The importance of spatial autocorrelation,extent and resolution in predicting forest bird occurrence

Concerns about declines in forest biodiversity underscore the need for accurate estimates of the distribution and abundance of organisms at large scales and at resolutions that are fine enough to be appropriate for management. This paper addresses three major objectives: (i) to determine whether the resolution of typical air photo-derived forest inventory is sufficient for the accurate prediction of site occupancy by forest birds. We compared prediction success of habitat models using air photo variables to models with variables derived from finer resolution, ground-sampled vegetation plots. (ii) To test whether incorporating spatial autocorrelation into habitat models via autologistic regression increases prediction success. (iii) To determine whether landscape structure is an important factor in predicting bird distribution in forest-dominated landscapes. Models were tested locally (Greater Fundy Ecosystem [GFE]) using cross-validation, and regionally using an independent data set from an area located ca. 250 km to the northwest (Riley Brook [RB]). We found significant positive spatial autocorrelation in the residuals of at least one habitat model for 76% (16/21) of species examined. In these cases, the logistic regression assumption of spatially independent errors was violated. Logistic models that ignored spatial autocorrelation tended to overestimate habitat effects. Though overall prediction success was higher for autologistic models than logistic models in the GFE, the difference was only significantly improved for one species. Further, the inclusion of spatial covariates did little to improve model performance in the geographically discrete study area. For 62% (13/21) of species examined, landscape variables were significant predictors of forest bird occurrence even after statistically controlling for stand-level variability. However, broad spatial extents explained less variation than local factors. In the GFE, 76% (16/21) of air photo and 81% (17/21) of ground plot models were accurate enough to be of practical utility (AUC > 0.7). When applied to RB, both model types performed effectively for 55% (11/20) of the species examined. We did not detect an overall difference in prediction success between air photo and ground plot models in either study area. We conclude that air photo data are as effective as fine resolution vegetation data for predicting site occupancy for the majority of species in this study. These models will be of use to forest managers who are interested in mapping species distributions under various timber harvest scenarios, and to protected areas planners attempting to optimize reserve function.     

Modeling species-habitat relationships with spatially autocorrelated observation data.

Spatial autocorrelation in wildlife observation data arises when extrinsic environmental processes and patterns that influence the spatial distribution of wildlife are themselves spatially structured, or when species are subject to intrinsic population processes, causing contagion or dispersion effects. Territoriality, Allee effects, dispersal limitations, and social clustering are examples of intrinsic processes. Both forms of autocorrelation can violate the assumptions of generalized linear regression models, resulting in biased estimation of model coefficients and diminished predictive performance. Such consequences may be avoided for extrinsic autocorrelation when autocorrelated environmental variables are available for use as model covariates, whereas intrinsic spatial autocorrelation requires an alternative modeling approach. The autologistic model provides an approach suited to the binary observations often obtained in wildlife surveys, but its performance has not been tested across widely varying sampling intensities or strengths of intrinsic spatial structure. Here we use simulated data to test the autologistic model under a range of sampling conditions. The autologistic model obtains better fits and substantially better predictive performance than the standard logistic regression model over the full range of sampling designs and intensities tested. We provide a simple Bayesian implementation of the autologistic model, which until now has not been achieved with standard statistical software alone. A step-by-step procedure is given for characterizing and modeling spatial autocorrelation in binary observation data, along with computer code for fitting autologistic models in WinBUGS, a freeware Bayesian analysis package. This approach avoids normal approximations to the pseudo-likelihood, in contrast to previous Bayesian applications of the autologistic model. We provide a sample application of the autologistic model, fitted to survey data for a gliding marsupial in southeastern Australia.     

The effect of species response form on species distribution model prediction and inference

Ecological theory and current evidence support the validity of various species response curves according to a variety of environmental gradients. Various methods have been developed for building species distribution models but it is not well known how these methods perform under various assumptions about the form of the underlying species response. It is also not well known how spatial correlation in species occurrence affects model performance. These effects were investigated by applying an environmental envelope method (BIOCLIM) and three regression-based methods: logistic regression (LR), generalized additive modelling (GAM), and classification and regression tree (CART) to simulated species occurrence data. Each simulated species was constructed as a sum of responses with varying weights. Three basic species response curves were assumed: Gaussian (bell-shaped), Beta (skew) and linear. The two non-linear responses conform to standard ecological niche theory. All three responses were applied in turn to three simulated environmental variables, each with varying degrees of spatial autocorrelation. GAM produced the most consistent model performance over all forms of simulated species response. BIOCLIM and CART were inclined to underrate the performance of variables with a linear response. BIOCLIM was less sensitive to data density. LR was susceptible to model misspecification. The use of a linear function in LR underestimated the performance of variables with non-linear species response and contributed to increased spatial autocorrelation in model residuals. Omission of important environmental variables with non-linear species response also contributed to increased spatial autocorrelation in model residuals. Adding a spatial autocovariate term to the LR model (autologistic model) reduced the spatial autocorrelation and improved model performance, but did not correct the misidentification of the dominant environmental determinant. This is to be expected since the autologistic approach was designed primarily for prediction and not for inference. Given that various forms of species response to environmental determinants arise commonly in nature: (1) higher order functions should always be tested when applying LR in modelling species distribution; (2) spatial autocorrelation in species distribution model residuals can indicate that environmental determinants with non-linear response are missing from the model; and (3) deficiencies in LR model performance due to model misspecification can be addressed by adding a spatial autocovariate to the model, but care should be taken when interpreting the coefficients of the model parameters.     

Spatial residual analysis of six modeling techniques

In recent years alternative modeling techniques have been used to account for spatial autocorrelations among data observations. They include linear mixed model (LMM), generalized additive model (GAM), multi-layer perceptron (MLP) neural network, radial basis function (RBF) neural network, and geographically weighted regression (GWR). Previous studies show these models are robust to the violation of model assumptions and flexible to nonlinear relationships among variables. However, many of them are non-spatial in nature. In this study, we utilize a local spatial analysis method (i.e., local Moran coefficient) to investigate spatial distribution and heterogeneity in model residuals from those modeling techniques with ordinary least-squares (OLS) as the benchmark. The regression model used in this study has tree crown area as the response variable, and tree diameter and the coordinates of tree locations as the predictor variables. The results indicate that LMM, GAM, MLP and RBF may improve model fitting to the data and provide better predictions for the response variable, but they generate spatial patterns for model residuals similar to OLS. The OLS, LMM, GAM, MLP and RBF models yield more residual clusters of similar values, indicating that trees in some sub-areas are either all underestimated or all overestimated for the response variable. In contrast, GWR estimates model coefficients at each location in the study area, and produces more accurate predictions for the response variable. Furthermore, the residuals of the GWR model have more desirable spatial distributions than the ones derived from the OLS, LMM, GAM, MLP and RBF models.     

Local spatial modeling of white-tailed deer distribution

Complex spatial heterogeneity of ecological systems is difficult to capture and interpret using global models alone. For this reason, recent attention has been paid to local spatial modeling techniques. We used one local modeling approach, geographically weighted regression (GWR), to investigate the effects of local spatial heterogeneity on multivariate relationships of white-tailed deer distribution using land cover patch metrics and climate factors. The results of these analyses quantify differences in the contributions of model parameters to estimates of deer density over space. A GWR model with local kernel bandwidth was compared to a GWR model with global kernel bandwidth and an ordinary least-squares regression (OLS) model with the same parameters to evaluate their relative abilities in modeling deer distributions. The results indicated that the GWR models predicted deer density better than the traditional ordinary least-squares model and also provided useful information regarding local environmental processes affecting deer distribution. GWR model comparisons showed that the local kernel bandwidth GWR model was more realistic than the global kernel bandwidth GWR model, as the latter exaggerated local spatial variation. The parameter estimates and model statistics (e.g., model R²) of the GWR models were mapped using geographic information systems (GIS) to illustrate local spatial variation in the regression relationship and to identify causes of large-scale model misspecifications and low estimation efficiencies.     

Crustacean habitat potential mapping in a tidal flat using remote sensing and GIS

To make a macrofaunal (crustacean) habitat potential map, the spatial distribution of ecological variables in the Hwangdo tidal flat, Korea, was explored. Spatial variables were mapped using remote sensing and a geographic information system (GIS) combined with field observations. A frequency ratio (FR) and logistic regression (LR) model were employed to map the macrofauna potential area for the Ilyoplax dentimerosa, a crustacean species. Spatial variables affecting the tidal macrofauna distribution were selected based on abundance and biomass and used within a spatial database derived from remotely sensed data of various types of sensors. The spatial variables included the intertidal digital elevation model (DEM), slope, distance from a tidal channel, tidal channel density, surface sediment facies, spectral reflectance of the near infrared (NIR) bands and the tidal exposure duration. The relation between the I. dentimerosa and each spatial variable was calculated using the FR and LR. The species was randomly divided into a training set (70%) to analyse habitat potential using FR and LR and a test set (30%) to validate the predicted habitat potential map. The relations were overlaid to produce a habitat potential map with the species potential index (SPI) value for each pixel. The potential habitat maps were compared with the surveyed habitat locations such as validation data set. The comparison results showed that the LR model (accuracy is 85.28%) is better in prediction than the FR (accuracy is 78.96%) model. The performance of models gave satisfactory accuracies. The LR provides the quantitative influence of variables on a potential habitat of species; otherwise, the FR shows the quantitative influence of a class in each variable. The combination of a GIS-based frequency ratio and logistic regression models and remote sensing with field observations is an effective method to determine locations favorable for macrofaunal species occurrences in a tidal flat.     

An innovative computer design for modeling forest landscape change in very large spatial extents with fine resolutions

Although forest landscape models (FLMs) have benefited greatly from ongoing advances of computer technology and software engineering, computing capacity remains a bottleneck in the design and development of FLMs. Computer memory overhead and run time efficiency are primary limiting factors when applying forest landscape models to simulate large landscapes with fine spatial resolutions and great vegetation detail. We introduce LANDIS PRO 6.0, a landscape model that simulates forest succession and disturbances on a wide range of spatial and temporal scales. LANDIS PRO 6.0 improves on existing forest landscape models with two new data structures and algorithms (hash table and run-length compression). The innovative computer design enables LANDIS PRO 6.0 to simulate very large (>10⁸ ha) landscapes with a 30-m spatial resolution, which to our knowledge no other raster forest landscape models can do. We demonstrate model behavior and performance through application to five nested forest landscapes with varying sizes (from 1 million to 100 million 0.09-ha cells) in the southern Missouri Ozarks. The simulation results showed significant and variable effects of changing spatial extent on simulated forest succession patterns. Results highlighted the utility of a model like LANDIS PRO 6.0 that is capable of efficiently simulating large landscapes and scaling up forest landscape processes to a common regional scale of analysis. The programming methodology presented here may significantly advance the development of next generation of forest landscape models.     

Modelling the distribution of plant species using the autologistic regression model

For modeling the distribution of plant species in terms of climate covariates, we consider an autologistic regression model for spatial binary data on a regularly spaced lattice. This model belongs to the class of autologistic models introduced by Besag (1974). Three estimation methods, the coding method, maximum pseudolikelihood method and Markov chain Monte Carlo method are studied and comparedvia simulation and real data examples. As examples, we use the proposed methodology to model the distributions of two plant species in the state of Florida.     

Modeling freshwater fish distributions using multiscale landscape data: A case study of six narrow range endemics

Species distribution models (SDMs) have become integral tools in scientific research and conservation planning. Despite progress in the assessment of various statistical models for use in SDMs, little has been done in way of evaluating appropriate ecological models. In this paper, we evaluate the multiscale filter framework as a suitable theoretical model for predicting freshwater fish distributions in the upper Green River system (Ohio River drainage), USA. The spatial distributions of six fishes with contrasting biogeographies were modeled using boosted regression trees and multiscale landscape data. Species biogeography did not appear to affect predictive performance and all models performed well statistically with receiver operating characteristic area under the curve (AUC) ranging from 0.87 to 0.98. Predictive maps show accurate estimations of ranges for five of six species based on historical collections. The relative influence of each type of environmental feature and spatial scale varied markedly with between species. A hierarchical effect was detected for narrowly distributed species. These species were highly influenced by soil composition at larger spatial scales and land use/land cover (LULC) patterns at more proximal scales. Conversely, LULC pattern was the most influential feature for widely distributed at all spatial scales. Using multiscale data capable of capturing hierarchical landscape influences allowed production of accurate predictive models and provided further insight into factors controlling freshwater fish distributions.     

Spatial uncertainty analysis of population models

This paper describes an approach for conducting spatial uncertainty analysis of spatial population models, and illustrates the ecological consequences of spatial uncertainty for landscapes with different properties. Spatial population models typically simulate birth, death, and migration on an input map that describes habitat. Typically, only a single “reference” map is available, but we can imagine that a collection of other, slightly different, maps could be drawn to represent a particular species’ habitat. As a first approximation, our approach assumes that spatial uncertainty (i.e., the variation among values assigned to a location by such a collection of maps) is constrained by characteristics of the reference map, regardless of how the map was produced. Our approach produces lower levels of uncertainty than alternative methods used in landscape ecology because we condition our alternative landscapes on local properties of the reference map. Simulated spatial uncertainty was higher near the borders of patches. Consequently, average uncertainty was highest for reference maps with equal proportions of suitable and unsuitable habitat, and no spatial autocorrelation. We used two population viability models to evaluate the ecological consequences of spatial uncertainty for landscapes with different properties. Spatial uncertainty produced larger variation among predictions of a spatially explicit model than those of a spatially implicit model. Spatially explicit model predictions of final female population size varied most among landscapes with enough clustered habitat to allow persistence. In contrast, predictions of population growth rate varied most among landscapes with only enough clustered habitat to support a small population, i.e., near a spatially mediated extinction threshold. We conclude that spatial uncertainty has the greatest effect on persistence when the amount and arrangement of suitable habitat are such that habitat capacity is near the minimum required for persistence.     

Predictability of marine nematode biodiversity

In this paper, we investigated: (1) the predictability of different aspects of biodiversity, (2) the effect of spatial autocorrelation on the predictability and (3) the environmental variables affecting the biodiversity of free-living marine nematodes on the Belgian Continental Shelf. An extensive historical database of free-living marine nematodes was employed to model different aspects of biodiversity: species richness, evenness, and taxonomic diversity. Artificial neural networks (ANNs), often considered as “black boxes”, were applied as a modeling tool. Three methods were used to reveal these “black boxes” and to identify the contributions of each environmental variable to the diversity indices. Since spatial autocorrelation is known to introduce bias in spatial analyses, Moran's I was used to test the spatial dependency of the diversity indices and the residuals of the model. The best predictions were made for evenness. Although species richness was quite accurately predicted as well, the residuals indicated a lack of performance of the model. Pure taxonomic diversity shows high spatial variability and is difficult to model. The biodiversity indices show a strong spatial dependency, opposed to the residuals of the models, indicating that the environmental variables explain the spatial variability of the diversity indices adequately. The most important environmental variables structuring evenness are clay and sand fraction, and the minimum annual total suspended matter. Species richness is also affected by the intensity of sand extraction and the amount of gravel of the sea bed.     

Development of an individual-based model to evaluate elk (Cervus elaphus nelsoni) movement and distribution patterns following the Cerro Grande Fire in north central New Mexico, USA

Though studies have modeled the effects of fires on elk, no studies have related the effects of post-fire landscape succession on ungulate movements and distribution using dynamic modeling techniques. The purpose of this study was to develop and test a spatially-explicit, stochastic, individual-based model (IBM) to evaluate potential movement and distribution patterns of elk (Cervus elaphus nelsoni) in relation to spatial and temporal aspects of the Cerro Grande Fire that burned north central New Mexico in May of 2000. Following extensive literature review, the SAVANNA Ecosystem Model was selected to simulate the underlying post-fire successional processes driving elk movement and distribution. Standard logisitic regression was used to analyze habitat-use patterns of ten elk from data collected using global positioning system radio collars while an additional five animals were used as an independent test set during model validation. Static variables in the form of roads, buildings, fences, and habitual use/memory were used to modify a map of impedance values based on the logistic regression of slope, aspect, and elevation. Integration with SAVANNA came through the application of a habitat suitability index (HSI), which combined movement rules written for the IBM and variables modified and produced by the dynamic ecological processes run in SAVANNA. Overall pattern analysis indicated that realistic migrational processes and habitat-use patterns emerged from movement rules incorporated into the IBM in response to advancing and receding snow when compared to the independent test set. Primary and secondary movement pathways emerged from the collective responses of simulated individuals. Using regression analyses, no significant differences between simulated animals and animals used in either model development or an independent test set revealed any differences in response to snow patterns. These considerations suggest the model was adequately corroborated based on existing data and outlined objectives.     

The Application of Neutral Landscape Models in Conservation Biology

Neutral landscape models, derived from percolation theory in the field of landscape ecology, are grid-based maps in which complex habitat distributions are generated by random or fractal algorithms. This grid-based representation of landscape structure is compatible with the raster-based format of geographical information systems (GIS), which facilitates comparisons between theoretical and real landscapes. Neutral landscape models permit the identification of critical thresholds in connectivity, which can be used to predict when landscapes will become fragmented. The coupling of neutral landscape models with generalized population models, such as metapopulation theory, provides a null model for generating predictions about population dynamics in fragmented landscapes. Neutral landscape models can contribute to the following applications in conservation: (1) incorporation of complex spatial patterns in (meta)population models; (2) identification of species' perceptions of landscape structure; (3) determination of landscape connectivity; (4) evaluation of the consequences of habitat fragmentation for population subdivision; (5) identification of the domain of metapopulation dynamics; (6) prediction of the occurrence of extinction thresholds; ( 7) determination of the genetic consequences of habitat fragmentation; and (8) reserve design and ecosystem management. This generalized, spatially explicit framework bridges the gap between spatially implicit, patch-based models and spatially realistic GIS applications which are usually parameterized for a single species in a specific landscape. Development of a generalized, spatially explicit framework is essential in conservation biology because we will not be able to develop individual models for every species of management concern.     

Simulating spatiotemporal dynamics of urbanization with multi-agent systems—A case study of the Phoenix metropolitan region, USA

Urbanization is a human-dominated process and has greatly impacted biodiversity, ecosystem processes, and regional climate. To understand the socioeconomic drivers of urbanization and project future urban landscape changes, multi-agent systems provide a powerful tool. We develop an agent-based model of urban growth for the Phoenix metropolitan region of the United States, which simulates the behavior of regional authorities, real estate developers, residents, and environmentalists. The BDI (Beliefs-Desires-Intentions) structure is employed to simulate the agents behavior and decision models. The heterogeneity of agents is reflected by adjusting parameters according to the agents’ beliefs, desires and preferences. Three scenarios, baseline, economic development priority and environmental protection, are developed and analyzed. The combination of multi-agent system and spatial regression model is employed to predict the future urban development of the Phoenix metropolitan region. Landscape metrics are used to compare the spatial patterns of the urban landscape resulting from different scenarios in different times. In general, with the rapid urban expansion, the shape of urban patches will become more regular as many of them become coalesced. The spatial analysis of urban development through modeling individual and group decisions and human-environment interactions with a multi-agent systems approach can enhance our understanding of the socioeconomic driving forces and mechanisms of urban development.     

Environmental filtering and land-use history drive patterns in biomass accumulation in a mediterranean-type landscape

Aboveground biomass (AGB) reflects multiple and often undetermined ecological and land-use processes, yet detailed landscape-level studies of AGB are uncommon due to the difficulty in making consistent measurements at ecologically relevant scales. Working in a protected mediterranean-type landscape (Jasper Ridge Biological Preserve, California, USA), we combined field measurements with remotely sensed data from the Carnegie Airborne Observatory's light detection and ranging (lidar) system to create a detailed AGB map. We then developed a predictive model using a maximum of 56 explanatory variables derived from geologic and historic-ownership maps, a digital elevation model, and geographic coordinates to evaluate possible controls over currently observed AGB patterns. We tested both ordinary least-squares regression (OLS) and autoregressive approaches. OLS explained 44% of the variation in AGB, and simultaneous autoregression with a 100-m neighborhood improved the fit to an r2 = 0.72, while reducing the number of significant predictor variables from 27 variables in the OLS model to 11 variables in the autoregressive model. We also compared the results from these approaches to a more typical field-derived data set; we randomly sampled 5% of the data 1000 times and used the same OLS approach each time. Environmental filters including incident solar radiation, substrate type, and topographic position were significant predictors of AGB in all models. Past ownership was a minor but significant predictor, despite the long history of conservation at the site. The weak predictive power of these environmental variables, and the significant improvement when spatial autocorrelation was incorporated, highlight the importance of land-use history, disturbance regime, and population dynamics as controllers of AGB.     

Evaluation of discrete host–parasitoid models for diamondback moth and Diadegma semiclausum field time population density series

The performance of discrete mathematical models to describe the population dynamics of diamondback moth (DBM) (Plutella xylostella L.) and its parasitoid Diadegma semiclausum was investigated. The parameter values for several well-known models (Nicholson–Bailey, Hassell and Varley, Beddington, Free and Lawton, May, Holling type 2, 3 and Getz and Mills functional responses) were estimated. The models were tested on 20 consecutive sets of time series data collected at 14 days interval for pest and parasitoid populations obtained from a highland cabbage growing area in eastern Kenya. Model parameters were estimated from minimized squared difference between the numerical solution of the model equations and the empirical data using Powell's method. Maximum calculated DBM growth rates varied between 0.02 and 0.07. The carrying capacity determined at 16.5 DBM/plant by the Beddington et al. model was within the range of field data. However, all the estimated parameter values relating to the parasitoid, including the instantaneous searching rate (0.07–0.28), per capita searching efficiency (0.20–0.27), search time (5.20–5.33), handling time (0.77–0.90), and parasitism aggregation index (0.33), were well outside the range encountered empirically. All models evaluated for DBM under Durbin–Watson criteria, except the May model, were not autocorrelated with respect to residuals. In contrast, the criteria applied to the parasitoid residuals showed strong autocorrelations. Thus, these models failed to estimate parasitoid dynamics. We conclude that the interactions of the DBM with its parasitoid cannot be explained by any of the models tested. Two factors may be associated with this failure. First, the parasitoid in this integrated biological control system may not be playing a major role in regulating DBM population. Second, and perhaps more likely, poor correlations reflect gross inadequacies in the theoretical assumptions that underlie the existing models.     

Applying the Patuxent Landscape Unit Model to human dominated ecosystems: the case of agriculture

Non-spatial dynamics are core to landscape simulations. Unit models simulate system interactions aggregated within one space unit of resolution used within a spatial model. For unit models to be applicable to spatial simulations they have to be formulated in a general enough way to simulate all habitat elements within the landscape. Within the Patuxent River watershed, human dominated land uses, such as agriculture and urban land, are already 50% of the current land use, while urban land is replacing forests, agriculture and wetlands at a rapid rate. The Patuxent Landscape Model (PLM) with the Patuxent General Unit Model as core (Pat-GEM) was developed as a predictive policy tool to estimate environmental impacts of such land use changes. The Pat-GEM is based on the General Ecosystem Model (GEM) developed by [Ecol. Modelling 88 1996 263]. Previous calibrations of the Pat-GEM for anthropogenic land uses have not been satisfactory due to the scarcity of appropriate data. This paper shows Pat-GEM simulations of biomass growth and nutrient uptake for crops typical within the Patuxent watershed. The Pat-GEM was expanded to include processes and fluxes that characterize agricultural land use. The most important extension was to include crop rotation into the model. Additionally, we refined the processes for planting, harvesting and fertilization by introducing specific growth parameters. Our revised Pat-GEM was calibrated against the results from Erosion Productivity Impact Calculator (EPIC) a widely used and calibrated agricultural model. We achieved high correlation between results generated with Pat-GEM and EPIC. The correlation coefficients (r²) varied between 0.87 and 0.98, with the simulation results for winter wheat showing the lowest correlation coefficients. Intercalibration using EPIC is a powerful method for calibrating the Pat-GEM model for agricultural land use. EPIC was able (a) to provide about 30% of the input data required for running the Pat-GEM model; and (b) to provide time series output data (with a daily time step) to calibrate the output variables biomass production and nutrient uptake.     

Modelling habitat overlap among sympatric mesocarnivores in southern Illinois,USA

Few researchers have developed large-scale habitat models for sympatric carnivore species. We created habitat models for red foxes (Vulpes vulpes), coyotes (Canis latrans) and bobcats (Lynx rufus) in southern Illinois, USA, using the Penrose distance statistic, remotely sensed landscape data, and sighting location data within a GIS. Our objectives were to quantify and spatially model potential habitat differences among species. Habitat variables were quantified for 1-km² buffered areas around mesocarnivore sighting locations. Following variable reduction procedures, five habitat variables (percentage of grassland patches, interspersion–juxtaposition of forest patches, mean fractal dimension of wetland patches and the landscape, and road density) were used for analysis. Only one variable differed (P < 0.05) between red fox and coyote sighting areas (road density) and bobcat and coyote sighting areas (mean fractal dimension of the landscape). However, all five variables differed between red fox and bobcat sighting areas, indicating considerable differences in habitat affiliation between this pair-group. Compared to bobcats, red fox sightings were affiliated with more grassland cover and larger grassland patches, higher road densities, lower interspersion and juxtaposition of forest patches, and lower mean fractal dimension of wetland patches. These differences can be explained by different life history requirements relative to specific cover types. We then used the Penrose distance statistic to create habitat models for red foxes and bobcats, respectively, based on the five-variable dataset. An independent set of sighting locations were used to validate these models; model fit was good with 65% of mesocarnivore locations within the top 50% of Penrose distance values. In general, red foxes were affiliated with mixtures of agricultural and grassland cover, whereas bobcats were associated with a combination of grassland, wetland, and forest cover. The greatest habitat overlap between red foxes and bobcats was found at the interface between forested areas and more open cover types. Our study provides insight into habitat overlap among sympatric mesocarnivores, and the distance-based modelling approach we used has numerous applications for modelling wildlife–habitat relationships over large scales.     

Simulating population variation and movement within fragmented landscapes: An application to the gopher tortoise (Gopherus polyphemus)

As the human activity footprint grows, land-use decisions play an increasing role in determining the future of plant and animal species. Studies have shown that urban and agricultural development cannot only harm species populations directly through habitat destruction, but also by destroying the corridors that connect habitat patches and populations within a metapopulation. Without these pathways, populations can encounter inbreeding depression and degeneration, which can increase death rates and lower rates of reproduction. This article describes the development and application of the FRAGGLE model, a spatial system dynamics model designed to calculate connectivity indices among populations. FRAGGLE can help planners and managers identify the relative contribution of populations associated with habitat patches to future populations in those patches, taking into account the importance of interstitial land to migration success. The model is applied to the gopher tortoise (Gopherus polyphemus), a threatened species whose southeastern U.S. distribution has diminished significantly within its native range due to agricultural and urban development over the last several decades. This model is parameterized with life history and movement traits of the gopher tortoise in order to simulate population demographics and spatial distribution within an area in west-central Georgia that supports a significant tortoise population. The implications of this simulation modeling effort are demonstrated using simple landscape representations and a hypothetical on land-use management scenario. Our findings show that development resulting in even limited habitat losses (10%) may lead to significant increases in fragmentation as measured by a loss in the rate of dispersions (31%) among area subpopulations.     

A multi-agent simulation to assess the risk of malaria re-emergence in southern France

A multi-agent simulation (MAS) was developed to assess the risk of malaria re-emergence in the Camargue in southern France, a non-endemic area where mosquitoes of the genus Anopheles (Culicidae) live. The contact rate between people and potential malaria vectors, or the human biting rate, is one of the key factor to predict the risk of re-emergence of malaria, would the parasite be introduced in the region. Our model (called MALCAM) represents the different agents that could influence malaria transmission in the Camargue – people, mosquitoes, animal hosts and the landscape – in a spatially explicit environment. The model simulates spatial and temporal variations in human biting rate at the landscape scale. These variations depend on the distribution of people and potential vectors, their behaviour and their interactions. A land use/cover map was used as a cellular-spatial support for the movements of and interactions between mobile agents. The model was tested for its sensitivity to variations in parameter values, and for the agreement between field observations and model predictions. The MALCAM model provides a tool to better understand the interactions between the multiple agents of the disease transmission system, and the land use and land cover factors that control the spatial heterogeneity in these interactions. It allows testing hypotheses and scenarios related to disease dynamics by varying the value of exogenous biological, geographical, or human factors. This application of agent-based modelling to a human vector-borne disease can be adapted to different diseases and regions.