From broadscale patterns to fine-scale processes: habitat structure influences genetic differentiation in the pitcher plant midge across multiple spatial scales

The spatial scale at which samples are collected and analysed influences the inferences that can be drawn from landscape genetic studies. We examined genetic structure and its landscape correlates in the pitcher plant midge, Metriocnemus knabi, an inhabitant of the purple pitcher plant, Sarracenia purpurea, across several spatial scales that are naturally delimited by the midge’s habitat (leaf, plant, cluster of plants, bog and system of bogs). We analysed 11 microsatellite loci in 710 M. knabi larvae from two systems of bogs in Algonquin Provincial Park (Canada) and tested the hypotheses that variables related to habitat structure are associated with genetic differentiation in this midge. Up to 54% of variation in individual‐based genetic distances at several scales was explained by broadscale landscape variables of bog size, pitcher plant density within bogs and connectivity of pitcher plant clusters. Our results indicate that oviposition behaviour of females at fine scales, as inferred from the spatial locations of full‐sib larvae, and spatially limited gene flow at broad scales represent the important processes underlying observed genetic patterns in M. knabi. Broadscale landscape features (bog size and plant density) appear to influence oviposition behaviour of midges, which in turn influences the patterns of genetic differentiation observed at both fine and broad scales. Thus, we inferred linkages among genetic patterns, landscape patterns and ecological processes across spatial scales in M. knabi. Our results reinforce the value of exploring such links simultaneously across multiple spatial scales and landscapes when investigating genetic diversity within a species.     

Changing importance of habitat structure across multiple spatial scales for three species of insects

Considerable scientific effort has gone into examining how the spatial structure of habitat influences organism distribution and abundance in both theoretical and applied contexts. An emerging conclusion from these works is that the overall amount of habitat in the landscape matters most for species persistence and that more local attributes of habitat structure such as the size and arrangement of patches is of secondary importance. In this study, we quantify how and when the effects of habitat configuration (patch size and isolation) influence the density of three species of insects (Order: Diptera; Wyeomyia smithii , Metriocnemus knabi , Fletcherimyia fletcheri ) whose larvae are found exclusively in identical habitats (the water-filled leaves of pitcher plants – Sarracenia purpurea ) in a system that is naturally patchy at multiple spatial scales. We illustrate that relationships with configuration exist regardless of the overall amount of habitat in the broader landscape, and that there are distinct changes in the relationship between insect density and habitat configuration across multiple spatial scales. In general, patch size is more important within the movement range of the individual and isolation is important at larger, aggregation scales. Thus we demonstrate that a) both the amount and configuration of habitat are important attributes of species distribution; b) responses to measures of configuration can be scaled to processes such as movement and c) that hierarchical frameworks extending across very broad scales are essential for understanding how species respond to habitat structure and their role in ecosystem function.     

Seascape drivers of Macrocystis pyrifera population genetic structure in the northeast Pacific

At small spatial and temporal scales, genetic differentiation is largely controlled by constraints on gene flow, while genetic diversity across a species' distribution is shaped on longer temporal and spatial scales. We assess the hypothesis that oceanographic transport and other seascape features explain different scales of genetic structure of giant kelp, Macrocystis pyrifera. We followed a hierarchical approach to perform a microsatellite‐based analysis of genetic differentiation in Macrocystis across its distribution in the northeast Pacific. We used seascape genetic approaches to identify large‐scale biogeographic population clusters and investigate whether they could be explained by oceanographic transport and other environmental drivers. We then modelled population genetic differentiation within clusters as a function of oceanographic transport and other environmental factors. Five geographic clusters were identified: Alaska/Canada, central California, continental Santa Barbara, California Channel Islands and mainland southern California/Baja California peninsula. The strongest break occurred between central and southern California, with mainland Santa Barbara sites forming a transition zone between the two. Breaks between clusters corresponded approximately to previously identified biogeographic breaks, but were not solely explained by oceanographic transport. An isolation‐by‐environment (IBE) pattern was observed where the northern and southern Channel Islands clustered together, but not with closer mainland sites, despite the greater distance between them. The strongest environmental association with this IBE pattern was observed with light extinction coefficient, which extends suitable habitat to deeper areas. Within clusters, we found support for previous results showing that oceanographic connectivity plays an important role in the population genetic structure of Macrocystis in the Northern hemisphere.     

Contrasting demographic and genetic estimates of dispersal in the endangered Coahuilan box turtle: a contemporary approach to conservation

The evolutionary viability of an endangered species depends upon gene flow among subpopulations and the degree of habitat patch connectivity. Contrasting population connectivity over ecological and evolutionary timescales may provide novel insight into what maintains genetic diversity within threatened species. We employed this integrative approach to evaluating dispersal in the critically endangered Coahuilan box turtle (Terrapene coahuila) that inhabits isolated wetlands in the desert‐spring ecosystem of Cuatro Ciénegas, Mexico. Recent wetland habitat loss has altered the spatial distribution and connectivity of habitat patches; and we therefore predicted that T. coahuila would exhibit limited movement relative to estimates of historic gene flow. To evaluate contemporary dispersal patterns, we employed mark–recapture techniques at both local (wetland complex) and regional (intercomplex) spatial scales. Gene flow estimates were obtained by surveying genetic variation at nine microsatellite loci in seven subpopulations located across the species’ geographical range. The mark–recapture results at the local spatial scale reveal frequent movement among wetlands that was unaffected by interwetland distance. At the regional spatial scale, dispersal events were relatively less frequent between wetland complexes. The complementary analysis of population genetic substructure indicates strong historic gene flow (global F_ST = 0.01). However, a relationship of genetic isolation by distance across the geographical range suggests that dispersal limitation exists at the regional scale. Our approach of contrasting direct and indirect estimates of dispersal at multiple spatial scales in T. coahuila conveys a sustainable evolutionary trajectory of the species pending preservation of threatened wetland habitats and a range‐wide network of corridors.     

Support for the habitat amount hypothesis from a global synthesis of species density studies

Decades of research suggest that species richness depends on spatial characteristics of habitat patches, especially their size and isolation. In contrast, the habitat amount hypothesis predicts that (1) species richness in plots of fixed size (species density) is more strongly and positively related to the amount of habitat around the plot than to patch size or isolation; (2) habitat amount better predicts species density than patch size and isolation combined, (3) there is no effect of habitat fragmentation per se on species density and (4) patch size and isolation effects do not become stronger with declining habitat amount. Data on eight taxonomic groups from 35 studies around the world support these predictions. Conserving species density requires minimising habitat loss, irrespective of the configuration of the patches in which that habitat is contained.     

The scale of genetic differentiation in the Dunes Sagebrush-Lizard (Sceloporus arenicolus), an endemic habitat specialist

The Dunes Sagebrush-Lizard (Sceloporus arenicolus) is a North American species endemic to sand-shinnery oak habitats of the Mescalero and Monahans sand dunes in eastern New Mexico and western Texas. This lizard is listed as Endangered in New Mexico and exhibits habitat specificity at several geographic scales. Dunes Sagebrush-Lizards are only found in topographically complex shinnery oak (Quercus havardii) dominated landscapes within their small geographic distribution and are not found in surrounding human-altered landscapes. Within suitable sand-shinnery oak habitat, individuals predominantly occupy non-vegetated sand dune blowouts and utilize blowouts with particular physical characteristics due to thermoregulatory, reproduction, and foraging requirements. Here, we examined historical and contemporary patterns of genetic differentiation with respect to the current distribution of suitable habitat at multiple spatial scales using mitochondrial DNA sequences and microsatellite data from individuals throughout the entire range. We found three genetic clusters of individuals generally concordant with geographic regions and low sequence divergence at mitochondrial loci suggesting a recent origin of these populations. We also found high levels of genetic structure at microsatellite loci among populations within each of these groups indicating restricted gene flow at intermediate scales. Despite high habitat specificity, we did not detect genetic structure among sand blowouts at finer spatial scales. Within each population, matrices comprised of both sand blowouts and vegetated shinnery oak patches are necessary for genetic connectivity, but the fine scale spatial arrangement of blowouts may not be as critical. We discuss our results with respect to the scale of landscape heterogeneity and habitat connectivity and consider the conservation implications for this threatened taxon.     

Islands,mainland, and terrestrial fragments: How isolation shapes plant diversity

The fragmentation of natural habitats is a major threat for biodiversity. However, the impact and spatial scale of natural isolation mechanisms leading to species loss, compared to anthropogenic fragmentation, are not clear, mainly due to differences between fragments and islands, such as matrix permeability. We studied a 500 km² Mediterranean region in France, including urban habitat fragments, continuous habitat, and continental‐shelf islands. On the basis of 295 floristic relevés, we built species–area relationships to compare isolation in fragments after urbanization, with continuous habitat and continental‐shelf islands. We assumed either no dispersal, infinite dispersal, or estimated intermediate levels of habitat reachability through graph theory. Isolation mechanisms occurred in fragments but with a lower strength than in near‐shore islands, and most importantly affected perennial plants. Annual plants were less affected, probably due to their smaller size and shorter life cycle. Isolation occurred at landscape level in fragments and at patch level in islands. The amount of reachable habitat (accounting for spatial configuration) explained local species richness in both systems, but the amount of habitat (no consideration of spatial configuration) was already a good predictor. These results suggest an important role of habitat amount around fragments in mitigating the isolation effects observed in near‐shore islands, and the importance of carefully considering different functional groups.     

Rampant drift in artificially fragmented populations of the endangered tidewater goby (Eucyclogobius newberryi)

Habitat fragmentation and its genetic consequences are a critically important issue in evaluating the evolutionary penalties of human habitat modification. Here, we examine the genetic structure and diversity in naturally subdivided and artificially fragmented populations of the endangered tidewater goby (Eucyclogobius newberryi), a small fish restricted to discrete coastal lagoons and estuaries in California, USA. We use five naturally fragmented coastal populations from a 300‐ km spatial scale as a standard to assess migration and drift relative to eight artificially fragmented bay populations from a 30‐ km spatial scale. Using nine microsatellite loci in 621 individuals, and a 522‐base fragment of mitochondrial DNA control region from 103 individuals, we found striking differences in the relative influences of migration and drift on genetic variation at these two scales. Overall, the artificially fragmented populations exhibited a consistent pattern of higher genetic differentiation and significantly lower genetic diversity relative to the naturally fragmented populations. Thus, even in a species characterized by habitat isolation and subdivision, further artificial fragmentation appears to result in substantial population genetic consequences and may not be sustainable.     

Rethinking patch size and isolation effects: the habitat amount hypothesis

I challenge (1) the assumption that habitat patches are natural units of measurement for species richness, and (2) the assumption of distinct effects of habitat patch size and isolation on species richness. I propose a simpler view of the relationship between habitat distribution and species richness, the ‘habitat amount hypothesis’, and I suggest ways of testing it. The habitat amount hypothesis posits that, for habitat patches in a matrix of non‐habitat, the patch size effect and the patch isolation effect are driven mainly by a single underlying process, the sample area effect. The hypothesis predicts that species richness in equal‐sized sample sites should increase with the total amount of habitat in the ‘local landscape’ of the sample site, where the local landscape is the area within an appropriate distance of the sample site. It also predicts that species richness in a sample site is independent of the area of the particular patch in which the sample site is located (its ‘local patch’), except insofar as the area of that patch contributes to the amount of habitat in the local landscape of the sample site. The habitat amount hypothesis replaces two predictor variables, patch size and isolation, with a single predictor variable, habitat amount, when species richness is analysed for equal‐sized sample sites rather than for unequal‐sized habitat patches. Studies to test the hypothesis should ensure that ‘habitat’ is correctly defined, and the spatial extent of the local landscape is appropriate, for the species group under consideration. If supported, the habitat amount hypothesis would mean that to predict the relationship between habitat distribution and species richness: (1) distinguishing between patch‐scale and landscape‐scale habitat effects is unnecessary; (2) distinguishing between patch size effects and patch isolation effects is unnecessary; (3) considering habitat configuration independent of habitat amount is unnecessary; and (4) delineating discrete habitat patches is unnecessary.     

Linking genetic and ecological differentiation in an ungulate with a circumpolar distribution

Genetic differentiation among populations may arise from the disruption of gene flow due to local adaptation to distinct environments and/or neutral accumulation of mutations and genetic drift resulted from geographical isolation. Quantifying the role of these processes in determining the genetic structure of natural populations remains challenging. Here, we analyze the relative contribution of isolation‐by‐resistance (IBR), isolation‐by‐environment (IBE), genetic drift and historical isolation in allopatry during Pleistocene glacial cycles on shaping patterns of genetic differentiation in caribou/reindeer populations Rangifer tarandus across the entire distribution range of the species. Our study integrates analyses at range‐wide and regional scales to partial out the effects of historical and contemporary isolation mechanisms. At the circumpolar scale, our results indicate that genetic differentiation is predominantly explained by IBR and historical isolation. At a regional scale, we found that IBR, IBE and population size significantly explained the spatial distribution of genetic variation among populations belonging to the Euro‐Beringian lineage within North America. In contrast, genetic differentiation among populations within the North American lineage was predominantly explained by IBR and population size, but not IBE. We also found discrepancies between genetic and ecotype designation across the Holarctic species distribution range. Overall, these results indicate that multiple isolating mechanisms have played roles in shaping the spatial distribution of genetic variation across the distribution range of a large mammal with high potential for gene flow. Considering multiple spatial scales and simultaneously testing a comprehensive suite of potential isolating mechanisms, our study contributes to understand the ecological and evolutionary processes underlying organism–landscape interactions.     

Assemblage-level responses of phyllostomid bats to tropical forest fragmentation: land-bridge islands as a model system

Aim Working within a system of high structural contrast between fragments and the surrounding matrix, we assessed patterns of species loss and changes in species composition of phyllostomid bats on artificial land‐bridge islands relative to mainland assemblages, and evaluated the responses of bats to forest edges. We further examined the relative influence of local‐scale characteristics (e.g. vegetation structure, island area) versus landscape attributes (e.g. forest cover, patch density) and the importance of spatial scale in determining phyllostomid species richness and composition on islands. Location Islands in Gatún Lake and adjacent mainland peninsulas in the Barro Colorado Nature Monument, Panama. Methods Bats were sampled over a 2‐year period on 11 islands as well as at forest‐edge and interior sites on adjacent mainland, resulting in > 8400 captures. Results The islands harboured a less diverse and structurally simplified phyllostomid bat fauna. Islands far from the mainland were especially species‐poor. This decline in species richness was associated with compositional shifts towards assemblages strongly dominated by frugivores with good dispersal abilities. Members of other ensembles, most importantly gleaning animalivores, were much less common or absent. Although overall species composition was not significantly altered, species richness at continuous forest‐edge sites was significantly lower compared with that at interior sites. Distance from the mainland and amount of forest cover in the landscape were the best predictors of species richness and assemblage composition. Responses were scale‐dependent. At the local scale, species richness was independent of island area but was correlated positively with distance from the mainland. In contrast, area effects became more important at larger spatial scales, suggesting that many species use multiple fragments. Main conclusions Our results underline the conservation value of small habitat remnants, which, even when embedded in a hostile matrix, can support a relatively diverse bat fauna, provided that there is a low degree of patch isolation and spatial proximity to larger tracts of continuous forest. Although the results at the assemblage level were inconclusive, we demonstrate that certain bat species and ensembles, particularly gleaning animalivores, exhibit high edge‐sensitivity. Our results point to habitat loss rather than changes in landscape configuration as the main process after isolation underlying phyllostomid bat responses, suggesting that conservation efforts should focus on habitat preservation instead of trying to minimize fragmentation per se at the expense of habitat amount.     

Variance in composition of inquiline communities in leaves of Sarracenia purpurea L. on multiple spatial scales

A survey of the abundances of species that inhabit the water-bearing leaves of the pitcher plant Sarracenia purpurea was conducted at several different spatial scales in northern Florida. Individual leaves are hosts to communities of inquiline species, including mosquitoes, midges, mites, copepods, cladocerans, and a diverse bacterial assemblage. Inquiline communities were quantified from four pitchers per plant, three plants per subpopulation, two subpopulations per population, and three populations. Species varied in abundance at different spatial scales. Variation in the abundances of mosquitoes and copepods was not significantly associated with any spatial scale. Midges varied in abundance at the level of populations; one population contained significantly more midges than the other two. Cladocerans varied at the level of the subpopulation, whereas mites varied at the level of the individual plants. Bacterial communities were described by means of Biolog plates, which quantify the types of carbon media used by the bacteria in each pitcher. Bacterial communities were found to vary significantly in composition among individual plants but not among populations or subpopulations. These results suggest that independent factors determining the abundances of individual species are important in determining community patterns in pitcher-plant inquilines.     

Habitat fragmentation influences genetic diversity and differentiation: Fine‐scale population structure of Cercis canadensis (eastern redbud)

Forest fragmentation may negatively affect plants through reduced genetic diversity and increased population structure due to habitat isolation, decreased population size, and disturbance of pollen‐seed dispersal mechanisms. However, in the case of tree species, effective pollen‐seed dispersal, mating system, and ecological dynamics may help the species overcome the negative effect of forest fragmentation. A fine‐scale population genetics study can shed light on the postfragmentation genetic diversity and structure of a species. Here, we present the genetic diversity and population structure of Cercis canadensis L. (eastern redbud) wild populations on a fine scale within fragmented areas centered around the borders of Georgia–Tennessee, USA. We hypothesized high genetic diversity among the collections of C. canadensis distributed across smaller geographical ranges. Fifteen microsatellite loci were used to genotype 172 individuals from 18 unmanaged and naturally occurring collection sites. Our results indicated presence of population structure, overall high genetic diversity (H_E = 0.63, H_O = 0.34), and moderate genetic differentiation (F_ST = 0.14) among the collection sites. Two major genetic clusters within the smaller geographical distribution were revealed by STRUCTURE. Our data suggest that native C. canadensis populations in the fragmented area around the Georgia–Tennessee border were able to maintain high levels of genetic diversity, despite the presence of considerable spatial genetic structure. As habitat isolation may negatively affect gene flow of outcrossing species across time, consequences of habitat fragmentation should be regularly monitored for this and other forest species. This study also has important implications for habitat management efforts and future breeding programs.     

SPATIAL SCALE OF GENETIC STRUCTURE AND AN INDIRECT ESTIMATE OF GENE FLOW IN EELGRASS,ZOSTERA MARINA

In this study, the first investigation of population structure in an aquatic angiosperm, I show that populations of a marine angiosperm (eelgrass, Zostera marina) are genetically differentiated at a number of spatial scales. I find also that there is no correspondence between geographic and genetic distances separating subpopulations, an increasingly common result in spatially stratified studies of genetic structure in marine invertebrates. F-statistics, calculated for two years from electrophoretic variation at five polymorphic allozyme loci, indicate significant genetic differentiation among sampling quadrats within each of two bays (θ = 0.064-0.208), between tide zones within a bay (θ = 0.025-0.157) and between bays (θ = 0.079). Spatial autocorrelation analysis was used to explore genetic differentiation at smaller spatial scales; estimated patch sizes (within which genetic individuals are randomly associated) indicated no appeciable genetic structure at scales less than 20 m × 20 m. Calculated values of F-statistics were a function of the spatial scale from which samples were drawn: increasing the size of the “subpopulation” included in calculation of fixation indices for the same “total” sample resulted in an increase in the magnitude of f (e.g., from 0.092 to 0.181) and a decrease in θ (e.g., from 0.186 to 0.025). On the basis of the best estimate of the spatial scale of subpopulations, the effective number of migrants per generation (N_em) ranges from 1.1 to 2.8. Genetic consequences of the disturbance regime in the eelgrass habitat sampled were extreme variation between years in the allele richness and proportion of heterozygotes in a sample and a positive relationship between the extinction probability of patches and the genetic variance among them. The changes in F-statistics as a function of sampling scale and the observation that θ among sampled quadrats was positively associated with the probability of extinction among quadrats indicated that indirect estimates of gene flow (N_em) calculated from θ should be cautiously interpreted in populations that may not yet be in drift-migration equilibrium.     

Ecological metrics predict connectivity better than geographic distance

We use microsatellite loci to examine genetic structure of the Florida scrub lizard (Sceloporus woodi) and test for the effects of landscape variables at the scale of neighboring patches. We evaluate ecological metrics of connectivity with genetics data, which to our knowledge is the first application of these particular metrics to landscape-level genetics studies in Florida scrub. Florida scrub is a highly threatened ecosystem in which habitat patches are remnants of a previously widespread xeric landscape. Analysis of mitochondrial DNA (mtDNA) has shown that landscape structure influenced the evolutionary history of the Florida scrub lizard (S. woodi) across its range. Our results concur with these mtDNA studies in documenting divergence between xeric ridge systems and also demonstrate divergence at very local scales. Both least-cost distance and pairwise isolation (a metric used in ecological studies that includes patch size, quality and a modified isolation index) were better predictors of genetic distance than Euclidean distance, indicating that mesic and hydric habitat influence spatial patterns in genetic variation. Our results support the need for focusing on spatial distribution of scrub habitat at the scale of neighboring patches, as well as regionally, in conservation management and restoration. Also, our study points to the value of integrating landscape ecology metrics into landscape genetics.     

Strong isolation by distance among local populations of an endangered butterfly species (Euphydryas aurinia)

The marsh fritillary (Euphydryas aurinia) is a critically endangered butterfly species in Denmark known to be particularly vulnerable to habitat fragmentation due to its poor dispersal capacity. We identified and genotyped 318 novel SNP loci across 273 individuals obtained from 10 small and fragmented populations in Denmark using a genotyping‐by‐sequencing (GBS) approach to investigate its population genetic structure. Our results showed clear genetic substructuring and highly significant population differentiation based on genetic divergence (F _ST) among the 10 populations. The populations clustered in three overall clusters, and due to further substructuring among these, it was possible to clearly distinguish six clusters in total. We found highly significant deviations from Hardy–Weinberg equilibrium due to heterozygote deficiency within every population investigated, which indicates substructuring and/or inbreeding (due to mating among closely related individuals). The stringent filtering procedure that we have applied to our genotype quality could have overestimated the heterozygote deficiency and the degree of substructuring of our clusters but is allowing relative comparisons of the genetic parameters among clusters. Genetic divergence increased significantly with geographic distance, suggesting limited gene flow at spatial scales comparable to the dispersal distance of individual butterflies and strong isolation by distance. Altogether, our results clearly indicate that the marsh fritillary populations are genetically isolated. Further, our results highlight that the relevant spatial scale for conservation of rare, low mobile species may be smaller than previously anticipated.     

Patch occupancy of two hemipterans sharing a common host plant

Aim Two hemipteran species were chosen as a study system for the comparative analysis of patch occupancy and spatial population structure of insects sharing a common host plant. This study tested whether (1) the incidence in the host plant patches differed between the two species, and (2) the two species exhibited a different spatial population structure, i.e. were they affected differentially by isolation and area of the host plant patches. Location The porphyry landscape north of Halle (Saale) in Germany comprising 506 patches of the host plant Brachypodium pinnatum. Methods The host plant patches were surveyed for the two hemipterans. To assess the influence of patch quality on species occurrence the patches were characterized by mean cover abundance of B. pinnatum, type of subsoil, slope, exposure, and shading. The spatial configuration of the patches was considered by patch area and isolation. The influence of the habitat factors and the spatial configuration on the occupancy of the two species was analysed by logistic regression. Results Adarrus multinotatus was found in 441 patches, while Neophilaenus albipennis was found in only 90 patches. While A. multinotatus showed virtually no relationship to the habitat factors, the occupancy of N. albipennis was influenced by subsoil type, cover abundance, and shading. The effects of area and isolation on occupancy of the patches also differed between the two species. The occupancy of N. albipennis was determined largely by area and isolation, whereas in A. multinotatus no considerable effect of spatial configuration was found. Main conclusions The study revealed a marked difference between the two hemipteran species in respect of spatial population structure. Adarrus multinotatus built up a ‘patchy population’, whereas N. albipennis showed a ‘metapopulation’ structure within the same set of patches in the same landscape. Spatial population structure was found to be not only a function of spatial configuration of habitat patches, but population structure differed between the habitat generalist A. multinotatus and the habitat specialist N. albipennis.     

Spatial Heterogeneity in Habitat Quality and Cross-Scale Interactions in Metapopulations

Abstract Integration of habitat heterogeneity into spatially realistic metapopulation approaches reveals the potential for key cross-scale interactions. Broad-scale environmental gradients and land-use practices can create autocorrelation of habitat quality of suitable patches at intermediate spatial scales. Patch occupancy then depends not only on habitat quality at the patch scale but also on feedbacks from surrounding neighborhoods of autocorrelated patches. Metapopulation dynamics emerge from how demographic and dispersal processes interact with relevant habitat heterogeneity. We provide an empirical example from a metapopulation of round-tailed muskrats (Neofiber alleni) in which habitat quality of suitable patches was spatially autocorrelated most strongly within 1,000 m, which was within the expected dispersal range of the species. After controlling for factors typically considered in metapopulation studies—patch size, local patch quality, patch connectivity—we use a cross-variogram analysis to demonstrate that patch occupancy by muskrats was correlated with habitat quality across scales ≤1,171 m. We also discuss general consequences of spatial heterogeneity of habitat quality for metapopulations related to potential cross-scale interactions. We focus on spatially correlated extinctions and metapopulation persistence, hierarchical scaling of source–sink dynamics, and dispersal decisions by individuals in relation to information constraints.     

Niche specificity influences gene flow across fine‐scale habitat mosaics in Succulent Karoo plants

While the tempo of diversification in biodiversity hotspots has received much attention, the spatial scale of diversification has often been overlooked. Addressing this deficiency requires understanding the drivers of population divergence and the spatial scales at which they operate in species‐rich clades and ecosystems. South Africa's Succulent Karoo (SK) hotspot provides an excellent system for such research, being both compact (ca. 110,000 km²) and home to spectacular in‐situ radiations, such as the ruschioid Aizoaceae. Here we use GBS to document genetic structure in two co‐occurring ruschioid species, at both coarse (>10 km) and fine (<500 m) spatial scales. Where Ruschia burtoniae shows strong between‐population genetic differentiation and no gene flow, Conophytum calculus shows weak differentiation, with high levels of admixture suggesting recent or ongoing gene flow. Community analysis and transplant experiments reveal that R. burtoniae occupies a narrow, low‐pH edaphic niche, and at scales of a few hundred metres, areas of elevated genetic turnover correspond to patches of edaphically unsuitable habitat. In contrast, C. calculus occupies a broader niche and exhibits isolation‐by‐distance without a habitat effect. We suggest that edaphic specialisation, coupled with highly restricted seed and pollen dispersal in heterogeneous landscapes, has played a major role in driving rapid diversification at small spatial scales in this system. However, the contrasting patterns in our study species show that these factors do not influence all organisms uniformly, being strongly modulated by lineage‐specific traits that influence both the spatial scale of gene flow and habitat specificity.     

Evidence of spatial genetic structure in a California bunchgrass population

We investigated the scale of genetic variation of purple needlegrass (Nassella pulchra), a species commonly used in California for grassland restoration. Common garden and field data revealed evidence of genetic differentiation between two intermixed microhabitats characterized by differences in soil depth and community composition. We assessed the genetic variation within a single population using randomly amplified polymorphic DNA (RAPD) data collected from clusters of five individuals in 40 locations. We found no evidence for genetic structure at the whole population level. At smaller spatial scales, however, we found strong evidence that genetic subdivision of the population occurs at the level of the maternal neighborhood. We suggest that the interaction between widespread pollen dispersal and restricted seed dispersal may be the primary factor generating these results; panmictic pollen dispersal will make detection of genetic patterning difficult at larger spatial scales while limited seed dispersal will generate local genetic structure. As a result, the detection of population genetic structure will depend on the spatial scale of analysis. Local selection gradients related to topography and soil depth are also likely to play a role in structuring local genetic variation. Since N. pulchra is widely used in California in grassland and woodland habitat restoration, we suggest that, as a general rule, care should be exercised in transferring germplasm for the purposes of conservation when little is known about the within-population genetic subdivision of a plant species. Received: 23 December 1996 / Accepted: 20 May 1997