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.     

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.     

Influence of grain size on species-habitat models

High resolution remote sensing data facilitate the use of small-scale habitat features such as trees or hedges in the analysis of species-habitat relationships. Such data potentially enable more accurate species-habitat mapping than lower resolution data. Here, for the first time, we systematically investigated this hypothesis by altering the spatial resolution from 1 m up to 1000 m grain size in species-habitat models of 13 bird species. The study area covered the Nidda river catchment in central Germany, a large heterogeneous landscape of 1620 km². A high resolution habitat map of the area was converted to coarser spatial and thematic resolutions in seven steps. We investigated how model performance responded to grain size, and we compared the differential effects of spatial resolution and thematic resolution on model performance. Explained deviance (D²) of the bird models generally decreased with coarser spatial resolution of the data, although it did not decrease monotonically in all species. On average across all species, model D² decreased from 41.5 at 1 m grain size to 15.9 at 1000 m grain size. Ten species were best modelled at 1 m, two species at 3 m and one species at 32 m grain size. Model performance degraded continuously with increasing grain size, both in habitat generalist and habitat specialist bird species, and was systematically lower in habitat generalists. The higher model performance observed at finer grain sizes was most likely caused by the combination of three factors: (1) high spatial accuracy of bird records and (2) a more precise location and delineation of habitat features and, (3) to a lesser degree, by more habitat types differentiated in maps of finer resolution. We conclude that higher spatial and thematic resolution data can be essential for deriving accurate predictions on bird distribution patterns from species-habitat models. Especially for bird species that are sensitive to specific land-use types or to small-scaled habitat features, a grain size of 1-3 m seems most promising.     

Incorporating spatial autocorrelation into cellular automata model: An application to the dynamics of Chinese tamarisk (Tamarix chinensis Lour.)

Spatial autocorrelation (SAC) is frequently encountered in most spatial data in ecology. Cellular automata (CA) models have been widely used to simulate complex spatial phenomena. However, little has been done to examine the impact of incorporating SAC into CA models. Using image-derived maps of Chinese tamarisk (Tamarix chinensis Lour.), CA models based on ordinary logistic regression (OLCA model) and autologistic regression (ALCA model) were developed to simulate landscape dynamics of T. chinensis. In this study, significant positive SAC was detected in residuals of ordinary logistic models, whereas non-significant SAC was found in autologistic models. All autologistic models obtained lower Akaike's information criterion corrected for small sample size (AICc) values than the best ordinary logistic models. Although the performance of ALCA models only satisfied the minimum requirement, ALCA models showed considerable improvement upon OLCA models. Our results suggested that the incorporation of the autocovariate term not only accounted for SAC in model residuals but also provided more accurate estimates of regression coefficients. The study also found that the neglect of SAC might affect the statistical inference on underlying mechanisms driving landscape changes and obtain false ecological conclusions and management recommendations. The ALCA model is statistically sound when coping with spatially structured data, and the adoption of the ALCA model in future landscape transition simulations may provide more precise probability maps on landscape transition, better model performance and more reasonable mechanisms that are responsible for landscape changes.     

The Importance of Being Spatial (and Reserved): Assessing Northern Spotted Owl Habitat Relationships with Hierarchical Bayesian Models

Abstract:  Regional conservation planning increasingly draws on habitat suitability models to support decisions regarding land allocation and management. Nevertheless, statistical techniques commonly used for developing such models may give misleading results because they fail to account for 3 factors common in data sets of species distribution: spatial autocorrelation, the large number of sites where the species is absent (zero inflation), and uneven survey effort. We used spatial autoregressive models fit with Bayesian Markov Chain Monte Carlo techniques to assess the relationship between older coniferous forest and the abundance of Northern Spotted Owl nest and activity sites throughout the species' range. The spatial random-effect term incorporated in the autoregressive models successfully accounted for zero inflation and reduced the effect of survey bias on estimates of species–habitat associations. Our results support the hypothesis that the relationship between owl distribution and older forest varies with latitude. A quadratic relationship between owl abundance and older forest was evident in the southern portion of the range, and a pseudothreshold relationship was evident in the northern portion of the range. Our results suggest that proposed changes to the network of owl habitat reserves would reduce the proportion of the population protected by up to one-third, and that proposed guidelines for forest management within reserves underestimate the proportion of older forest associated with maximum owl abundance and inappropriately generalize threshold relationships among subregions. Bayesian spatial models can greatly enhance the utility of habitat analysis for conservation planning because they add the statistical flexibility necessary for analyzing regional survey data while retaining the interpretability of simpler models.     

Null models reveal preferential sampling, spatial autocorrelation and overfitting in habitat suitability modelling

Nowadays, species are driven to extinction at a high rate. To reduce this rate it is important to delineate suitable habitats for these species in such a way that these areas can be suggested as conservation areas. The use of habitat suitability models (HSMs) can be of great importance for the delineation of such areas. In this study MaxEnt, a presence-only modelling technique, is used to develop HSMs for 223 nematode species of the Southern Bight of the North Sea. However, it is essential that these models are beyond discussion and they should be checked for potential errors. In this study we focused on two categories (1) errors which can be attributed to the database such as preferential sampling and spatial autocorrelation and (2) errors induced by the modelling technique such as overfitting, In order to quantify these adverse effects thousands of nulls models were created. The effect of preferential sampling (i.e. some areas where visited more frequenty than others) was investigated by comparing model outcomes based from null models sampling the actual sampling stations and null models sampling the entire mapping area (Raes and ter Steege, 2007). Overfitting is exposed by a fivefold cross-validation and the influence of spatial autocorrelation is assessed by separating test and training sets in space. Our results clearly show that all these effects are present: preferential sampling has a strong effect on the selection of non-random species models. Crossvalidation seems to have less influence on the model selection and spatial autocorrelation is also strongly present. It is clear from this study that predefined thresholds are not readily applicable to all datasets and additional tests are needed in model selection.     

The performance of state-of-the-art modelling techniques depends on geographical distribution of species

We explored the effects of prevalence, latitudinal range and clumping (spatial autocorrelation) of species distribution patterns on the predictive accuracy of eight state-of-the-art modelling techniques: Generalized Linear Models (GLMs), Generalized Boosting Method (GBM), Generalized Additive Models (GAMs), Classification Tree Analysis (CTA), Artificial Neural Network (ANN), Multivariate Adaptive Regression Splines (MARS), Mixture Discriminant Analysis (MDA) and Random Forest (RF). One hundred species of Lepidoptera, selected from the Distribution Atlas of European Butterflies, and three climate variables were used to determine the bioclimatic envelope for each butterfly species. The data set consisting of 2620 grid squares 30′ × 60′ in size all over Europe was randomly split into the calibration and the evaluation data sets. The performance of different models was assessed using the area under the curve (AUC) of a receiver operating characteristic (ROC) plot. Observed differences in modelling accuracy among species were then related to the geographical attributes of the species using GAM. The modelling performance was negatively related to the latitudinal range and prevalence, whereas the effect of spatial autocorrelation on prediction accuracy depended on the modelling technique. These three geographical attributes accounted for 19–61% of the variation in the modelling accuracy. Predictive accuracy of GAM, GLM and MDA was highly influenced by the three geographical attributes, whereas RF, ANN and GBM were moderately, and MARS and CTA only slightly affected. The contrasting effects of geographical distribution of species on predictive performance of different modelling techniques represent one source of uncertainty in species spatial distribution models. This should be taken into account in biogeographical modelling studies and assessments of climate change impacts.     

Predicting barrier passage and habitat suitability for migratory fish species

Fish migrate to spawn, feed, seek refuge from predators, and escape harmful environmental conditions. The success of upstream migration is limited by the presence of barriers that can impede the passage of fish. We used a spatially explicit modeling strategy to examine the effects of barriers on passage for 21 native and non-native migratory fish species and the amount of suitable habitat blocked for each species. Spatially derived physical parameter estimates and literature based fish capabilities and tolerances were used to predict fish passage success and habitat suitability. Both the fish passage and the habitat suitability models accurately predicted fish presence above barriers for most common, non-stocked species. The fish passage model predicted that barriers greater than or equal to 6 m block all migratory species. Chinook salmon (Oncorhynchus tshawytscha) was expected to be blocked the least. The habitat suitability model predicted that low gradient streams with intact habitat quality were likely to support the highest number of fish species. The fish passage and habitat suitability models were intended to be used by environmental managers as strategy development tools to prioritize candidate dams for field assessment and make decisions regarding the management of migratory fish populations.     

Landscape-level impact of tropical forest loss and fragmentation on bird occurrence in eastern Guatemala

Tropical forest destruction and fragmentation of habitat patches may reduce population persistence at the landscape level. Given the complex nature of simultaneously evaluating the effects of these factors on biotic populations, statistical presence/absence modelling has become an important tool in conservation biology. This study uses logistic regression to evaluate the independent effects of tropical forest cover and fragmentation on bird occurrence in eastern Guatemala. Logistic regression models were constructed for 10 species with varying response to habitat alteration. Predictive variables quantified forest cover, fragmentation and their interaction at three different radii (200, 500 and 1000 m scales) of 112 points where presence of target species was determined. Most species elicited a response to the 1000 m scale, which was greater than most species’ reported territory size. Thus, their presence at the landscape scale is probably regulated by extra-territorial phenomena, such as dispersal. Although proportion of forest cover was the most important predictor of species’ presence, there was strong evidence of area-independent and -dependent fragmentation effects on species presence, results that contrast with other studies from northernmost latitudes. Species’ habitat breadth was positively correlated with AIC model values, indicating a better fit for species more restricted to tropical forest. Species with a narrower habitat breadth also elicited stronger negative responses to forest loss. Habitat breadth is thus a simple measure that can be directly related to species’ vulnerability to landscape modification. Model predictive accuracy was acceptable for 4 of 10 species, which were in turn those with narrower habitat breadths.     

Comparison of statistical and theoretical habitat models for conservation planning: the benefit of ensemble prediction

Selection of a modeling approach is an important step in the conservation planning process, but little guidance is available. We compared two statistical and three theoretical habitat modeling approaches representing those currently being used for avian conservation planning at landscape and regional scales: hierarchical spatial count (HSC), classification and regression tree (CRT), habitat suitability index (HSI), forest structure database (FS), and habitat association database (HA). We focused our comparison on models for five priority forest-breeding species in the Central Hardwoods Bird Conservation Region: Acadian Flycatcher, Cerulean Warbler, Prairie Warbler, Red-headed Woodpecker, and Worm-eating Warbler. Lacking complete knowledge on the distribution and abundance of each species with which we could illuminate differences between approaches and provide strong grounds for recommending one approach over another, we used two approaches to compare models: rank correlations among model outputs and comparison of spatial correspondence. In general, rank correlations were significantly positive among models for each species, indicating general agreement among the models. Worm-eating Warblers had the highest pairwise correlations, all of which were significant (P < 0.05). Red-headed Woodpeckers had the lowest agreement among models, suggesting greater uncertainty in the relative conservation value of areas within the region. We assessed model uncertainty by mapping the spatial congruence in priorities (i.e., top ranks) resulting from each model for each species and calculating the coefficient of variation across model ranks for each location. This allowed identification of areas more likely to be good targets of conservation effort for a species, those areas that were least likely, and those in between where uncertainty is higher and thus conservation action incorporates more risk. Based on our results, models developed independently for the same purpose (conservation planning for a particular species in a particular geography) yield different answers and thus different conservation strategies. We assert that using only one habitat model (even if validated) as the foundation of a conservation plan is risky. Using multiple models (i.e., ensemble prediction) can reduce uncertainty and increase efficacy of conservation action when models corroborate one another and increase understanding of the system when they do not.     

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.     

Tradeoffs between forestry resource and conservation values under alternate policy regimes: A spatial analysis of the western Canadian boreal plains

An important element of resource management and conservation is an understanding of the tradeoffs between marketed products, such as timber, and measures of environmental quality, such as biodiversity. In this paper, we develop an integrated economic-ecological spatial optimization model that we then apply to evaluate alternate forest policies on a 560,000 km² study region of managed boreal forest in Alberta and British Columbia, Canada. The integrated model incorporates dynamic forest sector harvesting, current levels of oil and gas sector development, coarse-filter or habitat-based old forest indicators, a set of empirical forest bird abundance models, and statistical models of the natural and current fire regimes. Using our integrated model, economic tradeoff curves, or production possibility frontiers, are developed to illustrate the cost of achieving coarse-filter targets by a set time (50 years) within a 100-year time horizon. We found levels of ecological indicators and economic returns from the timber industry could both be increased if spatial constraints imposed by the current policy environment were relaxed; other factors being equal, this implies current policy should be revised. We explore the production possibility frontier's relationship to the range of natural variation of old forest habitat, and show how this range can be used to guide choices of preferred locations along the frontier. We also show that coarse-filter constraints on the abundance of certain habitat elements are sufficient to satisfy some fine-filter objectives, expressed as the predicted abundances of various species of songbirds.     

Testing the Generality of Bird-Habitat Models

Bird-habitat models are frequently used as predictive modeling tools—for example, to predict how a species will respond to habitat modifications. We investigated the generality of the predictions from this type of model. Multivariate models were developed for Golden Eagle (Aquila chrysaetos), Raven (Corvus corax), and Buzzard (Buteo buteo) living in northwest Scotland. Data were obtained for all habitat and nest locations within an area of 2349 km². This assemblage of species is relatively static with respect to both occupancy and spatial positioning. The area was split into five geographic subregions: two on the mainland and three on the adjacent Island of Mull, which has one of United Kingdom's richest raptor fauna assemblages. Because data were collected for all nest locations and habitats, it was possible to build models that did not incorporate sampling error. A range of predictive models was developed using discriminant analysis and logistic regression. The models differed with respect to the geographical origin of the data used for model development. The predictive success of these models was then assessed by applying them to validation data. The models showed a wide range of predictive success, ranging from only 6% of nest sites correctly predicted to 100% correctly predicted. Model validation techniques were used to ensure that the models' predictions were not statistical artefacts. The variability in prediction success seemed to result from methodological and ecological processes, including the data recording scheme and interregional differences in nesting habitat. The results from this study suggest that conservation biologists must be very careful about making predictions from such studies because we may be working with systems that are inherently unpredictable.     

Extinction debt of forest plants persists for more than a century following habitat fragmentation

Following habitat fragmentation individual habitat patches may lose species over time as they pay off their "extinction debt." Species with relatively low rates of population extinction and colonization ("slow" species) may maintain extinction debts for particularly prolonged periods, but few data are available to test this prediction. We analyzed two unusually detailed data sets on forest plant distributions and land-use history from Lincolnshire, United Kingdom, and Vlaams-Brabant, Belgium, to test for an extinction debt in relation to species-specific extinction and colonization rates. Logistic regression models predicting the presence-absence of 36 plant species were first parameterized using data from Lincolnshire, where forest cover has been relatively low (approximately 5-8%) for the past 1000 years. Consistent with extinction debt theory, for relatively slow species (but not fast species) these models systematically underpredicted levels of patch occupancy in Vlaams-Brabant, where forest cover was reduced from approximately 25% to <10% between 1775 and 1900 (it is presently 6.5%). As a consequence, the ability of the Lincolnshire models to predict patch occupancy in Vlaams-Brabant was worse for slow than for fast species. Thus, more than a century after forest fragmentation reached its current level an extinction debt persists for species with low rates of population turnover.     

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.     

GIS-based modeling of the geographic distribution of Quercus emoryi Torr. (Fagaceae) in México and identification of significant environmental factors influencing the species’ distribution

The greatest concentration of oak species in the world is believed to be found in Mexico. These species are potentially useful for reforestation because of their capacity to adapt to diverse environments. Knowledge of their geographic distribution and of species–environment relations is essential for decision-making in the management and conservation of natural resources. The objectives of this study were to develop a model of the distribution of Quercus emoryi Torr. in Mexico, using geographic information systems and data layers of climatic and other variables, and to determine the variables that significantly influence the distribution of the species. The study consisted of the following steps: (A) selection of the target species from a botanical scientific collection, (B) characterization of the collecting sites using images with values or categories of the variables, (C) model building with the overlay of images that meet the habitat conditions determined from the characterization of sites, (D) model validation with independent data in order to determine the precision of the model, (E) model calibration through adjustment of the intervals of some variables, and (F) sensitivity analysis using precision and concordance non-parametric statistics applied to pairs of images. Results show that the intervals of the variables that best describe the species’ habitat are the following: altitude from 1650 to 2750 amsl, slope from 0 to 66°; average minimum temperature of January from −12 to −3 °C; mean temperature of June from 11 to 25 °C; mean annual precipitation from 218 to 1225 mm; soil units: lithosol, eutric cambisol, haplic phaeozem, chromic luvisol, rendzina, luvic xerosol, mollic planosol, pellic vertisol, eutric regosol; type of vegetation: oak forest, oak–pine forest, pine forest, pine–oak forest, juniperus forest, low open forest, natural grassland and chaparral. The resulting model of the geographic distribution of Quercus emoryi in Mexico had the following values for non-parametric statistics of precision and agreement: Kappa index of 0.613 and 0.788, overall accuracy of 0.806 and 0.894, sensitivity of 0.650 and 0.825, specificity of 0.963, positive predictive value of 0.945 and 0.957 and negative predictive value of 0.733 and 0.846. Results indicate that the variable average minimum temperature of January, with a maximum value of −3 °C, is an important factor in limiting the species’ distribution.     

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.     

Spatial occurrence of a habitat-tracking saproxylic beetle inhabiting a managed forest landscape.

Because of the dynamic nature of many managed habitats, proper evaluation of conservation efforts calls for models that take into account both spatial and temporal habitat dynamics. We develop a metapopulation model for successional-type systems, in which habitat quality changes over time in a predictable fashion. The occupancy and recruitment of the predatory saproxylic (dependent on dead wood) beetle Harminius undulatus was studied in a managed boreal forest landscape, covering 24,449 ha, in central Sweden. In a first step, we analyzed the beetle's occupancy pattern in relation to stand characteristics, and the amounts of present and past habitat in the surrounding landscape. Managed forest is suitable habitat when > or =60 years old, and immediately after cutting, but not between the ages of 10 and 60 years. The observed occupancy of H. undulatus was positively correlated with the stand's age as habitat. We used a metapopulation model to predict the current probability of occurrence in each forest stand, given the spatiotemporal distribution of suitable forest stands during the last 50 years. Metapopulation parameters were estimated by matching predicted spatial distributions with observed spatial distributions. The model predicted observed spatial distributions better than a similar model that assumed constant habitat quality of each forest stand. Thus, metapopulation models for successional-type systems, such as dead wood dependent organisms in managed forest landscapes, should include habitat dynamics. An estimated 82% of the landscape-wide recruitment took place in managed stands, which covered 87% of the forest area, in comparison with 18% in unmanaged stands, which covered 13% of the forest area. Among the managed stand types, > or =60-year-old stands and 3-7-year-old clear-cuttings contributed to 79% of the total recruitment while 8-59-year-old stands only contributed 3%. The results suggest the following guidelines to improve conditions for H. undulatus and other species with similar habitat requirements: (1) the proportion of the landscape constituted by younger stands should not be allowed to grow too large, (2) the rotation period of managed stands should not be allowed to be too short, and (3) dead wood should be retained and created at final cutting.     

Cumulative ecological and socioeconomic effects of forest policies in coastal Oregon.

Forest biodiversity policies in multi-ownership landscapes are typically developed in an uncoordinated fashion with little consideration of their interactions or possible unintended cumulative effects. We conducted an assessment of some of the ecological and socioeconomic effects of recently enacted forest management policies in the 2.3-million-ha Coast Range Physiographic Province of Oregon. This mountainous area of conifer and hardwood forests includes a mosaic of landowners with a wide range of goals, from wilderness protection to high-yield timber production. We projected forest changes over 100 years in response to logging and development using models that integrate land use change and forest stand and landscape processes. We then assessed responses to those management activities using GIS models of stand structure and composition, landscape structure, habitat models for focal terrestrial and aquatic species, timber production, employment, and willingness to pay for biodiversity protection. Many of the potential outcomes of recently enacted policies are consistent with intended goals. For example, we project the area of structurally diverse older conifer forest and habitat for late successional wildlife species to strongly increase. 'Other outcomes might not be consistent with current policies: for example, hardwoods and vegetation diversity strongly decline within and across owners. Some elements of biodiversity, including streams with high potential habitat for coho salmon (Oncorhynchus kisutch) and sites of potential oak woodland, occur predominately outside federal lands and thus were not affected by the strongest biodiversity policies. Except for federal lands, biodiversity policies were not generally characterized in sufficient detail to provide clear benchmarks against which to measure the progress or success. We conclude that land management institutions and policies are not well configured to deal effectively with ecological issues that span broad spatial and temporal scales and that alternative policies could be constructed that more effectively provide for a mix of forest values from this region.     

How does the spatial structure of habitat loss affect the eco-epidemic dynamics?

Habitat loss is considered as one of the primary causes of species extinction, especially for a species that also suffers from an epidemic disease. Little attention has been paid to the combined effect of habitat loss and epidemic transmission on the species spatiotemporal dynamics. Here, a spatial model of the parasite–host/prey–predator eco-epidemiological system with habitat loss was studied. Habitat patches in the model, instead of undergoing a random loss, were spatially clustered by different degrees. Not only the quantity of habitat loss but also its clustering degree was shown to affect the equilibrium of the system. The infection rate and the probability of successful predation were keys to determine the spatial patterns of species. The epidemic disease is more likely to break out if only a small amount of suitable patches were lost. Counter-intuitively, infected preys are more sensitive to habitat loss than predators if the lost patches are highly clustered. This result is new to eco-epidemiology and implies a possibility of using spatial arrangement of suitable (or unsuitable) patches to control the spread of epidemics in the ecological system.