Genetic diversity and structure of the cycad Zamia loddigesii Miq. (Zamiaceae): implications for evolution and conservation

The genetic diversity and structure of four populations of the cycad Zamia loddigesii were studied throughout its range in Mexico. Allozyme electrophoresis of 15 loci was conducted. The mean number of alleles per locus was 1.80 ± 0.09, the percentage of polymorphic loci was 66.6 ± 5.4, and the expected heterozygosity was 0.266 ± 0.02. The results indicated that the genetic diversity was relatively higher, with respect to tropical tree species and other cycads. The genetic variation explained by differences among populations was 18%. On average, gene flow between paired populations was similar ( Nm  = 1.6) to other tropical forest trees and cycad species. Our results indicated that the geographical isolation among populations of Z. loddigesii generated allele loss, as well as a clinal variation in the frequencies of two loci ( MDH and MNR2 ), in relation to the latitudinal distribution of populations. The populations have become fragmented due to increasingly higher pressure of habitat conversion and disturbance. The importance of the establishment of sanctuaries and protected areas and a reduction in deforestation is highlighted in this research as a way of preserving the high genetic diversity of this and other endemic species.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 533–544.     

The influence of history and contemporary stream hydrology on the evolution of genetic diversity within species: an examination of microsatellite DNA variation in bull trout,Salvelinus confluentus (Pisces: Salmonidae)

An understanding of the relative roles of historical and contemporary factors in structuring genetic variation is a fundamental, but understudied aspect of geographic variation. We examined geographic variation in microsatellite DNA allele frequencies in bull trout (Salvelinus confluentus, Salmonidae) to test hypotheses concerning the relative roles of postglacial dispersal (historical) and current landscape features (contemporary) in structuring genetic variability and population differentiation. Bull trout exhibit relatively low intrapopulation microsatellite variation (average of 1.9 alleles per locus, average He = 0.24), but high levels of interpopulation divergence (F(ST) = 0.39). We found evidence of historical influences on microsatellite variation in the form of a decrease in the number of alleles and heterozygosities in populations on the periphery of the range relative to populations closer to putative glacial refugia. In addition, one region of British Columbia that was colonized later during deglaciation and by more indirect watershed connections showed less developed and more variable patterns of isolation by distance than a similar region colonized earlier and more directly from refugia. Current spatial and drainage interconnectedness among sites and the presence of migration barriers (falls and cascades) within individual streams were found to be important contemporary factors influencing historical patterns of genetic variability and interpopulation divergence. Our work illustrates the limited utility of equilibrium models to delineate population structure and patterns of genetic diversity in recently founded populations or those inhabiting highly heterogeneous environments, and it highlights the need for approaches incorporating a landscape context for population divergence. Substantial microsatellite DNA divergence among bull trout populations may also signal divergence in traits important to population persistence in specific environments.     

THE ECOLOGICAL GENETICS OF INTRODUCED POPULATIONS OF THE GIANT TOAD,BUFO MARINUS. III. GEOGRAPHICAL PATTERNS OF VARIATION

The allele frequencies at ten polymorphic loci are described from 31 Bufo marinus populations in the Moreton Bay region in southeastern Queensland, Australia and the variation of these is found to be non-random in all cases. The pattern of non-randomness varies among loci, being clinal in two instances. The allele frequencies at the same ten loci are also described for 12 populations sampled from throughout B. marinus' Australian range. The frequency variation on this larger geographical scale is non-random at all but two loci (Mpi and Hbdh) and also varies among loci, in this case being clinal in four instances. In both cases, the patterns of variation are most reasonably explained as having resulted from genetic drift occurring during the recent range expansion which B. marinus is known to have experienced in Australia. It seems that natural selection has played little, if any, role in generating the observed gene frequency patterns. These results emphasize the need for caution in interpreting geographical patterns of variation. They show that even when clinal patterns exist at some loci but not at others, one cannot conclude that the patterns result from natural selection, unless the demographic histories of the studied populations are known and are inconsistent with the alternative hypothesis that the patterns result from genetic drift.     

Divergence in coloration and ecological speciation in the Anolis marmoratus species complex

Adaptive divergence in coloration is expected to produce reproductive isolation in species that use colourful signals in mate choice and species recognition. Indeed, many adaptive radiations are characterized by differentiation in colourful signals, suggesting that divergent selection acting on coloration may be an important component of speciation. Populations in the Anolis marmoratus species complex from the Caribbean island of Guadeloupe display striking divergence in the colour and pattern of adult males that occurs over small geographic distances, suggesting strong divergent selection. Here we test the hypothesis that divergence in coloration results in reduced gene flow among populations. We quantify variation in adult male coloration across a habitat gradient between mesic and xeric habitats, use a multilocus coalescent approach to infer historical demographic parameters of divergence, and examine gene flow and population structure using microsatellite variation. We find that colour variation evolved without geographic isolation and in the face of gene flow, consistent with strong divergent selection and that both ecological and sexual selection are implicated. However, we find no significant differentiation at microsatellite loci across populations, suggesting little reproductive isolation and high levels of contemporary gene exchange. Strong divergent selection on loci affecting coloration probably maintains clinal phenotypic variation despite high gene flow at neutral loci, supporting the notion of a porous genome in which adaptive portions of the genome remain fixed whereas neutral portions are homogenized by gene flow and recombination. We discuss the impact of these findings for studies of colour evolution and ecological speciation.     

Microsatellite analysis of the phylogeography, Pleistocene history and secondary contact hypotheses for the killifish, Fundulus heteroclitus

The mummichog, Fundulus heteroclitus, exhibits extensive latitudinal clinal variation in a number of physiological and biochemical traits, coupled with phylogeographical patterns at mitochondrial and nuclear DNA loci that suggest a complicated history of spatially variable selection and secondary intergradation. This species continues to serve as a model for understanding local and regional adaptation to variable environments. Resolving the influences of historical processes on the distribution of genetic variation within and among extant populations of F. heteroclitus is crucial to a better understanding of how populations evolve in the context of contemporary environments. In this study, we analysed geographical patterns of genetic variation at eight microsatellite loci among 15 populations of F. heteroclitus distributed throughout the North American range of the species from Nova Scotia to Georgia. Genetic variation in Northern populations was lower than in Southern populations and was strongly correlated with latitude throughout the species range. The most common Northern alleles at all eight loci exhibited concordant latitudinal clinal patterns, and the existence of an abrupt transition zone in allele frequencies between Northern and Southern populations was similar to that observed for mitochondrial DNA and allozyme loci. A significant pattern of isolation by distance was observed both within and between northern and southern regions. This pattern was unexpected, particularly for northern populations, given the recent colonization history of post-Pleistocene habitats, and was inconsistent with either a recent northward population expansion or a geographically restricted northern Pleistocene refugium. The data provided no evidence for recent population bottlenecks, and estimates of historical effective population sizes suggest that post-Pleistocene populations have been large throughout the species distribution. These results suggest that F. heteroclitus was broadly distributed throughout most of its current range during the last glacial event and that the abrupt transition in allele frequencies that separate Northern and Southern populations may reflect regional disequilibrium conditions associated with the post-Pleistocene colonization history of habitats in that region.     

When can refuges mediate the genetic effects of fire regimes? A simulation study of the effects of topography and weather on neutral and adaptive genetic diversity in fire‐prone landscapes

Understanding how landscape heterogeneity mediates the effects of fire on biodiversity is increasingly important under global changes in fire regimes. We used a simulation experiment to investigate how fire regimes interact with topography and weather to shape neutral and selection‐driven genetic diversity under alternative dispersal scenarios, and to explore the conditions under which microrefuges can maintain genetic diversity of populations exposed to recurrent fire. Spatial heterogeneity in simulated fire frequency occurred in topographically complex landscapes, with fire refuges and fire‐prone “hotspots” apparent. Interannual weather variability reduced the effect of topography on fire patterns, with refuges less apparent under high weather variability. Neutral genetic diversity was correlated with long‐term fire frequency under spatially heterogeneous fire regimes, being higher in fire refuges than fire‐prone areas, except under high dispersal or low fire severity (low mortality). This generated different spatial genetic structures in fire‐prone and fire‐refuge components of the landscape, despite similar dispersal. In contrast, genetic diversity was only associated with time since the most recent fire in flat landscapes without predictable refuges and hotspots. Genetic effects of selection driven by fire‐related conditions depended on selection pressure, migration distance and spatial heterogeneity in fire regimes. Allele frequencies at a locus conferring higher fitness under successional environmental conditions followed a pattern of “temporal adaptation” to contemporary conditions under strong selection pressure and high migration. However, selected allele frequencies were correlated with spatial variation in long‐term mean fire frequency (relating to environmental predictability) under weak dispersal, low selection pressure and strong spatial heterogeneity in fire regimes.     

Genetic analysis of differentiation among breeding ponds reveals a candidate gene for local adaptation in Rana arvalis

One of the main questions in evolutionary and conservation biology is how geographical and environmental features of the landscape shape neutral and adaptive genetic variation in natural populations. The identification of genomic polymorphisms that account for adaptive variation can aid in finding candidate loci for local adaptation. Consequently, a comparison of spatial patterns in neutral markers and loci under selection may help disentangle the effects of gene flow, genetic drift and selection at the landscape scale. Many amphibians breed in wetlands, which differ in environmental conditions and in the degree of isolation, enhancing the potential for local adaptation. We used microsatellite markers to measure genetic differentiation among 17 local populations of Rana arvalis breeding in a network of wetlands. We found that locus RC08604 deviated from neutral expectations, suggesting that it is a good candidate for directional selection. We used a genetic network analysis to show that the allele distribution in this locus is correlated with habitat characteristics, whereas this was not the case at neutral markers that displayed a different allele distribution and population network in the study area. The graph approach illustrated the genomic heterogeneity (neutral loci vs. the candidate locus for directional selection) of gene exchange and genetic divergence among populations under directional selection. Limited gene flow between wetlands was only observed at the candidate genomic region under directional selection. RC08604 is partially located inside an up‐regulated thyroid‐hormone receptor (TRβ) gene coordinating the expression of other genes during metamorphosis and appears to be linked with variation in larval life‐history traits found among R. arvalis populations. We suggest that directional selection on genes coding larval life‐history traits is strong enough to maintain the divergence in these genomic regions, reducing the effective recombination of locally adapted alleles but not in other regions of the genome. Integrating this knowledge into conservation plans at the landscape scale will improve the design of management strategies to preserve adaptive genetic diversity in wetland networks.     

Response of a southern temperate marsupial, the Tasmanian pademelon (Thylogale billardierii), to historical and contemporary forest fragmentation

Despite only limited Pleistocene glacial activity in the southern hemisphere, temperate forest species experienced complex distributional changes resulting from the combined effects of glaciation, sea level change and increased aridity. The effects of these historical processes on population genetic structure are now overlain by the effects of contemporary habitat modification. In this study, 10 microsatellites and 629 bp of the mitochondrial control region were used to assess the effects of historical forest fragmentation and recent anthropogenic habitat change on the broad-scale population genetic structuring of a southern temperate marsupial, the Tasmanian pademelon. A total of 200 individuals were sampled from seven sites across Tasmania and two islands in Bass Strait. High mitochondrial and nuclear genetic diversity indicated the maintenance of large historical population sizes. There was weak phylogeographical structuring of haplotypes, although all King Island haplotypes and three Tasmanian haplotypes formed a divergent clade implying the mid-Pleistocene isolation of a far northwestern population. Both the mitochondrial and nuclear data indicated a division of Tasmanian populations into eastern and western regions. This was consistent with a historical barrier resulting from increased aridity in the lowland 'midlands' region during glacial periods, and with a contemporary barrier resulting from recent habitat modification in that region. In Tasmania, gene flow appears to have been relatively unrestricted during glacial maxima in the west, while in the east there was evidence for historical expansion from at least one large glacial refuge and recolonization of Flinders Island.     

Guppy populations differ in cannibalistic degree and adaptation to structural environments

There is considerable variation in cannibalism between different species and also between individuals of different species, although relatively little is known about what creates this variation. We investigated the degree of cannibalism in guppy (Poecilia reticulata) populations originating from high and low predation environments in Trinidad, and also how cannibalism was affected by the presence of refuges. Females from two populations were allowed to feed on juveniles from two populations in aquaria trials. The cannibalism was size-dependent and varied depending on both juvenile and female origin. Low predation females were more efficient cannibals and low predation juveniles were better at avoiding cannibalism compared to high predation guppies when no refuges were present. The high predation females were superior cannibals and the high predation juveniles were better at escaping cannibalism than the low predation guppies when refuges were present. We discuss whether the differences in cannibalism and response to refuge addition relate to predation-induced habitat shifts and differences in the guppies’ natural environment.     

Repeated sampling detects gene flow in a flightless ground beetle in a fragmented landscape

Secondary clines level down in the course of time if the gene flow is not interrupted. Temporally repeated sampling of populations in a cline allows the investigation not only of its occurrence but also of the estimation of the amount of ongoing gene flow. We reinvestigated an allozyme gradient in Carabus auronitens populations in the Westphalian Lowlands (northwestern Germany) 15 to 20 years after it had originally been recorded. A total of 977 individuals of this flightless woodland species from 29 sample sites were genotyped at the diallelic Est-1 locus in 2005-2006 and compared to former findings, collected in 1985-1994 from the same populations. Both data sets showed clinal variation. Pairwise differences between the samples of both data sets indicated significant decrease in the steepness of the cline during the past 15 to 20 years. The estimated average gene flow per generation is 0.6% of each beetle population. Ongoing gene flow in the flightless ground beetle C. auronitens led to a less pronounced cline despite a stable degree of fragmentation (and connectivity) of the landscape. Migration and gene flow were obviously enabled by the numerous hedgerows. The corridors are seen to be a prerequisite for migration between populations and for possible future range shifts of forest insect species.     

Latitudinal allozymic variability in natural populations ofZaprionus indianus from India

Twelve natural populations ofZ. indianus from India, collected along lines of latitude, were analyzed electrophoretically and the geographical pattern of variation was compared at 15 allozyme loci. Except forMdh-2, all the loci revealed extensive polymorphism. The allele frequency variation revealed a statistically significant correlation as well as regression coefficients with the latitude. The populations revealed extensive clinal variation at all the polymorphic loci, which depicted moderate to higherF _ST values. Thus, the observed genetic divergence in colonizing populations ofZ. indianus shows the role of natural selection that varies spatially along the north-south axis of the Indian subcontinent.Financial assistance from UGC, New Delhi, is gratefully acknowledged.     

The effects of genetic drift during range expansion on geographical patterns of variation: a computer simulation of the colonization of Australia by Bufo marinus

A computer simulation is performed of allele frequency changes resulting from genetic drift at eleven loci during the probable course of colonization of Australia by populations of the Giant Toad, Bufo marinus. The history of twelve populations for which allele frequency data are available is modelled. Account is taken of the likely pattern of relationship among the populations, the effective size of the populations (as indicated by the observed variance of allele frequencies) and the probable isolation of the populations from each other following their separation. In all simulations, allele frequencies at some loci show a significant association with latitude while others do not. In six of ten simulations, there is a significant association between degree of variation at a locus and the presence of a latitudinal cline of allele frequencies. There are also indications of this kind of association in simulations of a uni-directional range expansion. These results demonstrate that such associations, which were also observed in the data from the actual populations, can result from genetic drift during a range expansion, and therefore cannot be taken as evidence of the action of natural selection.     

Characterization of microsatellite loci in the fungus, Grosmannia clavigera, a pine pathogen associated with the mountain pine beetle

The largest forest pest epidemic in Canadian history caused by the mountain pine beetle (MPB) and its fungal associates has killed over 15 million hectares of forest. Sixty simple sequence repeat regions were identified from Grosmannia clavigera, an MPB associated fungus. Eight loci genotyped in 53 isolates from two populations in British Columbia, Canada revealed three to 10 alleles per locus and gene diversities of 0 to 0.79. All but two of these loci showed length polymorphism in Leptographium longiclavatum, a related MPB fungal associate. These microsatellites will be useful in population genetic studies of these fungi.     

Signatures of diversifying selection at EST‐SSR loci and association with climate in natural Eucalyptus populations

Understanding the environmental parameters that drive adaptation among populations is important in predicting how species may respond to global climatic changes and how gene pools might be managed to conserve adaptive genetic diversity. Here, we used Bayesian F_ST outlier tests and allele–climate association analyses to reveal two Eucalyptus EST‐SSR loci as strong candidates for diversifying selection in natural populations of a southwestern Australian forest tree, Eucalyptus gomphocephala (Myrtaceae). The Eucalyptus homolog of a CONSTANS‐like gene was an F_ST outlier, and allelic variation showed significant latitudinal clinal associations with annual and winter solar radiation, potential evaporation, summer precipitation and aridity. A second F_ST outlier locus, homologous to quinone oxidoreductase, was significantly associated with measures of temperature range, high summer temperature and summer solar radiation, with important implications for predicting the effect of temperature on natural populations in the context of climate change. We complemented these data with investigations into neutral population genetic structure and diversity throughout the species range. This study provides an investigation into selection signatures at gene‐homologous EST‐SSRs in natural Eucalyptus populations, and contributes to our understanding of the relationship between climate and adaptive genetic variation, informing the conservation of both putatively neutral and adaptive components of genetic diversity.     

The role of refuges in the persistence of Australian dryland mammals

Irruptive population dynamics are characteristic of a wide range of fauna in the world's arid (dryland) regions. Recent evidence indicates that regional persistence of irruptive species, particularly small mammals, during the extensive dry periods of unpredictable length that occur between resource pulses in drylands occurs as a result of the presence of refuge habitats or refuge patches into which populations contract during dry (bust) periods. These small dry‐period populations act as a source of animals when recolonisation of the surrounding habitat occurs during and after subsequent resource pulses (booms). The refuges used by irruptive dryland fauna differ in temporal and spatial scale from the refugia to which species contract in response to changing climate. Refuges of dryland fauna operate over timescales of months and years, whereas refugia operate on timescales of millennia over which evolutionary divergence may occur. Protection and management of refuge patches and refuge habitats should be a priority for the conservation of dryland‐dwelling fauna. This urgency is driven by recognition that disturbance to refuges can lead to the extinction of local populations and, if disturbance is widespread, entire species. Despite the apparent significance of dryland refuges for conservation management, these sites remain poorly understood ecologically. Here, we synthesise available information on the refuges of dryland‐dwelling fauna, using Australian mammals as a case study to provide focus, and document a research agenda for increasing this knowledge base. We develop a typology of refuges that recognises two main types of refuge: fixed and shifting. We outline a suite of models of fixed refuges on the basis of stability in occupancy between and within successive bust phases of population cycles. To illustrate the breadth of refuge types we provide case studies of refuge use in three species of dryland mammal: plains mouse (Pseudomys australis), central rock‐rat (Zyzomys pedunculatus), and spinifex hopping‐mouse (Notomys alexis). We suggest that future research should focus on understanding the species‐specific nature of refuge use and the spatial ecology of refuges with a focus on connectivity and potential metapopulation dynamics. Assessing refuge quality and understanding the threats to high‐quality refuge patches and habitat should also be a priority. To facilitate this understanding we develop a three‐step methodology for determining species‐specific refuge location and habitat attributes. This review is necessarily focussed on dryland mammals in continental Australia where most refuge‐based research has been undertaken. The applicability of the refuge concept and the importance of refuges for dryland fauna conservation elsewhere in the world should be investigated. We predict that refuge‐using mammals will be widespread particularly among dryland areas with unpredictable rainfall patterns.     

Genetics of Lupinus

Summary Preliminary surveys indicate that most populations of Lupinus succulentus are genetically monomorphic for three flower color loci, viz., BB PP DD. In one small geographic area, a number of populations were polymorphic for the D/d locus. In this case, clinal variation and seasonal variation were found. The S/s locus, affecting seed coat pattern, was polymorphic in a large majority of populations and genetic variation at this locus is present in all but a very few colonies. When subdivisions of five large populations were studied, genetic differentiation was found within two of these populations. Gene frequencies tended toward the limits of zero or one in populations from relatively undisturbed sites but tended toward intermediate values in the more disturbed ruderal populations presumed to be recently colonized. It is concluded that recently colonized populations are not always genetically uniform.This work supported in part by National Institutes of Health Grant GM 10476     

Disturbance and the role of refuges in mediterranean climate streams

Refuges protect plant and animal populations from disturbance. Knowledge of refuges from disturbance in mediterranean climate rivers (med-rivers) has increased the last decade. We review disturbance processes and their relationship to refuges in streams in mediterranean climate regions (med-regions). Med-river fauna show high endemicity and their populations are often exposed to disturbance; hence the critical importance of refuges during (both seasonal and supraseasonal) disturbances. Disturbance pressures are increasing in med-regions, in particular from climatic change, salinisation, sedimentation, water extraction, hydropower generation, supraseasonal drought, and wildfire. Med-rivers show annual cycles of constrained precipitation and predictable seasonal drying, causing the biota to depend on seasonal refuges, in particular, those that are spatially predictable. This creates a spatial and temporal mosaic of inundation that determines habitat extent and refuge function. Refuges of sufficient size and duration to maintain populations, such as perennially flowing reaches, sustain biodiversity and may harbour relict populations, particularly during increasing aridification, where little other suitable habitat remains in landscapes. Therefore, disturbances that threaten perennial flows potentially cascade disproportionately to reduce regional scale biodiversity in med-regions. Conservation approaches for med-river systems need to conserve both refuges and refuge connectivity, reduce the impact of anthropogenic disturbances and sustain predictable, seasonal flow patterns.     

Genetic structure of the forest herb Primula elatior in a changing landscape

To investigate whether changes in land use and associated forest patch turnover affected genetic diversity and structure of the forest herb Primula elatior, historical data on landscape changes were combined with a population genetic analysis using dominant amplified fragment length polymorphism markers. Based on nine topographic maps, landscape history was reconstructed and forest patches were assigned to two age classes: young (less than 35 years) and old (more than 35 years). The level of differentiation among Primula populations in recently established patches was compared with the level of differentiation among populations in older patches. Genetic diversity was independent of population size (P > 0.05). Most genetic variation was present within populations. Within-population diversity levels tended to be higher for populations located in older forests compared with those for populations located in young forests (Hj = 0.297 and 0.285, respectively). Total gene diversity was also higher for old than for young populations (Ht = 0.2987 and 0.2828, respectively). The global fixation index FST averaged over loci was low, but significant. Populations in older patches were significantly more differentiated from each other than were populations in recently established patches and they showed significant isolation by distance. In contrast, no significant correlations between pairwise geographical distance and FST were found for populations in recently established patches. The location of young and old populations in the studied system and altered gene flow because of increased population density and decreased inter-patch distances between extant populations may explain the observed lower genetic differentiation in the younger populations. This study exemplifies the importance of incorporating data on historical landscape changes in population genetic research at the landscape scale.     

Evolutionary genetics of Metridium senile. II. Geographic patterns of allozyme variation

Electrophoretic surveys have demonstrated that populations of the sea anemone Metridium senile along the northeast coast of the United States are polymorphic at four enzyme loci. Phosphoglucose isomerase (PGI) has two alleles in most populations, phosphoglucomutase (PGM) has three alleles, and two leucine aminopeptidase loci have two common alleles each. Phosphoglucose isomerase displays clinal variation and an apparent association with environmental temperature. Phosphoglucomutase shows clinal variation north of Cape Cod for two of the three alleles, while the two leucine aminopeptidase loci are not clinal. All loci show a great deal of variation in populations on Cape Cod, but there is no apparent systematic pattern to this variation. Temperature may be a selective agent in the maintenance of the PGI and PGM clines, although other possibilities cannot presently be completely excluded.Supported by Grant T-4 from the Health Research and Services Foundation, NSF DEB77-14442, NIH GM25809, and NIH GM28024.     

INTEGRATING LANDSCAPE GENOMICS AND SPATIALLY EXPLICIT APPROACHES TO DETECT LOCI UNDER SELECTION IN CLINAL POPULATIONS

Uncovering the genetic basis of adaptation hinges on the ability to detect loci under selection. However, population genomics outlier approaches to detect selected loci may be inappropriate for clinal populations or those with unclear population structure because they require that individuals be clustered into populations. An alternate approach, landscape genomics, uses individual‐based approaches to detect loci under selection and reveal potential environmental drivers of selection. We tested four landscape genomics methods on a simulated clinal population to determine their effectiveness at identifying a locus under varying selection strengths along an environmental gradient. We found all methods produced very low type I error rates across all selection strengths, but elevated type II error rates under “weak” selection. We then applied these methods to an AFLP genome scan of an alpine plant, Campanula barbata, and identified five highly supported candidate loci associated with precipitation variables. These loci also showed spatial autocorrelation and cline patterns indicative of selection along a precipitation gradient. Our results suggest that landscape genomics in combination with other spatial analyses provides a powerful approach for identifying loci potentially under selection and explaining spatially complex interactions between species and their environment.