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Phylogeographic structure of the fire ant Solenopsis invicta in its native South American range: roles of natural barriers and habitat connectivity.

We generated mitochondrial DNA (mtDNA) sequence data from 402 individuals of the fire ant Solenopsis invicta collected from 11 native populations and analyzed these data using a combination of demographic, phylogenetic, and phylogeographic methods to infer features of the evolutionary history of this species. Prior expectations regarding high levels of genetic structure and isolation by distance among populations were supported by the data, but we also discovered several unanticipated patterns. Our analyses revealed a major genetic break between S. invicta mtDNA haplotypes that coincides with the Mesopotamia wetlands region of South America, resulting in two higher level nested clade groupings. In addition, we identified contrasting patterns of genetic differentiation within these two major groups, which may reflect differences in connectivity of suitable habitat in different parts of the native range of S. invicta. Our study represents the first attempt to understand the phylogeographic history of S. invicta across its native range.

Animals↗

The colonization of Europe by the freshwater crustacean Asellus aquaticus (Crustacea: Isopoda) proceeded from ancient refugia and was directed by habitat connectivity.

Recent continental-scale phylogeographic studies have demonstrated that not all freshwater fauna colonized Europe from the classic Mediterranean peninsular refugia, and that northern or central parts of the continent were occupied before, and remained inhabited throughout the Pleistocene. The colonization history of the ubiquitous aquatic isopod crustacean Asellus aquaticus was assessed using mitochondrial COI and a variable part of nuclear 28S rDNA sequences. Phylogeographic analysis of the former suggested that dispersion proceeded possibly during late Miocene from the western part of the Pannonian basin. Several areas colonized from here have served as secondary refugia and/or origins of dispersion, well before the beginning of the Pleistocene. Postglacial large-scale range expansion was coupled with numerous separate local dispersions from different refugial areas. Connectivity of the freshwater habitat has played an important role in shaping the current distribution of genetic diversity, which was highest in large rivers. The importance of hydrographic connections for the maintenance of genetic contact was underscored by a discordant pattern of mtDNA and nuclear rDNA differentiation. Individuals from all over Europe, differing in their mtDNA to a level normally found between species or even genera (maximal within population nucleotide divergence reached 0.16 +/- 0.018), shared the same 28S rRNA gene sequence. Only populations from hydrographically isolated karst water systems in the northwestern Dinaric Karst had distinct 28S sequences. Here isolation seemed to be strong enough to prevent homogenization of the rRNA gene family, whereas across the rest of Europe genetic contact was sufficient for concerted evolution to act.

Animals↗

Habitat connectivity and ecosystem productivity: implications from a simple model.

The import of resources (food, nutrients) sustains biological production and food webs in resource-limited habitats. Resource export from donor habitats subsidizes production in recipient habitats, but the ecosystem-scale consequences of resource translocation are generally unknown. Here, I use a nutrient-phytoplankton-zooplankton model to show how dispersive connectivity between a shallow autotrophic habitat and a deep heterotrophic pelagic habitat can amplify overall system production in metazoan food webs. This result derives from the finite capacity of suspension feeders to capture and assimilate food particles: excess primary production in closed autotrophic habitats cannot be assimilated by consumers; however, if excess phytoplankton production is exported to food-limited heterotrophic habitats, it can be assimilated by zooplankton to support additional secondary production. Transport of regenerated nutrients from heterotrophic to autotrophic habitats sustains higher system primary production. These simulation results imply that the ecosystem-scale efficiency of nutrient transformation into metazoan biomass can be constrained by the rate of resource exchange across habitats and that it is optimized when the transport rate matches the growth rate of primary producers. Slower transport (i.e., reduced connectivity) leads to nutrient limitation of primary production in autotrophic habitats and food limitation of secondary production in heterotrophic habitats. Habitat fragmentation can therefore impose energetic constraints on the carrying capacity of aquatic ecosystems. The outcomes of ecosystem restoration through habitat creation will be determined by both functions provided by newly created aquatic habitats and the rates of hydraulic connectivity between them.

Animals↗

Connectivity, habitat heterogeneity, and population persistence in Ranunculus nodiflorus, an endangered species in France.

Here, we explore the role of habitat spatial structure in the maintenance of metapopulations of Ranunculus nodiflorus. This rare species grows in puddles that can be connected occasionally by flooded corridors. We monitored five locations in the Fontainebleau forest, France, since 2002 and recorded the presence of corridors among puddles and evaluated their impact on puddle demography and plant fitness. We showed that connections increased population size, by increasing both the number of puddles occupied by the species and the density of individuals within puddles, but seemed to have no direct influence on plant fitness. We found no evidence of a large persistent soil seed bank. Natural corridors are likely to decrease the extinction probability of the populations, most probably by allowing recolonization of empty puddles after extinctions. Therefore, the preservation of corridors appears crucial for the conservation of R. nodiflorus in its natural habitat.

Ecosystem↗

Effects of historical climate change, habitat connectivity, and vicariance on genetic structure and diversity across the range of the red tree vole (Phenacomys longicaudus) in the Pacific Northwestern United States.

Phylogeographical analyses conducted in the Pacific Northwestern United States have often revealed concordant patterns of genetic diversity among taxa. These studies demonstrate distinct North/South genetic discontinuities that have been attributed to Pleistocene glaciation. We examined phylogeographical patterns of red tree voles (Phenacomys longicaudus) in western Oregon by analysing mitochondrial control region sequences for 169 individuals from 18 areas across the species' range. Cytochrome b sequences were also analysed from a subset of our samples to confirm the presence of major haplotype groups. Phylogenetic network analyses suggested the presence of two haplotype groups corresponding to northern and southern regions of P. longicaudus' range. Spatial genetic analyses (samova and Genetic Landscape Shapes) of control region sequences demonstrated a primary genetic discontinuity separating northern and southern sampling areas, while a secondary discontinuity separated northern sampling areas into eastern and western groups divided by the Willamette Valley. The North/South discontinuity likely corresponds to a region of secondary contact between lineages rather than an overt barrier. Although the Cordilleran ice sheet (maximum approximately 12,000 years ago) did not move southward to directly affect the region occupied by P. longicaudus, climate change during glaciation fragmented the forest landscape that it inhabits. Signatures of historical fragmentation were reflected by positive associations between latitude and variables such as Tajima's D and patterns associated with location-specific alleles. Genetic distances between southern sampling areas were smaller, suggesting that forest fragmentation was reduced in southern vs. northern regions.

Animals↗

Loss of plant species richness and habitat connectivity in grasslands associated with agricultural change in Finland.

The drastic loss of seminatural grasslands and the decrease in species diversity in Europe during the 20th century are closely linked to social-economic factors. Development in agricultural production drives land-use changes, and thus controls the capacity of landscapes to maintain biodiversity. In this study, we link agricultural production changes to landscape fragmentation and species diversity. Our results show that the termination of grazing on seminatural grassland caused significant changes in landscape structure and a decline in the number of vascular plant species. The decline of grazed grasslands has been driven mainly by farm-level economic efficiency and profitability interests, which have been connected with agricultural policy measures. Since 1995, when Finland joined the European Union, the area of grazed patches in our study area has again increased as a result of a support scheme for the management of seminatural grasslands.

Agriculture↗

A spatial model to estimate habitat fragmentation and its consequences on long-term persistence of animal populations.

The increasing use of the landscape by humans has led to important diminutions of natural surfaces. The remaining patches of wild habitat are small and isolated from each other among a matrix of inhospitable land-uses. This habitat fragmentation, by disabling population movements and stopping their spread to new habitats, is a major threat to the survival of numerous plant and animal species. We developed a general model, adaptable for specific species, capable of identifying suitable habitat patches within fragmented landscapes and investigating the capacity of populations to move between these patches. This approach combines GIS analysis of a landscape, with spatial dynamic modeling. Suitable habitat is identified using a threshold area to perimeter ratio. Potential movement pathways of species between habitat patches are modeled using a cellular automaton. Habitat connectivity is estimated by overlaying habitat patches with movement pathways. The maximum potential population is calculated within and between connected habitat patches and potential risk of inbreeding within meta-populations is considered. The model was tested on a sample map and applied to scenario maps of predicted land-use change in the Peoria Tri-county region (IL). It (1) showed area of natural area alone was insufficient to estimate the consequences on animal populations; (2) underscored the necessity to use approaches investigating the effect of land-use change spatially through the landscape and the importance of considering species-specific life history characteristics; and (3) highlighted the model's potential utility as an indicator of species likelihood to be affected negatively by land-use scenarios and therefore requiring detailed investigation.

Animals↗

Two-dimensional gap analysis: a tool for efficient conservation planning and biodiversity policy implementation.

The maintenance of biodiversity by securing representative and well-connected habitat networks in managed landscapes requires a wise combination of protection, management, and restoration of habitats at several scales. We suggest that the integration of natural and social sciences in the form of "Two-dimensional gap analysis" is an efficient tool for the implementation of biodiversity policies. The tool links biologically relevant "horizontal" ecological issues with "vertical" issues related to institutions and other societal issues. Using forest biodiversity as an example, we illustrate how one can combine ecological and institutional aspects of biodiversity conservation, thus facilitating environmentally sustainable regional development. In particular, we use regional gap analysis for identification of focal forest types, habitat modelling for ascertaining the functional connectivity of "green infrastructures", as tools for the horizontal gap analysis. For the vertical dimension we suggest how the social sciences can be used for assessing the success in the implementation of biodiversity policies in real landscapes by identifying institutional obstacles while implementing policies. We argue that this interdisciplinary approach could be applied in a whole range of other environments including other terrestrial biota and aquatic ecosystems where functional habitat connectivity, nonlinear response to habitat loss and a multitude of economic and social interests co-occur in the same landscape.

Conservation of Natural Resources↗

Genetic differentiation and gene flow among populations of the alpine butterfly, Parnassius smintheus, vary with landscape connectivity.

Levels of gene flow among populations vary both inter- and intraspecifically, and understanding the ecological bases of variation in levels of gene flow represents an important link between the ecological and evolutionary dynamics of populations. The effects of habitat spatial structure on gene flow have received considerable attention; however, most studies have been conducted at a single spatial scale and without background data on how individual movement is affected by landscape features. We examined the influence of habitat connectivity on inferred levels of gene flow in a high-altitude, meadow-dwelling butterfly, Parnassius smintheus. For this species, we had background data on the effects of landscape structure on both individual movement and on small-scale population genetic differentiation. We compared genetic differentiation and patterns of isolation by distance, based on variation at seven microsatellite loci, among three regions representing two levels of connectivity of high-altitude, nonforested habitats. We found that reduced connectivity of habitats, resulting from more forest cover at high altitudes, was associated with greater genetic differentiation among populations (higher estimated FST), a breakdown of isolation by distance, and overall lower levels of inferred gene flow. These observed differences were consistent with expectations based on our knowledge of the movement behaviour of this species and on previous population genetic analyses conducted at the smaller spatial scale. Our results indicate that the role of gene flow may vary among groups of populations depending on the interplay between individual movement and the structure of the surrounding landscape.

Animals↗

Habitat barriers limit gene flow and illuminate historical events in a wide-ranging carnivore, the American puma.

We examined the effects of habitat discontinuities on gene flow among puma (Puma concolor) populations across the southwestern USA. Using 16 microsatellite loci, we genotyped 540 pumas sampled throughout the states of Utah, Colorado, Arizona, and New Mexico, where a high degree of habitat heterogeneity provides for a wide range of connective habitat configurations between subpopulations. We investigated genetic structuring using complementary individual- and population-based analyses, the latter employing a novel technique to geographically cluster individuals without introducing investigator bias. The analyses revealed genetic structuring at two distinct scales. First, strikingly strong differentiation between northern and southern regions within the study area suggests little migration between them. Second, within each region, gene flow appears to be strongly limited by distance, particularly in the presence of habitat barriers such as open desert and grasslands. Northern pumas showed both reduced genetic diversity and greater divergence from a hypothetical ancestral population based on Bayesian clustering analyses, possibly reflecting a post-Pleistocene range expansion. Bayesian clustering results were sensitive to sampling density, which may complicate inference of numbers of populations when using this method. The results presented here build on those of previous studies, and begin to complete a picture of how different habitat types facilitate or impede gene flow among puma populations.

Animals↗

The maintenance (or not) of polygenic variation by soft selection in heterogeneous environments.

On the basis of single-locus models, spatial heterogeneity of the environment coupled with strong population regulation within each habitat (soft selection) is considered an important mechanism maintaining genetic variation. We studied the capacity of soft selection to maintain polygenic variation for a trait determined by several additive loci, selected in opposite directions in two habitats connected by dispersal. We found three main types of stable equilibria. Extreme equilibria are characterized by extreme specialization to one habitat and loss of polymorphism. They are analogous to monomorphic equilibria in singe-locus models and are favored by similar factors: high dispersal, weak selection, and low marginal average fitness of intermediate genotypes. At the remaining two types of equilibria the population mean is intermediate but variance is very different. At fully polymorphic equilibria all loci are polymorphic, whereas at low-variance equilibria at most one locus remains polymorphic. For most parameters only one type of equilibrium is stable; the transition between the domains of fully polymorphic and low-variance equilibria is typically sharp. Low-variance equilibria are favored by high marginal average fitness of intermediate genotypes, in contrast to single-locus models, in which marginal overdominance is particularly favorable for maintenance of polymorphism. The capacity of soft selection to maintain polygenic variation is thus more limited than extrapolation from single-locus models would suggest, in particular if dispersal is high and selection weak. This is because in a polygenic model, variance can evolve independently of the mean, whereas in the single-locus two-allele case, selection for an intermediate mean automatically leads to maintenance of polymorphism.

Animals↗

How corridors reduce indigo bunting nest success.

Corridors are a popular strategy to conserve biodiversity and promote gene flow in fragmented landscapes. Corridor effectiveness has been bolstered by the fact that no empirical field studies have shown negative effects on populations or communities. I tested the hypothesis that corridors increase nest predation in connected habitat fragments relative to unconnected fragments. I evaluated this hypothesis in a large-scale experimental system of open-habitatfragments that varied in shape and connectivity. Corridors increased nest predation rates in connected fragments relative to unconnected fragments with lower edge:area ratios. Nest predation rates were similar between connected and unconnected fragments with higher edge:area ratios. These results suggest that the increase in predator activity is largely attributable to edge effects incurred through the addition of a corridor This is the first field study to demonstrate that corridors can negatively impact animal populations occupying connected fragments.

Animals↗

Testing simple indices of habitat proximity.

Simple measures of habitat proximity made primarily on the basis of land cover are widely used in the ecological literature to infer habitat connectivity, or the potential for animal movement among resource patches. However, such indices rarely have been tested against observations of animal movement or against more detailed biological models. We developed a priori expectations as to the types of study systems and organisms for which various habitat proximity indices would be best suited. We then used data from three study systems and four species to test which, if any, of the indices were good predictors of population-level responses. Our a priori expectations about index performance were not upheld. The indices that consider both habitat area and distance from the focal patch were highly correlated with each other, suggesting that they do index similar quantities. However, none of the indices performed well in predicting population response variables. The results suggest that the pattern of habitat cover alone may be insufficient to predict the process of animal movement.

Animal Migration↗

Criteria to assess and select sites for long-term avian monitoring in an urbanizing landscape.

A methodology was developed to prioritize the suitability of sites for long-term monitoring of avian populations, including vulnerable species, both to enhance assessment of changes in ecological resources and to facilitate land-use planning at the regional scale. This paper argues that a successful monitoring program begins with a site prioritization procedure that integrates scores based on spatial controls with ecological and socio-economic indicators, particularly those dependent on community involvement. The evaluation strategy in this study combines 1) spatial controls such as land ownership and accessibility, with 2) biological and habitat indicators such as vulnerable species and habitat connectivity, and 3) community and agency variables such as volunteer commitment and agency priorities. In total, a set of ten indicators was identified. This strategy was applied to predominantly agricultural landscapes, which are experiencing increasing human pressures, in three sub-watersheds of the Credit River, Southern Ontario. Specifically, bird populations were recorded during the breeding seasons of 2000-2002 in nine land units or habitat types including marsh, deciduous forest, and grasslands as mapped by Credit Valley Conservation (CVC) following Ecological Land Classification (ELC) guidelines. CVC selected sites for long-term monitoring in 2002 and the relationships between the scored (or ranked) sites and the selected long-term monitoring sites are discussed.

Agriculture↗

Environmentally induced dispersal under heterogeneous logistic growth.

We consider a single-species model which is composed of several habitats connected by linear migration rates and having logistic growth. A spatially varying, temporally constant environment is introduced by the non-homogeneity of its carrying capacity. Under this condition any type of purely diffusive behavior, characterized in our model by symmetric migration rates, produces an unbalanced population distribution, i.e. some locations receive more individuals than can be supported by the environmental carrying capacity, while others receive less. Using an evolutionarily stable strategy (ESS) approach we show that an asymmetric migration mechanism, induced by the heterogeneous carrying capacity of the environment, will be selected. This strategy balances the inflow and outflow of individuals in each habitat (balanced dispersal), as well as 'balancing' the spatial distribution relative to variation in carrying capacity (the Ideal Free Distribution from habitat selection theory). We show that several quantities are maximized or minimized by the evolutionarily stable dispersal strategy.

Animal Migration↗

Spatially-explicit matrix models. A mathematical analysis of stage-structured integrodifference equations.

This paper is concerned with mathematical analysis of the 'critical domain-size' problem for structured populations. Space is introduced explicitly into matrix models for stage-structured populations. Movement of individuals is described by means of a dispersal kernel. The mathematical analysis investigates conditions for existence, stability and uniqueness of equilibrium solutions as well as some bifurcation behaviors. These mathematical results are linked to species persistence or extinction in connected habitats of different sizes or fragmented habitats; hence the framework is given for application of such models to ecology. Several approximations which reduce the complexity of integrodifference equations are given. A simple example is worked out to illustrate the analytical results and to compare the behavior of the integrodifference model to that of the approximations.

Algorithms↗

Flower visitors and pollination in the Oriental (Indomalayan) Region.

Current knowledge of flower visitors and pollination in the Oriental Region is summarised. Much less is known about pollination than seed dispersal and the coverage of habitats and taxa in the region is very uneven. The available evidence suggests that pollination in lowland forests is dominated by highly social bees (mainly Trigona and Apis species), with beetles probably the next most important group, followed by other bees and flies. In comparison with the better-studied Neotropics, large solitary bees, moths, Lepidoptera and vertebrates are relatively less important. These differences are greatest in the canopy of the lowland dipterocarp forests of Southeast Asia, where they probably reflect the unique temporal pattern of floral resource availability resulting from 'general flowering' at supra-annual intervals. Apis bees (but not Trigona species) are also important in most montane, subtropical and non-forest habitats. Apart from the figs (Ficus spp.), there are few well-documented examples of plant species visited by a single potential pollinator and most plant-pollinator relationships in the region appear to be relatively generalised. The small sizes of most pollinators and the absence of direct human exploitation probably make pollination mutualisms less vulnerable to failure as a result of human impacts than dispersal mutualisms, but more subtle impacts, as a result of altered gene flows, are likely to be widespread. On current evidence, pollination systems in the Oriental Region do not require any specific conservation action, but this review reinforces arguments for making the preservation (or restoration) of habitat connectivity the major focus of Oriental conservation.

Animals↗

Habitat history improves prediction of biodiversity in rainforest fauna.

Patterns of biological diversity should be interpreted in light of both contemporary and historical influences; however, to date, most attempts to explain diversity patterns have largely ignored history or have been unable to quantify the influence of historical processes. The historical effects on patterns of diversity have been hypothesized to be most important for taxonomic groups with poor dispersal abilities. We quantified the relative stability of rainforests over the late Quaternary period by modeling rainforest expansion and contraction in 21 biogeographic subregions in northeast Australia across four time periods. We demonstrate that historical habitat stability can be as important, and in endemic low-dispersal taxa even more important, than current habitat area in explaining spatial patterns of species richness. In contrast, patterns of endemic species richness for taxa with high dispersal capacity are best predicted by using current environmental parameters. We also show that contemporary patterns of species turnover across the region are best explained by historical patterns of habitat connectivity. These results clearly demonstrate that spatially explicit analyses of the historical processes of persistence and colonization are both effective and necessary for understanding observed patterns of biodiversity.

Animals↗