antagonistic genetic correlations.     

Climate affects predator control of an herbivore outbreak

Herbivore outbreaks and the accompanying devastation of plant biomass can have enormous ecological effects. Climate directly affects such outbreaks through plant stress or alterations in herbivore life-history traits. Large-scale variation in climate can indirectly affect outbreaks through trophic interactions, but the magnitude of such effects is unknown. On the California coast, rainfall in years during and immediately previous to mass lupine mortality was two-thirds that of years without such mortality. However, neither mature lupines nor their root-feeding herbivores are directly affected by annual variation in rainfall. By increasing soil moisture to levels characteristic of summers following El Ni?o/Southern Oscillation (ENSO) events, we increased persistence of a predator (the entomopathogenic nematode Heterorhabditis marelatus). This led to suppression of an outbreak of the herbivorous moth Hepialus californicus, indirectly protecting bush lupine (Lupinus arboreus). Our results are consistent with the marine-oriented Menge-Sutherland hypothesis (Menge and Sutherland 1987) that abiotic stress has greater effects on higher than on lower trophic levels. The mechanisms producing these results differ from those proposed by Menge-Sutherland, however, highlighting differences between trophic processes in underground and terrestrial/marine food webs. Our evidence suggests that herbivore outbreaks and mass lupine mortality are indirectly affected by ENSO's facilitation of top-down control in this food web.     

Effects of founding genetic variation on adaptation to a novel resource

Population genetic theory predicts that adaptation in novel environments is enhanced by genetic variation for fitness. However, theory also predicts that under strong selection, demographic stochasticity can drive populations to extinction before they can adapt. We exposed wheat-adapted populations of the flour beetle (Tribolium castaneum) to a novel suboptimal corn resource, to test the effects of founding genetic variation on population decline and subsequent extinction or adaptation. As previously reported, genetically diverse populations were less likely to go extinct. Here, we show that among surviving populations, genetically diverse groups recovered faster after the initial population decline. Within two years, surviving populations significantly increased their fitness on corn via increased fecundity, increased egg survival, faster larval development, and higher rate of egg cannibalism. However, founding genetic variation only enhanced the increase in fecundity, despite existing genetic variation-and apparent lack of trade-offs-for egg survival and larval development time. Thus, during adaptation to novel habitats the positive impact of genetic variation may be restricted to only a few traits, although change in many life-history traits may be necessary to avoid extinction. Despite severe initial maladaptation and low population size, genetic diversity can thus overcome the predicted high extinction risk in new habitats.     

Migration-induced phenotypic divergence: the migration-selection balance of correlated traits

Genetically correlated traits are known to respond to indirect selection pressures caused by directional selection on other traits. It is however unclear how local adaptation in populations diverging along some phenotypic traits but not others is affected by the joint action of gene flow and genetic correlations among traits. This simulation study shows that although gene flow is a potent constraining mechanism of population adaptive divergence, it may induce phenotypic divergence in traits under homogeneous selection among habitats if they are genetically correlated with traits under divergent selection. This correlated phenotypic divergence is a nonmonotonous function of migration and increases with mutational correlation among traits. It also increases with the number of divergently selected traits provided their genetic autonomy relative to the uniformly selected trait is reduced by specific patterns of genetic covariances: populations with lower effective trait dimensionality are more likely to generate very large correlated divergence. The correlated divergence is likely to be picked up by Q(ST)-F(ST) analysis of population genetic differentiation and be erroneously ascribed to adaptive divergence under divergent selection. This study emphasizes the necessity to understand the interaction between selection and the genetic basis of adaptation in a multivariate rather than univariate context.     

Differential threshold effects of habitat fragmentation on gene flow in two widespread species of bush crickets

Effects of habitat fragmentation on genetic diversity vary among species. This may be attributed to the interacting effects of species traits and landscape structure. While widely distributed and abundant species are often considered less susceptible to fragmentation, this may be different if they are small sized and show limited dispersal. Under intensive land use, habitat fragmentation may reach thresholds at which gene flow among populations of small-sized and dispersal-limited species becomes disrupted. Here, we studied the genetic diversity of two abundant and widespread bush crickets along a gradient of habitat fragmentation in an agricultural landscape. We applied traditional (G(ST), θ) and recently developed (G'ST', D) estimators of genetic differentiation on microsatellite data from each of twelve populations of the grassland species Metrioptera roeselii and the forest-edge species Pholidoptera griseoaptera to identify thresholds of habitat fragmentation below which genetic population structure is affected. Whereas the grassland species exhibited a uniform genetic structuring (G(ST) = 0.020-0.033; D = 0.085-0.149) along the whole fragmentation gradient, the forest-edge species' genetic differentiation increased significantly from D < 0.063 (G(ST) < 0.018) to D = 0.166 (G(ST) = 0.074), once the amount of suitable habitat dropped below a threshold of 20% and its proximity decreased substantially at the landscape scale. The influence of fragmentation on genetic differentiation was qualitatively unaffected by the choice of estimators of genetic differentiation but quantitatively underestimated by the traditional estimators. These results indicate that even for widespread species in modern agricultural landscapes fragmentation thresholds exist at which gene flow among suitable habitat patches becomes restricted.     

PLEIOTROPY CAUSES LONG-TERM GENETIC CONSTRAINTS ON LIFE-HISTORY EVOLUTION IN BRASSICA RAPA

Fundamental, long-term genetic trade-offs constrain life-history evolution in wild crucifer populations. I studied patterns of genetic constraint in Brassica rapa by estimating genetic correlations among life-history components by quantitative genetic analyses among ten wild populations, and within four populations. Genetic correlations between age and size at first reproduction were always greater than +0.8 within and among all populations studied. Although quantitative genetics might provide insight about genetic constraints if genetic parameters remain approximately constant, little evidence has been available to determine the constancy of genetic correlations. I found strong and consistent estimates of genetic correlations between life-history components, which were very similar within four natural populations. Population differentiation also showed these same trade-offs, resulting from long-term genetic constraint. For some traits, evolutionary changes among populations were incompatible with a model of genetic drift. Historical patterns of natural selection were inferred from population differentiation, suggesting that correlated response to selection has caused some traits to evolve opposite to the direct forces of natural selection. Comparison with Arabidopsis suggests that these life-history trade-offs are caused by genes that regulate patterns of resource allocation to different components of fitness. Ecological and energetic models may correctly predict these trade-offs because there is little additive genetic variation for rates of resource acquisition, but resource allocation is genetically variable.