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Abstract A capacity for rapid adaptation should enhance the persistence of populations subject to temporal and spatial heterogeneity in natural selection, but examples from nature remain scarce. Plectritis congesta (Caprifoliaceae) is a winter annual that exhibits local adaptation to browsing by ungulates and hypothesized to show context‐dependent trade‐offs in traits affecting success in competition versus resistance or tolerance to browsing. We grew P. congesta from 44 insular populations historically exposed or naïve to ungulates in common gardens to (1) quantify genetic, plastic and competitive effects on phenotype; (2) estimate a capacity for rapid adaptation (evolvability); and (3) test whether traits favoured by selection with ungulates present were selected against in their absence. Plants from browsed populations bolted and flowered later, had smaller inflorescences, were less fecund and half as tall as plants from naïve populations on average, replicating patterns in nature. Estimated evolvabilities (3–36%) and narrow‐sense heritabilities (h2; 0.13–0.32) imply that differences in trait values as large as reported here can arise in 2–18 generations in an average population. Phenotypic plasticity was substantial, varied by browsing history and fruit phenotype and increased with competition. Fecundity increased with plasticity in flowering height given competition (0.47 ± 0.02 florets/cm, β ± se), but 23–77% faster in naïve plants bearing winged fruits (0.53 ± 0.04) than exposed‐wingless plants (0.43 ± 0.03) or exposed‐winged and naïve‐wingless plants (0.30 ± 0.03, each case). Our results support the hypothesis that context‐dependent variation in natural selection in P. congesta populations has conferred a substantial capacity for adaptation in response to selection in traits affecting success in competition versus resistance or tolerance to browsing in the absence versus presence of ungulates, respectively. Theory suggests that conserving adaptive capacity in P. congesta will require land managers to maintain spatial heterogeneity in natural selection, prevent local extinctions and maintain gene flow.

The amount of standing variation present within populations is a fundamental quantity of interest in population genetics, commonly represented by calculating the average number of differences between pairs of nucleotide sequences (nucleotide diversity, π). It is well understood that both background and positive selection can cause reductions in nucleotide diversity, but less clear how local adaptation affects it. Depending on the assumptions and parameters, some theoretical studies have emphasized how local adaptation can reduce nucleotide diversity, while others have shown that it can increase it. Here, we explore how local adaptation shapes genome-wide patterns in within-population nucleotide diversity, extending previous work to study the effects of polygenic adaptation, genotypic redundancy, and population structure. We show that local adaptation produces two very different patterns depending on the relative strengths of migration and selection, either markedly decreasing or increasing within-population diversity at linked sites at equilibrium. At low migration, regions of depleted diversity can extend large distances from the causal locus, with substantially more diversity eroded than expected with background selection. With higher migration, peaks occur over much smaller genomic distances but with much larger magnitude changes in diversity. Across spatially extended environmental gradients, both patterns can be found within a single species, with increases in diversity at the center of the range and decreases towards the periphery. Our results demonstrate that there is no universal diagnostic signature of local adaptation based on within-population nucleotide diversity, so it will not be broadly useful for explaining increased FST. However, given that neither background nor positive selection inflate diversity, when peaks are found they suggest local adaptation may be acting on a causal allele in the region., Competing Interests: Conflicts of interest The authors declare no conflict of interest., (© The Author(s) 2024. Published by Oxford University Press on behalf of The Genetics Society of America.)

A capacity for rapid adaptation should enhance the persistence of populations subject to temporal and spatial heterogeneity in natural selection, but examples from nature remain scarce. Plectritis congesta (Caprifoliaceae) is a winter annual that exhibits local adaptation to browsing by ungulates and hypothesized to show context‐dependent trade‐offs in traits affecting success in competition versus resistance or tolerance to browsing. We grew P. congesta from 44 insular populations historically exposed or naïve to ungulates in common gardens to (1) quantify genetic, plastic and competitive effects on phenotype; (2) estimate a capacity for rapid adaptation (evolvability); and (3) test whether traits favoured by selection with ungulates present were selected against in their absence. Plants from browsed populations bolted and flowered later, had smaller inflorescences, were less fecund and half as tall as plants from naïve populations on average, replicating patterns in nature. Estimated evolvabilities (3–36%) and narrow‐sense heritabilities (h2; 0.13–0.32) imply that differences in trait values as large as reported here can arise in 2–18 generations in an average population. Phenotypic plasticity was substantial, varied by browsing history and fruit phenotype and increased with competition. Fecundity increased with plasticity in flowering height given competition (0.47 ± 0.02 florets/cm, β ± se), but 23–77% faster in naïve plants bearing winged fruits (0.53 ± 0.04) than exposed‐wingless plants (0.43 ± 0.03) or exposed‐winged and naïve‐wingless plants (0.30 ± 0.03, each case). Our results support the hypothesis that context‐dependent variation in natural selection in P. congesta populations has conferred a substantial capacity for adaptation in response to selection in traits affecting success in competition versus resistance or tolerance to browsing in the absence versus presence of ungulates, respectively. Theory suggests that conserving adaptive capacity in P. congesta will require land managers to maintain spatial heterogeneity in natural selection, prevent local extinctions and maintain gene flow. [ABSTRACT FROM AUTHOR]

Adaptation to derived habitats often occurs from standing genetic variation. The maintenance within ancestral populations of genetic variants favourable in derived habitats is commonly ascribed to long‐term antagonism between purifying selection and gene flow resulting from hybridization across habitats. A largely unexplored alternative idea based on quantitative genetic models of polygenic adaptation is that variants favoured in derived habitats are neutral in ancestral populations when their frequency is relatively low. To explore the latter, we first identify genetic variants important to the adaptation of threespine stickleback fish (Gasterosteus aculeatus) to a rare derived habitat—nutrient‐depleted acidic lakes—based on whole‐genome sequence data. Sequencing marine stickleback from six locations across the Atlantic Ocean then allows us to infer that the frequency of these derived variants in the ancestral habitat is unrelated to the likely opportunity for gene flow of these variants from acidic‐adapted populations. This result is consistent with the selective neutrality of derived variants within the ancestor. Our study thus supports an underappreciated explanation for the maintenance of standing genetic variation, and calls for a better understanding of the fitness consequences of adaptive variation across habitats and genomic backgrounds. [ABSTRACT FROM AUTHOR]

Populations often inhabit multiple ecological patches and thus experience divergent selection, which can lead to local adaptation if migration is not strong enough to swamp locally adapted alleles. Conditions for the establishment of a locally advantageous allele have been studied in randomly mating populations. However, many species reproduce, at least partially, through self-fertilization, and how selfing affects local adaptation remains unclear and debated. Using a two-patch branching process formalism, we obtained a closed-form approximation under weak selection for the probability of establishment of a locally advantageous allele (P) for arbitrary selfing rate and dominance level, where selection is allowed to act on viability or fecundity, and migration can occur via seed or pollen dispersal. This solution is compared to diffusion approximation and used to investigate the consequences of a shift in a mating system on P, and the establishment of protected polymorphism. We find that selfing can either increase or decrease P, depending on the patterns of dominance in the two patches, and has conflicting effects on local adaptation. Globally, selfing favors local adaptation when locally advantageous alleles are (partially) recessive, when selection between patches is asymmetrical and when migration occurs through pollen rather than seed dispersal. These results establish a rigorous theoretical background to study heterogeneous selection and local adaptation in partially selfing species. [ABSTRACT FROM AUTHOR]

Recent work incorporating demographic-genetic interactions indicates the importance of population size, gene flow, and selection in influencing local adaptation. This work typically assumes that density-dependent survival affects individuals equally, but individuals in natural population rarely compete equally. Among-individual differences in resource use generate stronger competition between more similar phenotypes (frequency-dependent competition) but it remains unclear how this additional form of selection changes the interactions between population size, gene flow, and local stabilizing selection. Here, we integrate migration-selection dynamics with frequency-dependent competition. We developed a coupled demographic-quantitative genetic model consisting of two patches connected by dispersal and subject to local stabilizing selection and competition. Our model shows that frequency-dependent competition slightly increases local adaptation, greatly increases genetic variance within patches, and reduces the amount that migration depresses population size, despite the increased genetic variance load. The effects of frequency-dependence depend on the strength of divergent selection, trait heritability, and when mortality occurs in the life cycle in relation to migration and reproduction. Essentially, frequency-dependent competition reduces the density-dependent interactions between migrants and residents, the extent to which depends on how different and common immigrants are compared to residents. Our results add new dynamics that illustrate how competition can alter the effects of gene flow and divergent selection on local adaptation and population carrying capacities.

Ongoing adaptive evolution, and resulting "evolutionary rescue" of declining populations, requires additive genetic variation in fitness. Such variation can be increased by gene flow resulting from immigration, potentially facilitating evolution. But, gene flow could in fact constrain rather than facilitate local adaptive evolution if immigrants have low additive genetic values for local fitness. Local migration‐selection balance and micro‐evolutionary stasis could then result. However, key quantitative genetic effects of natural immigration, comprising the degrees to which gene flow increases the total local additive genetic variance yet counteracts local adaptive evolutionary change, have not been explicitly quantified in wild populations. Key implications of gene flow for population and evolutionary dynamics consequently remain unclear. Our quantitative genetic analyses of long‐term data from free‐living song sparrows (Melospiza melodia) show that mean breeding value for local juvenile survival to adulthood, a major component of fitness, increased across cohorts more than expected solely due to drift. Such micro‐evolutionary change should be expected given nonzero additive genetic variance and consistent directional selection. However, this evolutionary increase was counteracted by negative additive genetic effects of recent immigrants, which increased total additive genetic variance but prevented a net directional evolutionary increase in total additive genetic value. These analyses imply an approximate quantitative genetic migration‐selection balance in a major fitness component, and hence demonstrate a key mechanism by which substantial additive genetic variation can be maintained yet decoupled from local adaptive evolutionary change. [ABSTRACT FROM AUTHOR]

Since the New Synthesis, most migration‐selection balance theory has predicted that there should be negligible differentiation over small spatial scales (relative to dispersal), because gene flow should erode any effect of divergent selection. Nevertheless, there are classic examples of microgeographic divergence, which theory suggests can arise under specific conditions: exceptionally strong selection, phenotypic plasticity in philopatric individuals, or nonrandom dispersal. Here, we present evidence of microgeographic morphological variation within lake and stream populations of threespine stickleback (Gasterosteus aculeatus). It seems reasonable to assume that a given lake or stream population of fish is well‐mixed. However, we found this assumption to be untenable. We examined trap‐to‐trap variation in 34 morphological traits measured on stickleback from 16 lakes and 16 streams. Most traits varied appreciably among traps within populations. Both between‐trap distance and microhabitat characteristics such as depth and substrate explained some of the within‐population morphological variance. Microhabitat was also associated with genotype at particular loci but there was no genetic isolation by distance, implying that heritable habitat preferences may contribute to microgeographic variation. Our study adds to growing evidence that microgeographic divergence can occur across small spatial scales within individuals' daily dispersal neighborhood where gene flow is expected to be strong. [ABSTRACT FROM AUTHOR]

Adaptation in spatially heterogeneous environments results from the balance between local selection, mutation, and migration. We study the interplay among these different evolutionary forces and demography in a classical two-habitat scenario with asexual reproduction. We develop a new theoretical approach that goes beyond the Adaptive Dynamics framework, and allows us to explore the effect of high mutation rates on the stationary phenotypic distribution. We show that this approach improves the classical Gaussian approximation, and captures accurately the shape of this equilibrium phenotypic distribution in one- and two-population scenarios. We examine the evolutionary equilibrium under general conditions where demography and selection may be nonsymmetric between the two habitats. In particular, we show how migration may increase differentiation in a source-sink scenario. We discuss the implications of these analytic results for the adaptation of organisms with large mutation rates, such as RNA viruses. [ABSTRACT FROM AUTHOR]

Summary: Self‐fertilisation has consequences for variation across the genome as it reduces effective population size, effect recombination rates and pollen flow, with implications for local adaptation.We conducted simulations of divergent stabilising selection on a quantitative trait with drift, pollen flow, mutation, recombination and different outcrossing rates. We quantified trait divergence and the genetic architecture of adaptation. We conducted an FST outlier analysis to identify candidate loci and quantified the impact of mating system on detectability.Selfing promoted trait divergence mainly through reductions in pollen flow. Moreover, trait architecture became more diffuse with selfing. Average effect size of trait loci was lower, while the number of loci, and their clustering distance increased. The genetic architecture of selfers was also more diffuse than outcrossers for equivalent migration rates. However, when deleterious alleles were included, architectures became more concentrated in selfers, likely to be because of reductions in population size caused by mutational meltdown and impacts of background selection on Ne.Our simulations demonstrate that mating system has important impacts on adaptive divergence of traits and the genetic landscape underlying that divergence. Selfing has a significant effect on detectability of regions of the genome important for adaptation because of neutral divergence and diffuse trait architecture. [ABSTRACT FROM AUTHOR]

Evolutionary biologists typically represent clines as spatial gradients in a univariate character (or a principal-component axis) whose mean changes as a function of location along a transect spanning an environmental gradient or ecotone. This univariate approach may obscure the multivariate nature of phenotypic evolution across a landscape. Clines might instead be plotted as a series of vectors in multidimensional morphospace, connecting sequential geographic sites. We present amodel showing that clinesmay trace nonlinear paths that arc through morphospace rather than elongating along a single major trajectory. Arcing clines arise because different characters diverge at different rates or locations along a geographic transect. We empirically confirmthat some clines arc throughmorphospace, usingmorphological data fromthreespine stickleback sampled along eight independent transects from lakes down their respective outlet streams. In all eight clines, successive vectors of lake-stream divergence fluctuate in direction and magnitude in trait space, rather than pointing along a single phenotypic axis. Most clines exhibit surprisingly irregular directions of divergence as one moves downstream, although a few clines exhibit more directional arcs throughmorphospace. Our results highlight the multivariate complexity of clines that cannot be captured with the traditional graphical framework. We discuss hypotheses regarding the causes, and implications, of such arcing multivariate clines. [ABSTRACT FROM AUTHOR]

Background: Blindness has evolved repeatedly in cave-dwelling organisms, and many hypotheses have been proposed to explain this observation, including both accumulation of neutral loss-of-function mutations and adaptation to darkness. Investigating the loss of sight in cave dwellers presents an opportunity to understand the operation of fundamental evolutionary processes, including drift, selection, mutation, and migration. Results: Here we model the evolution of blindness in caves. This model captures the interaction of three forces: (1) selection favoring alleles causing blindness, (2) immigration of sightedness alleles from a surface population, and (3) mutations creating blindness alleles. We investigated the dynamics of this model and determined selection-strength thresholds that result in blindness evolving in caves despite immigration of sightedness alleles from the surface. We estimate that the selection coefficient for blindness would need to be at least 0.005 (and maybe as high as 0.5) for blindness to evolve in the model cave-organism, Astyanax mexicanus. Conclusions: Our results indicate that strong selection is required for the evolution of blindness in cave-dwelling organisms, which is consistent with recent work suggesting a high metabolic cost of eye development. [ABSTRACT FROM AUTHOR]

• Identifies strength of selection on locally-adaptive alleles. • Shows dependence of selection coefficients on allele frequencies. • Backed up by simulation of finite-population models. • Quantifies the advantages conferred by adaptive genes. Understanding the role of natural selection in driving evolutionary change requires accurate estimates of the strength of selection acting at the genetic level in the wild. This is challenging to achieve but may be easier in the case of populations in migration-selection balance. When two populations are at equilibrium under migration-selection balance, there exist loci whose alleles are selected different ways in the two populations. Such loci can be identified from genome sequencing by their high values of F ST. This raises the question of what is the strength of selection on locally-adaptive alleles. To answer this question we analyse a 1-locus 2-allele model of a population distributed between two niches. We show by simulation of selected cases that the outputs from finite-population models are essentially the same as those from deterministic infinite-population models. We then derive theory for the infinite-population model showing the dependence of selection coefficients on equilibrium allele frequencies, migration rates, dominance and relative population sizes in the two niches. An Excel spreadsheet is provided for the calculation of selection coefficients and their approximate standard errors from observed values of population parameters. We illustrate our results with a worked example, with graphs showing the dependence of selection coefficients on equilibrium allele frequencies, and graphs showing how F ST depends on the selection coefficients acting on the alleles at a locus. Given the extent of recent progress in ecological genomics, we hope our methods may help those studying migration-selection balance to quantify the advantages conferred by adaptive genes. [ABSTRACT FROM AUTHOR]

Abstract: Interbreeding between domesticated individuals and their wild conspecifics occurs in a range of species. The rate of gene flow into the wild population, estimated using genetic markers, is often smaller than the fraction of immigrants, as immigrants and their descendants generally have lower relative fitness. Here the difference between one-way migration rate and gene flow (effective migration rate) is explored using quantitative genetic simulations. We model a trait undergoing stabilizing selection in the recipient population, influenced by an infinite number of loci, each with small effect. Immigrants have suboptimal trait values, and differ in allele frequency at an unlinked neutral marker locus. We derive an analytical approximation for the effective migration rate, and show that in the limiting case of low migration rates, the ratio between effective and actual migration rate approximately equals the ratio between mean fitness of immigrants and in the admixed population. This ignores indirect selection on the marker locus within the admixed population due to covariance with the trait value, which may be substantial when the genetic difference between the populations is large. For higher migration rates, the gene flow/migration ratio rises with increasing migration rate, inflating the rate of gene flow. [Copyright &y& Elsevier]

How do mutation and gene flow influence population persistence, niche expansion and local adaptation in spatially heterogeneous environments? In this article, we analyse a demographic and evolutionary model of adaptation to an environment containing two habitats in equal frequencies, and we bridge the gap between different theoretical frameworks. Qualitatively, our model yields four qualitative types of outcomes: (i) global extinction of the population, (ii) adaptation to one habitat only, but also adaptation to both habitats with, (iii) specialized phenotypes or (iv) with generalized phenotypes, and we determine the conditions under which each equilibrium is reached. We derive new analytical approximations for the local densities and the distributions of traits in each habitat under a migration-selection-mutation balance, compute the equilibrium values of the means, variances and asymmetries of the local distributions of phenotypes, and contrast the effects of migration and mutation on the evolutionary outcome. We then check our analytical results by solving our model numerically, and also assess their robustness in the presence of demographic stochasticity. Although increased migration results in a decrease in local adaptation, mutation in our model does not influence the values of the local mean traits. Yet, both migration and mutation can have dramatic effects on population size and even lead to metapopulation extinction when selection is strong. Niche expansion, the ability for the population to adapt to both habitats, can also be prevented by small migration rates and a reduced evolutionary potential characterized by rare mutation events of small effects; however, niche expansion is otherwise the most likely outcome. Although our results are derived under the assumption of clonal reproduction, we finally show and discuss the links between our model and previous quantitative genetics models. [ABSTRACT FROM AUTHOR]

The balance between stabilizing selection and migration of maladapted individuals has formerly been modeled using a variety of quantitative genetic models of increasing complexity, including models based on a constant expressed genetic variance and models based on normality. The infinitesimal model can accommodate nonnormality and a nonconstant genetic variance as a result of linkage disequilibrium. It can be seen as a parsimonious one-parameter model that approximates the underlying genetic details well when a large number of loci are involved. Here, the performance of this model is compared to several more realistic explicit multilocus models, with either two, several or a large number of alleles per locus with unequal effect sizes. Predictions for the deviation of the population mean from the optimum are highly similar across the different models, so that the non-Gaussian infinitesimal model forms a good approximation. It does, however, generally estimate a higher genetic variance than the multilocus models, with the difference decreasing with an increasing number of loci. The difference between multilocus models depends more strongly on the effective number of loci, accounting for relative contributions of loci to the variance, than on the number of alleles per locus. [ABSTRACT FROM AUTHOR]

There is a long tradition in population genetics of exploring the maintenance of variation under migration-selection balance using deterministic models that assume infinite population size. With finite population size, stochastic dynamics can greatly reduce the potential for the maintenance of polymorphism, but this has yet to be explored in detail. Here, classical two-patch models are extended to predict: (1) the probability of a locally beneficial mutation rising in frequency in the patch where it is favored and (2) the critical threshold migration rate above which the maintenance of polymorphism is much less likely. Individual-based simulations show that these approximations provide accurate predictions across a wide range of parameter space. [ABSTRACT FROM AUTHOR]

Many ecologically important traits have a complex genetic basis, with the potential formutations at many different genes to shape the phenotype. Even so, studies of local adaptation in heterogeneous environments sometimes find that just a few quantitative trait loci (QTL) of large effect can explain a large percentage of observed differences between phenotypically divergent populations. As high levels of gene flow can swamp divergence at weakly selected alleles, migration-selection-drift balance may play an important role in shaping the genetic architecture of local adaptation. Here, we use analytical approximations and individual-based simulations to explore how genetic architecture evolves when two populations connected by migration experience stabilizing selection toward different optima. In contrast to the exponential distribution of allele effect sizes expected under adaptation without migration (Orr 1998), we find that adaptation with migration tends to result in concentrated genetic architectures with fewer, larger, and more tightly linked divergent alleles. Even if many small alleles contribute to adaptation at the outset, they tend to be replaced by a few large alleles under prolonged bouts of stabilizing selection with migration. All else being equal, we also find that stronger selection can maintain linked clusters of locally adapted alleles over much greater map distances than weaker selection. The common empirical finding of QTL of large effect is shown to be expected with migration in a heterogeneous landscape, and these QTL may often be composed of several tightly linked alleles of smaller effect. [ABSTRACT FROM AUTHOR]

Adaptive divergence between adjoining populations reflects a balance between the diversifying effect of divergent selection and the potentially homogenizing effect of gene flow. In most models of migration-selection balance, gene flow is assumed to reflect individuals' inherent capacity to disperse, without regard to the match between individuals' phenotypes and the available habitats. However, habitat preferences can reduce dispersal between contrasting habitats, thereby alleviating migration load and facilitating adaptive divergence. We tested whether habitat preferences contribute to adaptive divergence in a classic example of migration-selection balance: parapatric lake and stream populations of three-spine stickleback ( Gasterosteus aculeatus). Using a mark-transplant-recapture experiment on morphologically divergent parapatric populations, we showed that 90% of lake and stream stickleback returned to their native habitat, reducing migration between habitats by 76%. Furthermore, we found that dispersal into a nonnative habitat was phenotype dependent. Stream fish moving into the lake were morphologically more lake-like than those returning to the stream (and the converse for lake fish entering the stream). The strong native habitat preference documented here increases the extent of adaptive divergence between populations two- to fivefold relative to expectations with random movement. These results illustrate the potential importance of adaptive habitat choice in driving parapatric divergence. [ABSTRACT FROM AUTHOR]

Gene flow among populations is typically thought to be antagonistic to population differentiation and local adaptation. However, this assumes that dispersing individuals disperse randomly with respect to their ability to use the environment. Yet dispersing individuals often sample and compare environments and settle in those environments that best match their phenotype, causing directed gene flow, which can in fact promote population differentiation and adaptation. We refer to this process as “matching habitat choice.” Although this process has been acknowledged by several researchers, no synthesis or perspective on its potentially widespread importance exists. Here we synthesize empirical and theoretical studies, and offer a new perspective that matching habitat choice can have significant effects on important and controversial topics. We discuss the potential implications of matching habitat choice for the degree and rate of local adaptation, the evolution of niche width, adaptive peak shifts, speciation in the presence of gene flow, and on our view and interpretation of measures of natural selection. Because of its potential importance for such a wide range of topics, we call for heightened empirical and theoretical attention for this neglected dimension in evolutionary and ecological studies. [ABSTRACT FROM AUTHOR]

Dispersal moves individuals from patches where their immediate ancestors were successful to sites where their genotypes are untested. As a result, dispersal generally reduces fitness, a phenomenon known as "migration load." The strength of migration load depends on the pattern of dispersal and can be dramatically lessened or reversed when individuals move preferentially toward patches conferring higher fitness. Evolutionary ecologists have long modeled nonrandom dispersal, focusing primarily on its effects on population density over space, the maintenance of genetic variation, and reproductive isolation. Here, we build upon previous work by calculating how the extent of local adaptation and the migration load are affected when individuals differ in their dispersal rate in a genotype-dependent manner that alters their match to their environment. Examining a one-locus, two-patch model, we show that local adaptation occurs through a combination of natural selection and adaptive dispersal. For a substantial portion of parameter space, adaptive dispersal can be the predominant force generating local adaptation. Furthermore, genetic load may be largely averted with adaptive dispersal whenever individuals move before selective deaths occur. Thus, to understand the mechanisms driving local adaptation, biologists must account for the extent and nature of nonrandom, genotype-dependent dispersal, and the potential for adaptation via spatial sorting of genotypes.

Numerous studies of ecological genetics have found that alleles contributing to local adaptation sometimes cluster together, forming "genomic islands of divergence." Divergence hitchhiking theory posits that these clusters evolve by the preferential establishment of tightly linked locally adapted mutations, because such linkage reduces the rate that recombination breaks up locally favorable combinations of alleles. Here, I use calculations based on previously developed analytical models of divergence hitchhiking to show that very few clustered mutations should be expected in a single bout of adaptation, relative to the number of unlinked mutations, suggesting that divergence hitchhiking theory alone may often be insufficient to explain empirical observations. Using individual-based simulations that allow for the transposition of a single genetic locus from one position on a chromosome to another, I then show that tight clustering of the loci involved in local adaptation tends to evolve on biologically realistic time scales. These results suggest that genomic rearrangements may often be an important component of local adaptation and the evolution of genomic islands of divergence. More generally, these results suggest that genomic architecture and functional neighborhoods of genes may be actively shaped by natural selection in heterogeneous environments. Because small-scale changes in gene order are relatively common in some taxa, comparative genomic studies could be coupled with studies of adaptation to explore how commonly such rearrangements are involved in local adaptation.

Dobzhansky's model of epistatic selection assumes that viable genotypes form "clusters" in genotype space so that populations can evolve from one state to a reproductively isolated state following a "ridge" of well-fit genotypes without crossing any deep adaptive valleys. Recently, the importance of Dobzhansky-type models in evolutionary studies has been reemphasized by Gavrilets (1997a) and Gavrilets and Gravner (1997) who argue that the existence of "ridges" of well-fit genotypes connecting reproductively isolated genotypes is actually a general property of multidimensional adaptive landscapes. Using rigorous techniques and numerical simulations, I analyze clines in the frequencies of selected and neutral alleles maintained by a balance of migration and Dobzhansky-type epistatic selection acting on two diallelic loci. I show that Dobzhansky-type epistatic selection can build up a very strong barrier to neutral gene flow. I describe properties of clines that are indicative of Dobzhansky-type selection., (© 1997 The Society for the Study of Evolution.)