The priority value of scrubland habitats for carnivore conservation in Mediterranean ecosystems

Carnivores are umbrella species with a key role in ecosystems. In the Mediterranean, they face with several conservation problems, mainly habitat loss or transformation. In this region, scrubland habitats are considered of minor conservation relevance as compared to mature forest formations. Conservation of scrublands in Mediterranean ecosystems is also difficult because to control fires, they are removed over large areas. For carnivores, scrublands may be essential to guarantee shelter and food. Here, we analyzed the importance of scrublands and other habitat variables in determining the richness of medium-sized carnivores in a typical Mediterranean area of central Spain (Monfragüe Natural Park). The Park was divided into plots of 2 × 2 km (n = 30). In each plot, a 2 km-length survey route was walked searching for carnivore scats. We recorded the number of species of carnivores and we related this value to several habitat variables. Habitat variables were summarized by means of Principal Component Analysis (PCA). Habitat models were developed by, using multiple regression models, including PCA factors and the habitat type as predictors. The only variable included in the final model was the first PCA factor. Carnivore richness was positively associated with large tree, shrub, and rock cover areas, which is the typical habitat structure of scrublands. This habitat offers the best available area for shelter and may be considered as a key element for carnivore conservation in the Mediterranean region. To reconciliate carnivore conservation and forestry management, we proposed small-scale clearances as a management alternative to typical large scrubland removals.     

Critical thresholds associated with habitat loss: a review of the concepts, evidence, and applications

A major conservation concern is whether population size and other ecological variables change linearly with habitat loss, or whether they suddenly decline more rapidly below a “critical threshold” level of habitat. The most commonly discussed explanation for critical threshold responses to habitat loss focus on habitat configuration. As habitat loss progresses, the remaining habitat is increasingly fragmented or the fragments are increasingly isolated, which may compound the effects of habitat loss. In this review we also explore other possible explanations for apparently nonlinear relationships between habitat loss and ecological responses, including Allee effects and time lags, and point out that some ecological variables will inherently respond nonlinearly to habitat loss even in the absence of compounding factors. In the literature, both linear and nonlinear ecological responses to habitat loss are evident among simulation and empirical studies, although the presence and value of critical thresholds is influenced by characteristics of the species (e.g. dispersal, reproduction, area/edge sensitivity) and landscape (e.g. fragmentation, matrix quality, rate of change). With enough empirical support, such trends could be useful for making important predictions about species' responses to habitat loss, to guide future research on the underlying causes of critical thresholds, and to make better informed management decisions. Some have seen critical thresholds as a means of identifying conservation targets for habitat retention. We argue that in many cases this may be misguided, and that the meaning (and utility) of a critical threshold must be interpreted carefully and in relation to the response variable and management goal. Despite recent interest in critical threshold responses to habitat loss, most studies have not used any formal statistical methods to identify their presence or value. Methods that have been used include model comparisons using Akaike information criterion (AIC) or t‐tests, and significance testing for changes in slope or for polynomial effects. The judicious use of statistics to help determine the shape of ecological relationships would permit greater objectivity and more comparability among studies.     

Fauna habitat modelling and mapping: A review and case study in the Lower Hunter Central Coast region of NSW

Abstract Habitat models are now broadly used in conservation planning on public lands. If implemented correctly, habitat modelling is a transparent and repeatable technique for describing and mapping biodiversity values, and its application in peri‐urban and agricultural landscape planning is likely to expand rapidly. Conservation planning in such landscapes must be robust to the scrutiny that arises when biodiversity constraints are placed on developers and private landholders. A standardized modelling and model evaluation method based on widely accepted techniques will improve the robustness of conservation plans. We review current habitat modelling and model evaluation methods and provide a habitat modelling case study in the New South Wales central coast region that we hope will serve as a methodological template for conservation planners. We make recommendations on modelling methods that are appropriate when presence‐absence and presence‐only survey data are available and provide methodological details and a website with data and training material for modellers. Our aim is to provide practical guidelines that preserve methodological rigour and result in defendable habitat models and maps. The case study was undertaken in a rapidly developing area with substantial biodiversity values under urbanization pressure. Habitat maps for seven priority fauna species were developed using logistic regression models of species‐habitat relationships and a bootstrapping methodology was used to evaluate model predictions. The modelled species were the koala, tiger quoll, squirrel glider, yellow‐bellied glider, masked owl, powerful owl and sooty owl. Models ranked sites adequately in terms of habitat suitability and provided predictions of sufficient reliability for the purpose of identifying preliminary conservation priority areas. However, they are subject to multiple uncertainties and should not be viewed as a completely accurate representation of the distribution of species habitat. We recommend the use of model prediction in an adaptive framework whereby models are iteratively updated and refined as new data become available.     

Habitat loss-induced tipping points in metapopulations with facilitation

Habitat loss is known to pervade extinction thresholds in metapopulations. Such thresholds result from a loss of stability that can eventually lead to collapse. Several models have been developed to understand the nature of these transitions and how they are affected by the locality of interactions, fluctuations or external drivers. Most models consider the impact of grazing or aridity as a control parameter that can trigger sudden shifts, once critical values are reached. Others explore instead the role played by habitat loss and fragmentation. Here we consider a minimal model incorporating facilitation between the individuals of the same species along with habitat destruction, with the aim of understanding how local cooperation and habitat loss interact with each other. A mathematical model incorporating facilitation and habitat destruction is derived, along with a spatially explicit simulation model. It is found that a catastrophic shift is expected for increasing levels of habitat loss, but the bifurcation becomes continuous when dispersal is local. Under these conditions, spatial patchiness is found and the qualitative change from discontinuous to continuous results are in agreement with previous studies on ecological systems. Our results suggest that species exhibiting facilitation and displaying short-range dispersal will be markedly more capable of avoiding catastrophic tipping points.     

The importance of distance to resources in the spatial modelling of bat foraging habitat

Many bats are threatened by habitat loss, but opportunities to manage their habitats are now increasing. Success of management depends greatly on the capacity to determine where and how interventions should take place, so models predicting how animals use landscapes are important to plan them. Bats are quite distinctive in the way they use space for foraging because (i) most are colonial central-place foragers and (ii) exploit scattered and distant resources, although this increases flying costs. To evaluate how important distances to resources are in modelling foraging bat habitat suitability, we radio-tracked two cave-dwelling species of conservation concern (Rhinolophus mehelyi and Miniopterus schreibersii) in a Mediterranean landscape. Habitat and distance variables were evaluated using logistic regression modelling. Distance variables greatly increased the performance of models, and distance to roost and to drinking water could alone explain 86 and 73% of the use of space by M. schreibersii and R. mehelyi, respectively. Land-cover and soil productivity also provided a significant contribution to the final models. Habitat suitability maps generated by models with and without distance variables differed substantially, confirming the shortcomings of maps generated without distance variables. Indeed, areas shown as highly suitable in maps generated without distance variables proved poorly suitable when distance variables were also considered. We concluded that distances to resources are determinant in the way bats forage across the landscape, and that using distance variables substantially improves the accuracy of suitability maps generated with spatially explicit models. Consequently, modelling with these variables is important to guide habitat management in bats and similarly mobile animals, particularly if they are central-place foragers or depend on spatially scarce resources.     

The Applicability of Habitat Evaluation Methodologies in Ecological Risk Assessment

Traditionally, ecological risk assessments (ERAs) have emphasized risks to individual organisms or populations of species. Although habitats may be a potential target for chemical stressors, and are considered in the framework for ERAs, the actual use of habitat evaluation methods in this process is limited. Habitats obviously represent an important entity to protect since damaged aquatic and wildlife habitats may be totally irretrievable over a human life span compared to deleterious biochemical and physiological changes which may be reversible within the life cycle of an organism, if exposure is terminated. Habitat methods have been largely used as management tools to evaluate impacts of planned water and land development projects. Habitat evaluation methods represent a structured, systematic and logical approach to determine changes to habitats because they consider important life requisites and environmental variables limiting to species. Their use in the ERA process will provide a means to differentiate habitat changes resulting from physical, chemical and/or biological factors or a combination of such factors. In addition, minimal and optimum habitat suitability can be determined for different habitat variables under different chemical exposure scenarios. The objectives of this paper are to review several available habitat evaluation methods and discuss their use in risk assessment. Particular emphasis is given to USFWS's Habitat Evaluation Procedures (HEPs) and the Instream Flow Incremental Method (IFIM).     

论自然保护与资源开发的策略

物种保护的根本途径是保护其生境,生境的完整性遭到破坏则物种将趋于灭绝［１,２］。不少国家的政府及非政府组织正投入巨资设置保护区试图将至少部分生物多样性资源加以永久保存［３］,但这些目标的达到往往是基于如下假设：即保护区将不受干扰、原封不动的加以保存并与经济活动及周围环境难以达成任何形式的妥协。然而越来越多的证据表明,在全球范围遭受人类干扰的背景下无论是保护区内还是保护区外的生境均将趋于退化,因此现行建立保护区的模式并不能有效阻止生物多样性灾难性的损失［４］。特别对于发展中国家,人口、资源、环境的…     

Are Regional Habitat Models Useful at a Local-Scale? A Case Study of Threatened and Common Insectivorous Bats in South-Eastern Australia

Habitat modelling and predictive mapping are important tools for conservation planning, particularly for lesser known species such as many insectivorous bats. However, the scale at which modelling is undertaken can affect the predictive accuracy and restrict the use of the model at different scales. We assessed the validity of existing regional-scale habitat models at a local-scale and contrasted the habitat use of two morphologically similar species with differing conservation status (Mormopterus norfolkensis and Mormopterus species 2). We used negative binomial generalised linear models created from indices of activity and environmental variables collected from systematic acoustic surveys. We found that habitat type (based on vegetation community) best explained activity of both species, which were more active in floodplain areas, with most foraging activity recorded in the freshwater wetland habitat type. The threatened M. norfolkensis avoided urban areas, which contrasts with M. species 2 which occurred frequently in urban bushland. We found that the broad habitat types predicted from local-scale models were generally consistent with those from regional-scale models. However, threshold-dependent accuracy measures indicated a poor fit and we advise caution be applied when using the regional models at a fine scale, particularly when the consequences of false negatives or positives are severe. Additionally, our study illustrates that habitat type classifications can be important predictors and we suggest they are more practical for conservation than complex combinations of raw variables, as they are easily communicated to land managers.     

Ecological filtering or random extinction? Beta‐diversity patterns and the importance of niche‐based and neutral processes following habitat loss

Although both niche‐based and neutral processes are involved in community assembly, most models on the effects of habitat loss are stochastic, assuming neutral communities mainly affected by ecological drift and random extinction. Given that habitat loss is considered the most important driver of the current biodiversity crisis, unraveling the processes underlying the effects of habitat loss is critical from both a theoretical and an applied perspective. Here we unveil the importance of niche‐based and neutral processes to species extinction and community assembly across a gradient of habitat loss, challenging the predictions of neutral models. We draw on a large dataset containing the distribution of 3653 individuals of 42 species, representing 35% of the small mammal species of the Atlantic Forest hotspot, obtained in 68 sites across three continuously‐forested landscapes and three adjacent 10 000‐ha fragmented landscapes differing in the amount of remaining forest (50%, 30% and 10%). By applying a null‐model approach, we investigated β‐diversity patterns by detecting deviations of observed community similarity from the similarity between randomly assembled communities. Species extinction following habitat loss was decidedly non‐random, in contrast to the notion that fragmented communities are mainly driven by ecological drift. Instead, habitat loss led to a strong biotic homogenization. Moreover, species composition changed abruptly at the same level of landscape‐scale habitat loss that has already been associated with a drastic decline in species richness. Habitat loss, as other anthropogenic disturbances, can thus be seen as a strong ecological filter that increases (rather than decreases) the importance of deterministic processes in community assembly. As such, critical advances for the development of conservation science lie on the incorporation of the relevant niche traits associated with extinction proneness into models of habitat loss. The results also underscore the fundamental importance of pro‐active measures to prevent human‐modified landscapes surpassing critical ecological thresholds.     

Habitat suitability models for the conservation of thermophilic grasshoppers and bush crickets—simple or complex?

One goal of conservation biology is the assessment of effects of land use change on species distribution. One approach for identifying the factors, which determine habitat suitability for a species are statistical habitat distribution models. These models are quantitative and can be used for predictions in management scenarios. However, they often have one major shortcoming, which is their complexity. This means that they need several, often costly-to-determine parameters for predictions of species occurrence. We first used habitat suitability models to investigate and determine habitat preferences of three different Orthoptera species. Second, we compared the predictive powers of simple habitat suitability models considering only the ‘habitat type’ as predictor with more complex models taking different habitat factors into account. We found that the habitat type is the most reliable and robust factor, which determines the occurrence of the species studied. Thus, analyses of habitat suitability can easily be carried out on the basis of existing vegetation maps for the conservation of the three species under study. Our results can serve as a basis for the estimation of spatio-temporal distribution and survival probabilities of the species studied and might also be valuable for other species living in dry grasslands.     

Howler monkey tolerance to habitat shrinking: Lifetime warranty or death sentence?

Habitat loss and fragmentation are major threats to the conservation of nonhuman primates. Given that species differ in their responses to fragmented landscapes, identifying the factors that enable them to cope with altered environments or that cause their extirpation is critical to design conservation management strategies. Howler monkeys (Alouatta spp.) are good models for studying the strategies of tolerant arboreal taxa and how they cope with spatial restriction, because they live in habitats ranging from vast pristine forests to small disturbed fragments and orchards. While some aspects of their ecology and behavior are conserved, others vary in predictable ways in response to habitat shrinking and decreasing resource availability. We argue that the ability of individual howler monkeys to inhabit low-quality environments does not guarantee the long-term persistence of the small populations that live under these conditions. Their local extirpation explains why few forest fragments below a given area threshold are frequently inhabited in landscapes where recolonization and gene flow are compromised by long isolation distances or less permeable matrices. In sum, howlers’ ability to cope with habitat restriction at the individual level in the short-term may mask the inevitable fate of isolated populations, thereby compromising the persistence of the species at a regional scale in the long-term if howlers’ need for protection in large forests is undervalued.     

Habitat conservation prioritization: A floristic approach applied to a Mediterranean wetland network

The importance of plant communities for conservation purposes is recognized in their inclusion in the “Habitat Directive” (92/43 EEC), that relates habitat types to plant community syntaxonomic units. However, habitat definitions in the Habitat Directive lead to several inaccuracies in local habitat characterization. Several wetland plant communities (and their corresponding habitats), rare in the Mediterranean basin, are not included in the Habitat Directive. This study proposes criteria for assessing the conservation importance of habitats. It deals with plant community types at the alliance level, as promising units for setting conservation priorities. The principle criteria considered as drivers for setting alliance conservation values are the distribution and abundance of plant species of conservation interest and their fidelity to a plant community. Multivariate methods were used, and a quantitative floristic index of alliance conservation priority was created. This procedure was applied to an important wetland in central Italy. Results emphasize: (i) high conservation values of some alliances not listed in the Habitat Directive, confirming various gaps in the current conservation law affecting Mediterranean wetlands; (ii) that habitats widely distributed in other biogeographical areas, may greatly underestimate their conservation importance in Mediterranean region; (iii) need to consider regional peculiarities when setting conservation priorities.     

Local native plant diversity responds to habitat loss and fragmentation over different time spans and spatial scales

Separating the threats that habitat loss and habitat fragmentation pose to biodiversity is challenging because these processes usually occur simultaneously. Additionally, their importance may be underestimated due to time-delayed extinction. In central Texas savannas, woody plant encroachment reduces the amount of habitat available to herbaceous species while fragmenting remaining habitat. We examined the relationships between present species richness and present and past habitat amount and fragmentation (measured as fractal dimension) using a series of aerial photographs taken over nearly 60 years. We show that woody plant encroachment, a common phenomenon in savannas worldwide, reduces the diversity of herbaceous vegetation through both habitat loss and fragmentation. Habitat loss has the strongest impact on species richness over short time spans and small spatial scales in these savannas. Habitat fragmentation, however, has the strongest impact over longer time spans and larger spatial scales and generates long-term extinction debts. We also demonstrate that examining habitat loss and habitat fragmentation across different time periods and at different spatial scales is essential for understanding their joint and individual effects on plant community composition.     

通过遗传多样性探讨极小种群野生植物的致濒机理及保护策略: 以裸子植物为例

遗传多样性是生物多样性的重要组成部分, 然而由于资源的过度开发利用和生境的破碎化影响了物种的遗传多样性, 甚至威胁到物种的生存适应性和生物多样性。极小种群野生植物是亟待保护的国家重点保护濒危植物,遗传多样性研究对揭示极小种群致濒机理及保护策略具有重要意义。生境破碎化会造成物种遗传多样性降低、种群间分化增加、基因流减少等, 使种群濒危。但在某些物种中, 繁殖特征、进化历史等生物和生态因素的不同也可能造成近期生境破碎化后遗传效应的延迟。裸子植物进化历史悠久, 包含许多孑遗物种, 由于生活史周期长, 遭受生境破碎化后可能短期内显示不出遗传效应的改变, 但长期很难恢复。本文以裸子植物为例综述了濒危植物的遗传多样性研究的案例, 探讨了濒危裸子植物应对环境恶化的维持机制、致濒因素和保护方案, 旨在说明通过遗传多样性研究充分认识极小种群致濒机理对高效保护极小种群野生植物的重要性。     

Detecting Fragmentation Extinction Thresholds for Forest Understory Plant Species in Peninsular Spain

Ecological theory predicts that fragmentation aggravates the effects of habitat loss, yet empirical results show mixed evidences, which fail to support the theory instead reinforcing the primary importance of habitat loss. Fragmentation hypotheses have received much attention due to their potential implications for biodiversity conservation, however, animal studies have traditionally been their main focus. Here we assess variation in species sensitivity to forest amount and fragmentation and evaluate if fragmentation is related to extinction thresholds in forest understory herbs and ferns. Our expectation was that forest herbs would be more sensitive to fragmentation than ferns due to their lower dispersal capabilities. Using forest cover percentage and the proportion of this percentage occurring in the largest patch within UTM cells of 10-km resolution covering Peninsular Spain, we partitioned the effects of forest amount versus fragmentation and applied logistic regression to model occurrences of 16 species. For nine models showing robustness according to a set of quality criteria we subsequently defined two empirical fragmentation scenarios, minimum and maximum, and quantified species’ sensitivity to forest contraction with no fragmentation, and to fragmentation under constant forest cover. We finally assessed how the extinction threshold of each species (the habitat amount below which it cannot persist) varies under no and maximum fragmentation. Consistent with their preference for forest habitats probability occurrences of all species decreased as forest cover contracted. On average, herbs did not show significant sensitivity to fragmentation whereas ferns were favored. In line with theory, fragmentation yielded higher extinction thresholds for two species. For the remaining species, fragmentation had either positive or non-significant effects. We interpret these differences as reflecting species-specific traits and conclude that although forest amount is of primary importance for the persistence of understory plants, to neglect the impact of fragmentation for some species can lead them to local extinction.     

Forest Loss and the Biodiversity Threshold: An Evaluation Considering Species Habitat Requirements and the Use of Matrix Habitats

Habitat loss is the main driver of the current biodiversity crisis, a landscape-scale process that affects the survival of spatially-structured populations. Although it is well-established that species responses to habitat loss can be abrupt, the existence of a biodiversity threshold is still the cause of much controversy in the literature and would require that most species respond similarly to the loss of native vegetation. Here we test the existence of a biodiversity threshold, i.e. an abrupt decline in species richness, with habitat loss. We draw on a spatially-replicated dataset on Atlantic forest small mammals, consisting of 16 sampling sites divided between forests and matrix habitats in each of five 3600-ha landscapes (varying from 5% to 45% forest cover), and on an a priori classification of species into habitat requirement categories (forest specialists, habitat generalists and open-area specialists). Forest specialists declined abruptly below 30% of forest cover, and spillover to the matrix occurred only in more forested landscapes. Generalists responded positively to landscape heterogeneity, peaking at intermediary levels of forest cover. Open area specialists dominated the matrix and did not spillover to forests. As a result of these distinct responses, we observed a biodiversity threshold for the small mammal community below 30% forest cover, and a peak in species richness just above this threshold. Our results highlight that cross habitat spillover may be asymmetrical and contingent on landscape context, occurring mainly from forests to the matrix and only in more forested landscapes. Moreover, they indicate the potential for biodiversity thresholds in human-modified landscapes, and the importance of landscape heterogeneity to biodiversity. Since forest loss affected not only the conservation value of forest patches, but also the potential for biodiversity-mediated services in anthropogenic habitats, our work indicates the importance of proactive measures to avoid human-modified landscapes to cross this threshold.     

A Revised Conservation Assessment of Dipterocarps in Sabah

Borneo has experienced a rapid decline in the extent of forest cover, which has reduced the amount of habitat available for many plant and animal species. The precise impact of habitat loss on the conservation status of dipterocarp trees is uncertain. We use three contrasting techniques, the extent of occurrence, area of occupancy and ecological niche models derived using maxent , in conjunction with a current land‐use map of Sabah, to derive estimates of habitat loss and infer a regional IUCN Red List conservation status for 33 Sabah dipterocarp species. Estimates of habitat loss differed significantly according to the methods employed and between species on different habitat types. Proportion of habitat loss determined from the ecological niche models varied from 21 percent for Shorea micans to 99.5 percent for Dipterocarpus lamellatus. Thirty‐two of the 33 dipterocarp species analyzed in this study would have their Sabah populations classified as Threatened (equal to a habitat loss of > 30%) under the A2 IUCN Red List criterion. Dipterocarps that occur in lowland forests have experienced greater habitat loss than upland/lower montane or ultramafic species. In addition, species with the lowest predicted area within their historic distributions had the highest proportion of habitat lost, which provides a rationale for targeting conservation effort on the species with narrow distributions. We recommend the ecological niche modeling approach as a rapid assessment tool for reconstructing species’ historic distributions during conservation assessments of tropical trees.     

Anticipated climate and land‐cover changes reveal refuge areas for Borneo's orang‐utans

Habitat loss and climate change pose a double jeopardy for many threatened taxa, making the identification of optimal habitat for the future a conservation priority. Using a case study of the endangered Bornean orang‐utan, we identify environmental refuges by integrating bioclimatic models with projected deforestation and oil‐palm agriculture suitability from the 1950s to 2080s. We coupled a maximum entropy algorithm with information on habitat needs to predict suitable habitat for the present day and 1950s. We then projected to the 2020s, 2050s and 2080s in models incorporating only land‐cover change, climate change or both processes combined. For future climate, we incorporated projections from four model and emission scenario combinations. For future land cover, we developed spatial deforestation predictions from 10 years of satellite data. Refuges were delineated as suitable forested habitats identified by all models that were also unsuitable for oil palm – a major threat to tropical biodiversity. Our analyses indicate that in 2010 up to 260 000 km² of Borneo was suitable habitat within the core orang‐utan range; an 18–24% reduction since the 1950s. Land‐cover models predicted further decline of 15–30% by the 2080s. Although habitat extent under future climate conditions varied among projections, there was majority consensus, particularly in north‐eastern and western regions. Across projections habitat loss due to climate change alone averaged 63% by 2080, but 74% when also considering land‐cover change. Refuge areas amounted to 2000–42 000 km² depending on thresholds used, with 900–17 000 km² outside the current species range. We demonstrate that efforts to halt deforestation could mediate some orang‐utan habitat loss, but further decline of the most suitable areas is to be expected given projected changes to climate. Protected refuge areas could therefore become increasingly important for ongoing translocation efforts. We present an approach to help identify such areas for highly threatened species given environmental changes expected this century.     

The generality of habitat suitability models: A practical test with two insect groups

For the design and declaration of conservation areas as well as for planning habitat management it is important to quantitatively know the habitat preferences of the focal species. To take into account the requirements of as many species as possible, it would be of great advantage if one would either (i) find one or several species whose habitat requirements cover those of a large number of other species or if one could (ii) identify a common set of habitat parameters that is important for the occurrence of many species. Ideally such common habitat parameters should be easy to measure. Only then they may be of practical value in applied conservation biology.In this study, we compared the habitat preferences of different insect species (grasshoppers, bush crickets, butterflies, moths) in the same region by applying identical methods. To identify common explanatory variables that predict the occurrence probability of these species, we first tested the transferability of the specific ‘species models’ to other species within the same insect group. We tested how well the incidence of one species can be predicted by the occurrence probability of another species. The ‘best’ models within each group were then tested for transferability between the different groups. Additionally, we tested the predictive power of the predictor variable ‘habitat type’ as an easy and often available measure for conservation practice.Although in the different ‘species models’ different key factors determine habitat suitability, some models were successfully transferred and were able to reasonably predict the distribution of other species. The habitat preferences of the burnet moth Zygaena carniolica were particularly well suited for the prediction of suitable habitats for all other species. In addition, the predictor variable ‘habitat type’ played a dominant role in all models. Models using this aggregated predictor variable may well predict suitable habitat for all species.