Landscape effects on anuran pond occupancy in an agricultural countryside: barrier-based buffers predict distributions better than circular buffers

Species movement and occupancy of habitat patches are dependent on landscape permeability. Some land-use types (e.g., roads) may be barriers to animal movement. Analyses of the effect of landscape structure on patch occupancy usually use circular buffers around focal patches. The main assumption of this methodological approach is that species are affected by a particular landscape element equally in every direction from a given patch. This assumption is likely not to hold if animal movement is restricted by barriers because barriers reduce movement patterns and reshape the ideal circular buffer into a noncircular buffer. We developed a method to determine the effect of landscape variables oil the distribution of two amphibian species that explicitly takes dispersal barriers into account. We extracted landscape variables within (i) circular buffers (CB) and (ii) barrier-based buffers (BBB). BBB were produced by reducing the boundaries of CB according to major impassable barriers. The BBB approach almost doubled the explained deviance of multiple regression models in comparison with the CB approach. Moreover. CB and BBB models included different predictor variables. We Suggest that the BBB approach is more Useful than the traditional CB analyses of species-habitat relationships because ecological barriers are explicitly taken into account..

Dispersal modelling

Séverine Vuilleumier Varisco

In human-dominated landscapes, populations and species extinctions are directly related to habitat destruction and fragmentation. To provide genetic diversity as well as population viability, individual exchanges among isolated populations must be maintained. Therefore, animal dispersal processes in fragmented landscape become an important topic for ecologists, and ecological networks planning has become one of the major challenges for landscape planners. Identification of habitat patches as well as assessment of the effect of ecological networks is badly needed. Since little information on the effect of landscape heterogeneities on animal dispersal is available, simulation models are being developed. As dispersal pattern and success strongly depend on the spatial context, species' interactions with landscapes, species behaviour and species ability to disperse, these models must be able to simulate them explicitly. This research work therefore aims first at developing methods and models that allow realistic animal dispersal simulations in fragmented landscapes. Second it aims at evaluating the effect of landscape heterogeneities and animal behaviour on dispersal and on species persistence. Additionally, the ability of such a model to estimate gene flow is analysed. To carry on this research, the following fields have been explored: landscape ecology, metapopulation dynamic, animal behaviour, genetics, Geographical Information Systems, modelling approaches and programming. A method, based on properties provided by Geographical Information Systems software, is first proposed to generate ecological networks by simulating animal dispersal according to animal movement constraints induced by human infrastructures. The resulting maps provide a spatial identification of ecological networks, corridors and conflicting areas. This model has proved to be a useful and straightforward tool for landscape planning, even if this model, similar to other present-day models used in dispersal simulations, presents numerous technical and scientific limitations. To improve models for animal dispersal, a feature-oriented landscape model associated with an expert system has been developed. Its conceptualisation, its formalism, its data structure and its object-oriented design implementation provide a very accurate representation of landscape features and simulation of complex interactions between model entities (individuals and landscape features) based on simple rules. It allows the spatial identification of simulated processes. The ability of this model to incorporate states, relations and transition rules between entities makes it applicable to simulate large ranges of dispersal processes according to specific behaviour and/or landscape uses. To analyse the influence of landscape heterogeneities and species behaviour on dispersal and their incidence on metapopulation dynamics, the proposed feature-oriented model has been coupled with an animal model. The latter assigns different cognitive and dispersal abilities to individuals. Based on simulations according to three movement strategies (corresponding to the cognitive abilities of the simulated species), two measures evaluate the effect of cognitive abilities on dispersal: the colonisation probability between habitat patches and the ecological distance (due to landscape heterogeneities). These measures give an estimation of metapopulation structures (the habitat patches belonging to the metapopulation) and metapopulation dynamics induced by the landscape heterogeneities (for example, the habitat patches which release individuals). The complexity of dispersal processes, considering species behaviours and dispersal abilities, can therefore be reproduced and analysed at different levels. This application has shown the importance of animal behaviour on metapopulation dynamics and structure. Since tracking animals and providing sufficient data remain difficult, calibration and validation procedures of dispersal models are difficult to perform. One approach proposed here is to measure one of the consequences of dispersal: genetic differentiation among populations. Geographical distances are in general used to explain a part of the genetic differentiations. But as our fundamental assumption states that landscape heterogeneities and spatial arrangements of landscape features may strongly affect dispersal successes, genetic distance between populations must be better explained by the estimate of a model which considers these factors. We have tested this assumption with the greater white-toothed shrew (Crocidura russula). Scenarios considering various behaviours and dispersal abilities of C. russula have been performed. Relating measures of genetic, geographical and ecological distances (the latter emerge from scenario simulation results) highlights the model capability to reproduce dispersal of C. russula by explaining a greater part of the genetic differentiation than that explained by the geographical distances. This application has not only pointed out the ability of the model to quantify connectivity between habitat patches but also the difficulty to relate gene dispersal and individual dispersal.

EPFL2003

Amphibian conservation in human shaped environments

Flavio Zanini

Global biodiversity is experiencing a worrying decline. Habitats destruction, associated to their degradation and fragmentation are among the greatest causes. Amphibians are particularly interesting because they are more threatened and decline more rapidly than either birds or mammals. In this context, the objective of our research is to improve some methodological approaches and offer practical scientific bases for decision making in landscape management and amphibian conservation. Our study focuses on fragmented Swiss landscapes. We developed a method that uses land-cover data and expert knowledge to enable a spatially explicit assessment of 1) the temporal changes in the nature conservation value of the landscape and 2) the rehabilitation potential of the landscape. We applied this Geographical Information Systems (GIS) based approach in the Swiss Rhone plain and we used the years 1900 as the reference state. The method constitutes a helpful tool for communication, decision-making and biological conservation management in landscape planning. Effective and optimal species management strategies can only be formulated after relationships between species distribution and environmental factors have been identified. Concerning amphibians, several approaches exist but they generally suffer from two limitations: 1) the spatial autocorrelation (i.e. the dependency between two observations) in data is rarely explicitly analyzed, even if it may affect the accuracy of species-habitat relationships models. We showed how this spatial autocorrelation can be measured and included in logistic models with the example of the agile frog (Rana dalmatina) in north-eastern Switzerland. We used the Moran's I and the autologistic model (i.e. a logistic model including a measure of the spatial arrangement of the response variables). We found that if spatial autocorrelation is not considered, then conclusions on species-habitat relationships can be incorrect. 2) The effect of landscape on amphibian occurrence in ponds is often assumed to be equal in every direction (isotropic). However, barriers and inhospitable surfaces may reduce movement patterns and the area around ponds accessible to species. This implies that the ideal circular area has in reality a shape depending on the surrounding landscape. We developed a method to determine the effect of habitat variables on amphibian species distribution, considering physical barriers in their movement around ponds. We studied two amphibian species: the common toad (Bufo bufo) and the common frog (Rana temporaria) in the Rhone plain. We demonstrated that reducing the boundaries of circular area following barriers, allowed to compute landscape predictors which better explained species distribution. These results suggested that the proposed approach is more pertinent than the traditional circular buffers analysis. Our results stress the necessity to consider barriers and ecological corridors in species distribution models in order to avoid incorrect inferences. Species distribution models are usually established for a single region. It is generally unknown whether the identified relationships between species distribution and environmental variables can be directly transferred to another geographical area. We examined landscape-level habitat relationships for six amphibian species by measuring correlations with their presence in 655 ponds of five different regions. We analyzed several models by using the information-theoretic approach and the Akaike Information criterion (AIC). For five out of six species, the best models predict that site occupation probability depends on region. Our results suggest that caution is needed when predictions of species occurrence and species management strategies are done using models built in other geographic regions. We also observed that connectivity was generally more explicative than landscape variables. In addition, we found that the spatial scale at which habitat affected species occurrence varied from pond to several km around ponds. Management strategies for amphibian conservation should be conducted taking into account the geographic context, connectivity of ponds and habitat characteristics at multiple spatial scales. Finally, we demonstrated that the landscape, separating patches in metapopulation models, has to be considered in order to avoid incorrect conclusions on population viability analyses. We explored how patch occupancy is sensitive to Euclidean (shortest) versus a landscape-based distance (least-cost). We found: 1) from a theoretical standpoint, that inter-patch landscape affects patch occupancy; 2) from a practical and conservation standpoints, which patches should be considered in priority for landscape management. The approach was illustrated in the case of two metapopulations of the Yellow-bellied Toad in the Rhone plain. We applied successfully the developed practical approaches to the case of several amphibian species, but they can doubtlessly be extended to any species functioning on a spatially defined patch basis (e.g. pond, nesting place, den …), structured as a metapopulation and affected by landscape structure during movement. By improving and combining spatially explicit approaches, we are more likely to provide wildlife managers with tools for valuable decision making.

EPFL2006

Theoretical and Experimental Studies of Dendritic Metacommunities

Francesco Carrara

The present thesis deals with the understanding of the origins and the mechanisms of maintenance of biodiversity in natural landscapes, in particular by identifying key processes that define large-scale patterns of abundance and diversity. Biological communities often occur in spatially structured habitats where connectivity directly affects dispersal and metacommunity processes. Recent theoretical work suggests that dispersal constrained by the connectivity of specific habitat structures affects diversity patterns and species interactions. This is particularly relevant in dendritic networks epitomized by fluvial ecological corridors. This thesis addresses whether connectivity alone can explain observed features of biodiversity and selectively promote different components of community composition in river-like landscapes, such as local species richness or the among-community similarity. The relevance of this thesis lies in the major ecological challenges posed by the topic, and its fundamental importance to conservation biology. The studies pursued herein are also deemed relevant because of the influence of the spatial connectivity and dispersal on population dynamics and of the relevance of biodiversity to ecosystem functioning. Mechanisms of species coexistence were investigated with a blend of theoretical tools (broadly related to statistical mechanics and the theory of stochastic processes) and experimental work using laboratory microbial communities. The research tools ranged from aspects of modern coexistence theory in a local perspective to the recent concept of the metacommunity in spatial ecology, within a unified framework. The study of biodiversity in riverine ecosystems guided by observational data has been addressed by combining theoretical metacommunity models with laboratory experiments. The results are diverse. First, they show experimentally that connectivity per se shapes key components of biodiversity in metacommunities. Local dispersal in isotropic lattice landscapes homogenizes local species richness and leads to pronounced spatial persistence. By contrast, dispersal along dendritic landscapes leads to higher variability in local diversity and among-community composition. Although headwaters exhibit relatively lower species richness, they are crucial for the maintenance of regional biodiversity. By suitably arranging patch sizes within river-like networks the effect of local habitat capacity (i.e., the patch size) and dendritic connectivity on biodiversity can be experimentally disentangled in aquatic microcosm metacommunities. It is shown in this thesis that species coexistence and community assembly depend on intricate, non-trivial interactions of local community capacity and network positioning. Furthermore, an interaction of spatial arrangement of habitat capacity and dispersal along river-like networks also affects a key ecosystem descriptor, namely regional evenness. High regional evenness in community composition is found only in landscapes preserving geomorphological scaling properties of patch sizes. In riverine environments some of the rarer species sustained regionally more abundant populations and were better able to track their own niche requirements compared to landscapes with homogeneous patch size or landscapes with spatially uncorrelated patch size. All the experimental results were supported and extended by a theoretical analysis where the above mechanisms have been generalized. This thesis provides the first direct experimental evidence that spatially constrained dendritic connectivity is a key factor for community composition and population persistence in riverine landscapes. As such, this thesis assesses a longstanding issue in spatial community ecology. It offers unique insights into the ecological forces structuring natural communities in a key ecosytem, and demonstrates principles that can be further tested in theoretical metacommunity models possibly to be extended to real riverine ecosystems. Taken together, the analyses show how the structure of ecological networks interacts with the spatial environmental matrix in determining biodiversity patterns and the functioning of biological communities. The analyses also suggest that altering the natural linkage between dendritic connectivity and patch size strongly affects community properties at multiple scales. The first part of this thesis (chapters 2 and 3) addresses key aspects of biodiversity-ecosystem functioning research where the combination of theory-guided experiments and theoretical investigations shows how a stochastic implementation of population dynamics proves fundamental for key community properties such as species persistence and community stability. The diversity-productivity and diversity-stability relationships are explored. Both experimental findings and the results of a stochastic model fitted to the experimental interaction matrix, suggest the emergence of strong stabilizing forces when species from different functional groups interact in the same environment, increasing species coexistence and community biomass production. The last part (chapter 6) provides a synthesis of this thesis work, in that it aims at unifying aspects from niche-theory, usually adopted in spatially implicit models, with those characteristic of a spatially explicit context from a typical real-life mountainous regions. It is dedicated to the possible explanation for a macroecological pattern routinely observed from organisms in different domains of life, that is, the mid-elevational peak in local species richness. Guided by empirical observations on diversity of macroinvertebrates in Swiss river basins, a theoretical ansatz is provided which is deemed to capture the essential geomorphological drivers and controls relating species-fitness to altitude. A set of overarching conclusions and perspectives for future research are discussed in the concluding chapter.

EPFL2013