Percolation theory suggests some general features in range margins across environmental gradients

The margins within the geographic range of species are often specific in terms of ecological and evolutionary processes, and can strongly influence the species' reaction to climate change. One of the frequently observed features at range margins is fragmentation, caused internally by population dynamics or externally by the limited availability of suitable habitat sites. We study both causes, and describe the transition from a connected to a fragmented state across space by means of a gradient metapopulation model. The main features of our approach are the following. 1) Inhomogeneities can occur at two spatial scales: there is a broad-scale gradient, which can be patterned by fine-scale heterogeneities. The latter is implemented by dispersing a variable number of small obstacles over the terrain, which can be penetrable or unpenetrable by the spreading species. 2) We study the occupancy of this terrain in a steady-state on two temporal scales: in snapshots and by long-term averages. The simulations reveal some general scaling laws that are applicable in various environments, independently of the mechanism of fragmentation. The edge of the connected region (the hull) is a fractal with dimension 7/4. Its width and length changes with the gradient according to universal scaling laws, that are characteristic for percolation transitions. The results suggest that percolation theory is a powerful tool for understanding the structure of range margins in a broad variety of real-life scenarios, including those in which the environmental gradient is combined with fine-scale heterogeneity. This provides a new method for comparing the range margins of different species in various geographic regions, and monitoring range shifts under climate change.Comment: 17 pages, 5 figure

Dynamics of Population on the Verge of Extinction

Theoretical considerations suggest that extinction in dispersal-limited populations is necessarily a threshold-like process that is analogous to a critical phase transition in physics. We use this analogy to find robust, common features in the dynamics of extinctions, and suggest early warning signals which may indicate that a population is endangered. As the critical threshold of extinction is approached, the population spontaneously fragments into discrete subpopulations and, consequently, density regulation fails. The population size declines and its spatial variance diverges according to scaling laws. Therefore, we can make robust predictions exactly in the range where prognosis is vital, on the verge of extinction

Intermediate landscape disturbance maximizes metapopulation density

The viability of metapopulations in fragmented landscapes has become a central theme in conservation biology. Landscape fragmentation is increasingly recognized as a dynamical process: in many situations, the quality of local habitats must be expected to undergo continual changes. Here we assess the implications of such recurrent local disturbances for the equilibrium density of metapopulations. Using a spatially explicit lattice model in which the considered metapopulation as well as the underlying landscape pattern change dynamically, we show that equilibrium metapopulation density is maximized at intermediate frequencies of local landscape disturbance. On both sides around this maximum, the metapopulation may go extinct. We show how the position and shape of the intermediate viability maximum is responding to changes in the landscape's overall habitat quality and the population's propensity for local extinction. We interpret our findings in terms of a dual effect of intensified landscape disturbances, which on the one hand exterminate local population and on the other hand enhance a metapopulation's capacity for spreading between habitat clusters

Cooperation and competition in heterogeneous environments: The evolution of resource sharing in clonal plants

Plant species show great variation in the degree of physiological integration between developmental units (modules). When this physiological integration is minimal, individual modules are self-supporting and compete with other modules. When there is greater physiological integration, modules remain physiologically connected and 'cooperate' by sharing resources like water, nutrients and photoassimilates taken up from their local environments. Thus, local differences in habitat quality can be diminished within a group of modules. Here we examine how the evolutionarily optimal amount of integration depends on habitat type - with habitats being characterized by the proportion of resource-rich and resource-poor sites and by the turnover rate between them. Two main questions are addressed: First, how does spatial heterogeneity influence natural selection for or against integration? Second, can adaptation, under reasonable ecological conditions, stabilize partial integration? A non-spatial version of the model, which assumes well-mixed populations, predicts the complete physiological independence of modules as the only evolutionarily stable outcome in any realistic habitat type. By contrast, a spatially explicit version of the model reveals the adaptive advantage of integration in typical high-risk habitats, where resource-rich sites are sparsely distributed in space and transient in time. We conclude that habitat diversity without spatial population structure is sufficient to explain the evolutionary loss of physiological integration. But only the additional consideration of spatial population structure can convincingly explain any backward transition and the stable existence of partial integration

Invasion impact is conditioned by initial vegetation states

Abstract Biological invasion is a crucial problem in the world because of its negative consequences for protected areas. The degradation stage of vegetation might affect the success of invasion. One of the most abundant and threatening invasive species is the common milkweed (Asclepias syriaca L.) which has invaded already 23 countries of Europe and in several habitat types its further spreading is promoted by climate change. Pannonian sand grassland is one of the most threatened habitat by common milkweed invasion. Therefore, invasion in sand grassland vegetation is an important issue. However, the effects of the invasive plant in the open sand grassland are rather controversial. In order to clarify the existing contradictory results, the study was carried out in a strictly protected area, near Fülöpháza (Hungary) in a reserve core area in a UNESCO biosphere reserve. Microcoenological study was applied to determinate the fine-scale community characteristics of non-invaded and invaded stands in natural and seminatural vegetation and data were processed by Juhász-Nagy's information theory models. Shannon diversity of species combinations (compositional diversity) which describes the ways of the coexistence of species, and the number of realized species combinations were used for measuring beta diversity. Differences between stands were analyzed by two-way ANOVA. The maximum compositional diversity of species and main life-forms (annuals, perennials and cryptogams) did not differ significantly between the non-invaded and invaded stands. In contrast, significantly larger characteristic areas of compositional diversity were detected in the invaded stands. Based on these results, it could be concluded that diversity of species combinations did not change but those values have shifted to coarser scales in case of invaded stands. The direction of this change suggests a kind of impoverishment in the presence of Asclepias. Thus, it is worth mentioning from the invasion management point of view that protection of the habitats against disturbance is a more cost-effective and successful way than protection against the establishment or extirpation of invasive species, since disturbance facilitates the invasions throughout the impoverishment of the community

Ecological Invasion, Roughened Fronts, and a Competitor's Extreme Advance: Integrating Stochastic Spatial-Growth Models

Both community ecology and conservation biology seek further understanding of factors governing the advance of an invasive species. We model biological invasion as an individual-based, stochastic process on a two-dimensional landscape. An ecologically superior invader and a resident species compete for space preemptively. Our general model includes the basic contact process and a variant of the Eden model as special cases. We employ the concept of a "roughened" front to quantify effects of discreteness and stochasticity on invasion; we emphasize the probability distribution of the front-runner's relative position. That is, we analyze the location of the most advanced invader as the extreme deviation about the front's mean position. We find that a class of models with different assumptions about neighborhood interactions exhibit universal characteristics. That is, key features of the invasion dynamics span a class of models, independently of locally detailed demographic rules. Our results integrate theories of invasive spatial growth and generate novel hypotheses linking habitat or landscape size (length of the invading front) to invasion velocity, and to the relative position of the most advanced invader.Comment: The original publication is available at www.springerlink.com/content/8528v8563r7u2742

Differential Influence of Clonal Integration on Morphological and Growth Responses to Light in Two Invasive Herbs

Background and aims: In contrast to seeds, high sensitivity of vegetative fragments to unfavourable environments may limit the expansion of clonal invasive plants. However, clonal integration promotes the establishment of propagules in less suitable habitats and may facilitate the expansion of clonal invaders into intact native communities. Here, we examine the influence of clonal integration on the morphology and growth of ramets in two invasive plants, Alternanthera philoxeroides and Phyla canescens, under varying light conditions. Methods: In a greenhouse experiment, branches, connected ramets and severed ramets of the same mother plant were exposed under full sun and 85 % shade and their morphological and growth responses were assessed. Key results: The influence of clonal integration on the light reaction norm (connection6light interaction) of daughter ramets was species-specific. For A. philoxeroides, clonal integration evened out the light response (total biomass, leaf mass per area, and stem number, diameter and length) displayed in severed ramets, but these connection6light interactions were largely absent for P. canescens. Nevertheless, for both species, clonal integration overwhelmed light effect in promoting the growth of juvenile ramets during early development. Also, vertical growth, as an apparent shade acclimation response, was more prevalent in severed ramets than in connected ramets. Finally, unrooted branches displayed smaller organ size and slower growth than connected ramets, but the pattern of light reaction was similar, suggesting mothe

How Past and Present Influence the Foraging of Clonal Plants?

Clonal plants spreading horizontally and forming a network structure of ramets exhibit complex growth patterns to maximize resource uptake from the environment. They respond to spatial heterogeneity by changing their internode length or branching frequency. Ramets definitively root in the soil but stay interconnected for a varying period of time thus allowing an exchange of spatial and temporal information. We quantified the foraging response of clonal plants depending on the local soil quality sampled by the rooting ramet (i.e. the present information) and the resource variability sampled by the older ramets (i.e. the past information). We demonstrated that two related species, Potentilla reptans and P. anserina, responded similarly to the local quality of their environment by decreasing their internode length in response to nutrient-rich soil. Only P. reptans responded to resource variability by decreasing its internode length. In both species, the experience acquired by older ramets influenced the plastic response of new rooted ramets: the internode length between ramets depended not only on the soil quality locally sampled but also on the soil quality previously sampled by older ramets. We quantified the effect of the information perceived at different time and space on the foraging behavior of clonal plants by showing a non-linear response of the ramet rooting in the soil of a given quality. These data suggest that the decision to grow a stolon or to root a ramet at a given distance from the older ramet results from the integration of the past and present information about the richness and the variability of the environment