Geographic Mosaic Of Coevolution Research Articles

Conspicuous coloration of toxin-resistant predators implicates additional trophic interactions in a predator-prey arms race.

Antagonistic coevolution between natural enemies can produce highly exaggerated traits, such as prey toxins and predator resistance. This reciprocal process of adaptation and counter-adaptation may also open doors to other evolutionary novelties not directly involved in the phenotypic interface of coevolution. We tested the hypothesis that predator-prey coevolution coincided with the evolution of conspicuous coloration on resistant predators that retain prey toxins. In western North America, common garter snakes (Thamnophis sirtalis) have evolved extreme resistance to tetrodotoxin (TTX) in the coevolutionary arms race with their deadly prey, Pacific newts (Taricha spp.). TTX-resistant snakes can retain large amounts of ingested TTX, which could serve as a deterrent against the snakes' own predators if TTX toxicity and resistance are coupled with a conspicuous warning signal. We evaluated whether arms race escalation covaries with bright red coloration in snake populations across the geographic mosaic of coevolution. Snake colour variation departs from the neutral expectations of population genetic structure and covaries with escalating clines of newt TTX and snake resistance at two coevolutionary hotspots. In the Pacific Northwest, bright red coloration fits an expected pattern of an aposematic warning to avian predators: TTX-resistant snakes that consume highly toxic newts also have relatively large, reddish-orange dorsal blotches. Snake coloration also seems to have evolved with the arms race in California, but overall patterns are less intuitively consistent with aposematism. These results suggest that interactions with additional trophic levels can generate novel traits as a cascading consequence of arms race coevolution across the geographic mosaic.

A geographic mosaic of coevolution between Eurosta solidaginis (Fitch) and its host plant tall goldenrod Solidago altissima (L.).

A geographic mosaic of coevolution has produced local reciprocal adaptation in tall goldenrod, Solidago altissima (L.), and the goldenrod ball-gall fly, Eurosta solidaginis (Fitch 1855). The fly is selected to induce gall diameters that minimize mortality from natural enemies, and the plant is selected to limit gall growth that reduces plant fitness. We conducted a double reciprocal transplant experiment where S. altissima and E. solidaginis from three sites were grown in gardens at each site to partition the gall morphology variation into fly genotype, plant genotype, and the environment components. The host plant gall diameter induced by each E. solidaginis population was adapted to inhibit local natural enemies from ovipositing on or consuming enclosed larvae. Reciprocally, increasing the gall size induced by the local fly population increased the resistance of the local plant host population to gall growth. Differences among sites in natural enemies produced a mosaic of hotspots of coevolutionary arms races between flies selecting for greater gall diameter and plants for smaller diameters, and coldspots where there is no selection on plant or fly for a change in gall diameter. In contrast, the geographic variations of gall length and gall shape were not due to coevolutionary interactions.

Strong Gene Flow Undermines Local Adaptations in a Host Parasite System.

Simple SummaryThe co-evolution of hosts and parasites depends on their ability to adapt to each other’s defense and counter-defense mechanisms. The strength of selection on those mechanisms may vary among populations, resulting in a geographical mosaic of co-evolution. The boreo-montane paper wasp Polistes biglumis and its parasite Polistes atrimandibularis exemplify this type of co-evolutionary system. Here, we used genetic markers to examine the genetic population structures of these wasps in the western Alps. We found that both host and parasite populations displayed similar levels of genetic variation. In the host species, populations located near to each other were genetically similar; in both the host and the parasite species populations farther apart were significantly different. Thus, apparent dispersal barriers (i.e., high mountains) did not seem to restrict gene flow across populations as expected. Furthermore, there were no major differences in gene flow between the two species, perhaps because P. atrimandibularis parasitizes both alpine and lowland host species and annually migrates between alpine and lowland populations. The presence of strong gene flow in a system where local populations experience variable levels of selection pressure challenges the classical hypothesis that restricted gene flow is required for local adaptations to evolve.The co-evolutionary pathways followed by hosts and parasites strongly depend on the adaptive potential of antagonists and its underlying genetic architecture. Geographically structured populations of interacting species often experience local differences in the strength of reciprocal selection pressures, which can result in a geographic mosaic of co-evolution. One example of such a system is the boreo-montane social wasp Polistes biglumis and its social parasite Polistes atrimandibularis, which have evolved local defense and counter-defense mechanisms to match their antagonist. In this work, we study spatial genetic structure of P. biglumis and P. atrimandibularis populations at local and regional scales in the Alps, by using nuclear markers (DNA microsatellites, AFLP) and mitochondrial sequences. Both the host and the parasite populations harbored similar amounts of genetic variation. Host populations were not genetically structured at the local scale, but geographic regions were significantly differentiated from each other in both the host and the parasite in all markers. The net dispersal inferred from genetic differentiation was similar in the host and the parasite, which may be due to the annual migration pattern of the parasites between alpine and lowland populations. Thus, the apparent dispersal barriers (i.e., high mountains) do not restrict gene flow as expected and there are no important gene flow differences between the species, which contradict the hypothesis that restricted gene flow is required for local adaptations to evolve.

The geographic mosaic of arms race coevolution is closely matched to prey population structure.

Reciprocal adaptation is the hallmark of arms race coevolution. Local coadaptation between natural enemies should generate a geographic mosaic pattern where both species have roughly matched abilities across their shared range. However, mosaic variation in ecologically relevant traits can also arise from processes unrelated to reciprocal selection, such as population structure or local environmental conditions. We tested whether these alternative processes can account for trait variation in the geographic mosaic of arms race coevolution between resistant garter snakes (Thamnophis sirtalis) and toxic newts (Taricha granulosa). We found that predator resistance and prey toxin levels are functionally matched in co‐occurring populations, suggesting that mosaic variation in the armaments of both species results from the local pressures of reciprocal selection. By the same token, phenotypic and genetic variation in snake resistance deviates from neutral expectations of population genetic differentiation, showing a clear signature of adaptation to local toxin levels in newts. Contrastingly, newt toxin levels are best predicted by genetic differentiation among newt populations, and to a lesser extent, by the local environment and snake resistance. Exaggerated armaments suggest that coevolution occurs in certain hotspots, but prey population structure seems to be of particular influence on local phenotypic variation in both species throughout the geographic mosaic. Our results imply that processes other than reciprocal selection, like historical biogeography and environmental pressures, represent an important source of variation in the geographic mosaic of coevolution. Such a pattern supports the role of “trait remixing” in the geographic mosaic theory, the process by which non‐adaptive forces dictate spatial variation in the interactions among species.

A small‐tiled geographic mosaic of coevolution between Eurosta solidaginis and its natural enemies and host plant

AbstractWe identified a geographic mosaic of Eurosta solidaginis fly traits produced by coevolution of the stem gall‐forming fly with both its natural enemies and its host plants at small geographic scales in the presence of gene flow. These tritrophic interactions between the fly with its natural enemies and with its host‐plant Solidago altissima produced what has been termed a small‐tiled geographic mosaic of coevolution. Selection on gall diameter and length varies between prairie and forest habitats due to differences in host plants and natural‐enemy communities. At the prairie–forest ecotone where prairie and forest habitats are intermixed, we found that geographic selection mosaics on gall diameter and length varied on a scale of a few kilometers. Gall diameter variation among sites correlated with selection on gall diameters, indicating local adaptation. In contrast, gall lengths did not correlate with selection, indicating that gene flow may have prevented local adaptation of this trait. Eastern (forest) and western (prairie) subspecies of E. solidaginis have been proposed based on fly wing patterns, and these had intermediate forms in the ecotone indicating gene flow between these subspecies. Variation in wing patterns correlated with gall diameter, indicating that gene flow between prairie and forest fly populations may influence the distribution of gall traits. The ratio of forest to prairie vegetation increases with latitude, but there was no indication of latitudinal clines in gall or wing traits. Our results indicate that selection for differentiation in coevolved traits is strong enough to overcome gene flow in small tiles of habitat. The result is that ecological forces produce a dynamic mosaic of genetically differentiated locally adapted populations. It also indicates that prairie and forest host races of E. solidaginis form a mosaic hybrid zone in this region.

Geographical variation in cone traits of Pinus nigra as result of crossbill predation: implications for assisted migration

The interaction between Spanish Black pine (Pinus nigra subsp. salzmannii) and common crossbill (Loxia curvirostra) could have resulted in a geographic mosaic of coevolution in the Iberian Peninsula. In turn, this coevolutionary interaction might limit the future applicability of assisted migration as a tool to adapt forests of Spanish Black pine to ongoing climate change. Several cone and seed traits were measured in 740 cones collected at six stands distributed across the range of Spanish Black pine. Data were analysed by means of linear-mixed effects models, canonical analysis of principal coordinates and random-forest analysis. Results showed that cones from the northern pine populations exhibited more defensive traits against crossbill predation than cones from the central and southern populations, resulting in a geographic mosaic of selection. Assisted migration of P. nigra would involve moving southern populations toward the north of the species’ range and would result in introducing the genotypes of pines with poorly defended cones into areas where crossbills are comparatively more abundant. The resulting risks for the regeneration prospects of the translocated trees are discussed.

The geographic mosaic of coevolution in mutualistic networks

Ecological interactions shape adaptations through coevolution not only between pairs of species but also through entire multispecies assemblages. Local coevolution can then be further altered through spatial processes that have been formally partitioned in the geographic mosaic theory of coevolution. A major current challenge is to understand the spatial patterns of coadaptation that emerge across ecosystems through the interplay between gene flow and selection in networks of interacting species. Here, we combine a coevolutionary model, network theory, and empirical information on species interactions to investigate how gene flow and geographical variation in selection affect trait patterns in mutualistic networks. We show that gene flow has the surprising effect of favoring trait matching, especially among generalist species in species-rich networks typical of pollination and seed dispersal interactions. Using an analytical approximation of our model, we demonstrate that gene flow promotes trait matching by making the adaptive landscapes of different species more similar to each other. We use this result to show that the progressive loss of gene flow associated with habitat fragmentation may undermine coadaptation in mutualisms. Our results therefore provide predictions of how spatial processes shape the evolution of species-rich interactions and how the widespread fragmentation of natural landscapes may modify the coevolutionary process.

Diversification of Trait Combinations in Coevolving Plant and Insect Lineages.

Closely related species often have similar traits and sometimes interact with the same species. A crucial problem in evolutionary ecology is therefore to understand how coevolving species diverge when they interact with a set of closely related species from another lineage rather than with a single species. We evaluated geographic differences in the floral morphology of all woodland star plant species (Lithophragma, Saxifragaceae) that are pollinated by Greya (Prodoxidae) moths. Flowers of each woodland star species differed depending on whether plants interact locally with one, two, or no pollinating moth species. Plants of one species grown in six different environments showed few differences in floral traits, suggesting that the geographic differences are not due significantly to trait plasticity. Greya moth populations also showed significant geographic divergence in morphology, depending on the local host and on whether the moth species co-occurred locally. Divergence in the plants and the moths involved shifts in combinations of partially correlated traits, rather than any one trait. The results indicate that the geographic mosaic of coevolution can be amplified as coevolving lineages diversify into separate species and come together in different combinations in different ecosystems.

Genome divergence and diversification within a geographic mosaic of coevolution.

Despite substantial interest in coevolution's role in diversification, examples of coevolution contributing to speciation have been elusive. Here, we build upon past studies that have shown both coevolution between South Hills crossbills and lodgepole pine (Pinus contorta), and high levels of reproductive isolation between South Hills crossbills and other ecotypes in the North American red crossbill (Loxia curvirostra) complex. We used genotyping by sequencing to generate population genomic data and applied phylogenetic and population genetic analyses to characterize the genetic structure within and among nine of the ecotypes. Although genome-wide divergence was slight between ecotypes (FST =0.011-0.035), we found evidence of relative genetic differentiation (as measured by FST ) between and genetic cohesiveness within many of them. As expected for nomadic and opportunistic breeders, we detected no evidence of isolation by distance. The one sedentary ecotype, the South Hills crossbill, was genetically most distinct because of elevated divergence at a small number of loci rather than pronounced overall genome-wide divergence. These findings suggest that mechanisms related to recent local coevolution between South Hills crossbills and lodgepole pine (e.g. strong resource-based density dependence limiting gene flow) have been associated with genome divergence in the face of gene flow. Our results further characterize a striking example of coevolution driving speciation within perhaps as little as 6000years.

Invasive species, disrupted chemical community dynamics and future adaptations: Commentary on Chabaane et al. 2015

Chabaane et al. (2015) describe the subtle disruptive chemical changes that an alien herbivore (a moth caterpillar, Spodoptera littoralis) may cause in a native community. This disruption is described across trophic links from the plant it eats (a mustard family species, Brassica rapa) to native herbivores (the cabbage white butterfly, Pieris brassicae) and their native natural enemies (parasitic wasps, Cotesia glomerata). This paper reinforces the concern that new insect species arriving in natural communities may potentially have a detrimental impact: inducing unexpected changes in plant chemistry; altering competitive interactions with other herbivores; and changing the dynamics of the natural enemies of such herbivores. Extreme disruptions by ‘invasive’ Lepidoptera include pest population outbreaks that may cause large-scale defoliations which affect the entire ecosystem by cascading chemical, physiological and ecological changes across all trophic levels. In consequence altering the functional dynamics in various ways (Redman & Scriber 2000; Kenis et al. 2009; Gandi & Herms 2010). The study by Chabaane et al. (2015) shows how subtle changes in chemical cues from alien herbivores may result in major changes with cascading responses of the native plant–herbivore–parasite community. Whether such chemically based disruptions such as those described by Chabaane et al. (2015) result in local population outbreaks, and whether such changes become part of the future community dynamics, depends on the intensity and variability of natural selection that occurs. All of these biotic interactions resulting from invasive species are shaped by a combination of ecological and evolutionary processes (Moran & Alexander 2014). The fundamental component at all levels (organism, population, community and ecosystem) is the underlying genetic variability, which both interacts with and shapes all aspects of the phenotypes (phytochemical, biochemical, behavioural, physiological and morphological traits) and the potential for rapid adaptation and introgressive gene flow (Brasier 2001; Scriber 2011; Franks & Hoffmann 2012; Pauls et al. 2013). Plant genotypes can shape communities and drive evolutionary change (‘community genetics’; Smith et al. 2011). Similarly, underlying genetics shape interactions and natural selection across different environmental gradients at the landscape and ecosystem levels (‘geographic mosaic of co-evolution’; Thompson 2005). Understanding the underlying chemical, ecological and genetic sources of evolutionary responses to invasive species in the face of rapid environmental changes (such as global warming) will become one of the most significant challenges in the next decade (Moran & Alexander 2014; Scriber 2014). There is clearly a need for additional studies such as Chabaane et al. (2015) because the impacts of introduced species may run contrary to expectation. Illustrated by the fact that the alien (potentially invasive) herbivore species caused disruptive impacts on the community via induced changes in plant chemistry. This occurred despite the fact that exotic herbivores themselves were not actually used as hosts by the native parasitoids. Chabaane et al. (2015) show that exotic herbivores may invade new environments where they can significantly alter the native communities. This is done via both direct (induced changes in plant phytochemistry or nutritional ecology; Scriber 2002; Gandi & Herms 2010) and indirect (cascading) impacts on species across trophic levels in the ecosystem (Redman & Scriber 2000; Dicke & Baldwin 2012; Stam et al. 2014). The chemical basis of these plant/herbivore/enemy interactions has historically focused on host plant resistance, herbivore counter-adaptations and chemical defenses derived from the plants or herbivores in a relatively recent field of chemical ecology (Price et al. 1980; Feeny 1992; Gols 2014). Changes in other abiotic factors (e.g. climate and local drought conditions, as well as stoichiochemistry; Mattson & Scriber 1987; Mattson & Haack 1987; Elser et al. 2000; Chunco 2014) can also greatly alter the chemical ecology of the community, including interactions of plants, herbivores and their enemies (predators, parasites, and pathogens: Stamp 1993; Ayres & Lombardero 2000; Singer & Stireman 2005). However, Chabaane et al. (2015) show that changes in the chemical environment are not always ecologically predictable. More ground-breaking research as described in the interactions of chemical components among native and exotics across local populations and trophic levels will be required in the future (Chabaane et al. 2015). However, the elucidation of the genetic basis of ecological ripples caused by allelochemical interactions will likely be the next major step in better understanding the autecological (organism & population) causes of synecological responses in the community and ecosystem (Whitham et al. 2003; Wheat et al. 2007; Stam et al. *Correspondence author: E-mail: scriber@msu.edu

The Spatial Structure of Antagonistic Species Affects Coevolution in Predictable Ways

A current challenge in evolutionary ecology is to assess how the spatial structure of interacting species shapes coevolution. Previous work on the geographic mosaic of coevolution has shown that coevolution depends on the spatial structure, the strength of selection, and gene flow across populations. We used spatial subgraphs and coevolutionary models to evaluate how spatial structure and the location of coevolutionary hotspots (sites in which reciprocal selection occurs) and coldspots (sites in which unidirectional selection occurs) contribute to the dynamics of coevolution and the maintenance of polymorphisms. Specifically, we developed a new approach based on the Laplacian matrices of spatial subgraphs to explore the tendency of interacting species to evolve toward stable polymorphisms. Despite the complex interplay between gene flow and the strength of reciprocal selection, simple rules drive coevolution in small groups of spatially structured interacting populations. Hotspot location and the spatial organization of coldspots are crucial for understanding patterns in the maintenance of polymorphisms. Moreover, the degree of spatial variation in the outcomes of the coevolutionary process can be predicted from the network pattern of gene flow among sites. Our work provides us with novel tools that can be used in the field or the laboratory to predict the effects of spatial structure on coevolutionary trajectories.

Genotype × genotype interactions between the toxic cyanobacterium Microcystis and its grazer, the waterflea Daphnia.

Toxic algal blooms are an important problem worldwide. The literature on toxic cyanobacteria blooms in inland waters reports widely divergent results on whether zooplankton can control cyanobacteria blooms or cyanobacteria suppress zooplankton by their toxins. Here we test whether this may be due to genotype × genotype interactions, in which interactions between the large-bodied and efficient grazer Daphnia and the widespread cyanobacterium Microcystis are not only dependent on Microcystis strain or Daphnia genotype but are specific to genotype × genotype combinations. We show that genotype × genotype interactions are important in explaining mortality in short-time exposures of Daphnia to Microcystis. These genotype × genotype interactions may result in local coadaptation and a geographic mosaic of coevolution. Genotype × genotype interactions can explain why the literature on zooplankton–cyanobacteria interactions is seemingly inconsistent, and provide hope that zooplankton can contribute to the suppression of cyanobacteria blooms in restoration projects.

Low levels of population genetic structure in Pinus contorta (Pinaceae) across a geographic mosaic of co‐evolution

Population genetic analyses provide information on the population context in which evolutionary processes operate and are important for understanding the evolution of geographically variable traits. Earlier studies showed that cone structure of lodgepole pine in the Rocky Mountains diverged among populations because of geographic variation in coevolutionary interactions involving mammalian and avian seed predators. Analyses of population genetic variation are needed to determine whether this divergence has arisen despite extensive gene flow and whether populations to the east and west of the Rocky Mountains have evolved convergent phenotypes independently. We investigated genetic structuring across 22 stands of lodgepole pine in the central Rocky Mountains and in isolated peripheral populations that experience different seed predators and exhibit parallel divergence in cone traits using a set of nine simple sequence repeats and 235 AFLP loci. Our analyses reveal high levels of genetic diversity within and low genetic differentiation among populations. Nonetheless, geographic and genetic distances were correlated, and isolated populations to the east and west of the Rocky Mountains had higher levels of differentiation than did populations in the central part of the range. These data indicate not only that adaptive divergence of cone traits across a geographic mosaic of coevolution has occurred despite minimal genetic differentiation, but also that isolated populations to the east and west of the Rocky Mountains have evolved distinctive cones independently and in parallel. The population structure quantified here will inform future research aimed at detecting genetic variants associated with divergent adaptive traits.

Interactions among plants and evolution

Summary 1. Interactions among plants and their consumers, pollinators and dispersers are central to evolutionary theory, but interactions among plants themselves have received much less attention. Thus focusing more attention on the evolutionary role of plant–plant interactions may provide greater insight into the processes that organize communities. 2. Here, we integrate divergent themes in the literature in an effort to provide a synthesis of empirical evidence and ideas about how plant interactions may affect evolution and how evolution may affect plant interactions. 3. First, we discuss the idea of niche partitioning evolving through competitive interactions among plants, the idea of niche construction evolving through facilitative interactions, and the connections between these ideas and more recent research on diversity and ecosystem function and trait‐based community organization. 4. We then review how a history of coexistence within a region might affect competitive outcomes and explore the mechanisms by which plants exert selective forces on each other. Next, we consider recent research on invasions suggesting that plant interactions can reflect regional evolutionary trajectories. Finally, we place these lines of research into the context of extended phenotypes and the geographic mosaic of co‐evolution. 5. Synthesis. Our synthesis of separate lines of inquiry is a step towards understanding the evolutionary importance of interactions among plants, and suggests that the evolutionary consequences of interactions contribute to communities that are more than assemblages of independent populations.

A geographical mosaic of coevolution in a slave‐making host–parasite system

Three different isolated populations of the slave-making ant Rossomyrmex minuchae, sympatric with its obligate host Proformica longiseta, are known from the high mountains of southern Spain. To test the prediction that the slave-maker and its host represent a coevolutionary geographical mosaic, we studied the variation in the cuticular hydrocarbons (CHCs) as the trait most likely to show the selection mosaic, plus trait remixing by the gene flow in the populations of each species by means of microsatellites. We found within populations, host and parasite had more similar CHC profiles than between the populations or between parasites and allopatric hosts. The differences between the CHC profiles of the host and parasite, which may be responsible for the level of tolerance towards the parasite, varied between the populations suggesting the existence of a selection mosaic of coevolution. Furthermore, P.longiseta showed higher gene flow than R.minuchae, which would allow local variation in the coevolution of the host and parasite while allowing some trait remixing.

Spatial heterogeneity, frequency-dependent selection and polymorphism in host-parasite interactions

BackgroundGenomic and pathology analysis has revealed enormous diversity in genes involved in disease, including those encoding host resistance and parasite effectors (also known in plant pathology as avirulence genes). It has been proposed that such variation may persist when an organism exists in a spatially structured metapopulation, following the geographic mosaic of coevolution. Here, we study gene-for-gene relationships governing the outcome of plant-parasite interactions in a spatially structured system and, in particular, investigate the population genetic processes which maintain balanced polymorphism in both species.ResultsFollowing previous theory on the effect of heterogeneous environments on maintenance of polymorphism, we analysed a model with two demes in which the demes have different environments and are coupled by gene flow. Environmental variation is manifested by different coefficients of natural selection, the costs to the host of resistance and to the parasite of virulence, the cost to the host of being diseased and the cost to an avirulent parasite of unsuccessfully attacking a resistant host. We show that migration generates negative direct frequency-dependent selection, a condition for maintenance of stable polymorphism in each deme. Balanced polymorphism occurs preferentially if there is heterogeneity for costs of resistance and virulence alleles among populations and to a lesser extent if there is variation in the cost to the host of being diseased. We show that the four fitness costs control the natural frequency of oscillation of host resistance and parasite avirulence alleles. If demes have different costs, their frequencies of oscillation differ and when coupled by gene flow, there is amplitude death of the oscillations in each deme. Numerical simulations show that for a multiple deme island model, costs of resistance and virulence need not to be present in each deme for stable polymorphism to occur.ConclusionsOur theoretical results confirm the importance of empirical studies for measuring the environmental heterogeneity for genetic costs of resistance and virulence alleles. We suggest that such studies should be developed to investigate the generality of this mechanism for the long-term maintenance of genetic diversity at host and parasite genes.

The potential of a population genomics approach to analyse geographic mosaics of plant--insect coevolution

A central issue in the evolutionary ecology of species interactions is coevolution, which involves the reciprocal selection between individuals of interacting species. Understanding the importance of coevolution in shaping species interactions requires the consideration of spatial variation in their strength. This is exactly what the, recently developed, geographic mosaic theory of coevolution addresses. Another major development in the study of population ecology is the introduction of the population genomics approach in this field of research. This approach addresses spatial processes through molecular methods. It is of particular interest that population genomics is especially applicable to natural populations of non-model species. We describe how population genomics can be used in the context of the geographic mosaic of coevolution, specifically to identify coevolutionary hot-spots, and to attribute genetic variation found at specific loci to processes of selection versus trait remixing. The proposed integration of the population genomics approach with the conceptual framework of the geographic mosaic of coevolution is illustrated with a few selected, particularly demonstrative, examples from the realm of insect--plant interactions.

Genetic diversity, population structure and sex‐biased dispersal in three co‐evolving species

Genetic diversity and spatial structure of populations are important for antagonistic coevolution. We investigated genetic variation and population structure of three closely related European ant species: the social parasite Harpagoxenus sublaevis and its two host species Leptothorax acervorum and Leptothorax muscorum. We sampled populations in 12 countries and analysed eight microsatellite loci and an mtDNA sequence. We found high levels of genetic variation in all three species, only slightly less variation in the host L. muscorum. Using a newly introduced measure of differentiation (Jost's D(est)), we detected strong population structuring in all species and less male-biased dispersal than previously thought. We found no phylogeographic patterns that could give information on post-glacial colonization routes - northern populations are as variable as more southern populations. We conclude that conditions for Thompson's geographic mosaic of coevolution are ideal in this system: all three species show ample genetic variation and strong population structure.

Fire Drives Transcontinental Variation in Tree Birch Defense against Browsing by Snowshoe Hares

Fire has been the dominant disturbance in boreal America since the Pleistocene, resulting in a spatial mosaic in which the most fire occurs in the continental northwest. Spatial variation in snowshoe hare (Lepus americanus) density reflects the fire mosaic. Because fire initiates secondary forest succession, a fire mosaic creates variation in the abundance of early successional plants that snowshoe hares eat in winter, leading to geographic variation in hare density. We hypothesize that fire is the template for a geographic mosaic of natural selection: where fire is greatest and hares are most abundant, hare browsing has most strongly selected juvenile-phase woody plants for defense. We tested the hypothesis at multiple spatial scales using Alaska birch (Betula neoalaskana) and white birch (Betula papyrifera). We also examined five alternative hypotheses for geographic variation in antibrowsing defense. The fire-hare-defense hypothesis was supported at transcontinental, regional, and local scales; alternative hypotheses were rejected. Our results link transcontinental variation in species interactions to an abiotic environmental driver, fire. Intakes of defense toxins by Alaskan hares exceed those by Wisconsin hares, suggesting that the proposed selection mosaic may coincide with a geographic mosaic of coevolution.

John N. Thompson, The Geographic Mosaic of Coevolution

John N. Thompson, The Geographic Mosaic of Coevolution