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.     

Independent effects of fragmentation on forest songbirds: an organism-based approach.

The degree to which spatial patterns influence the dynamics and distribution of populations is a central question in ecology. This question is even more pressing in the context of rapid habitat loss and fragmentation, which threaten global biodiversity. However, the relative influence of habitat loss and landscape fragmentation, the spatial patterning of remaining habitat, remains unclear. If landscape pattern affects population size, managers may be able to design landscapes that mitigate habitat loss. We present the results of a mensurative experiment designed to test four habitat loss vs. fragmentation hypotheses. Unlike previous studies, we measured landscape structure using quantitative, spatially explicit habitat distribution models previously developed for two species: Blackburnian Warbler (Dendroica fusca) and Ovenbird (Seiurus aurocapilla). We used a stratified sampling design that reduced the confounding of habitat amount and fragmentation variables. Occurrence and reoccurrence of both species were strongly influenced by characteristics at scales greater than the individual territory, indicating little support for the random-sample hypothesis. However, the type and spatial extent of landscape influence differed. Both occurrence and reoccurrence of Blackburnian Warblers were influenced by the amount of poor-quality matrix at 300- and 2000-m spatial extents. The occurrence and reoccurrence of Ovenbirds depended on a landscape pattern variable, patch size, but only in cases when patches were isolated. These results support the hypothesis that landscape pattern is important for some species only when the amount of suitable habitat is low. Although theoretical models have predicted such an interaction between landscape fragmentation and composition, to our knowledge this is the first study to report empirical evidence of such nonlinear fragmentation effects. Defining landscapes quantitatively from an organism-based perspective may increase power to detect fragmentation effects, particularly in forest mosaics where boundaries between patches and matrix are ambiguous. Our results indicate that manipulating landscape pattern may reduce negative impacts of habitat loss for Ovenbird, but not Blackburnian Warbler. We emphasize that most variance in the occurrence of both species was explained by local scale or landscape composition variables rather than variables reflecting landscape pattern.     

Conservation of Fragmented Populations

In this paper we argue that landscape spatial structure is of central importance in understanding the effects of fragmentation on population survival. Landscape spatial structure is the spatial relationships among habitat patches and the matrix in which they are embedded. Many general models of subdivided populations make the assumptions that (1) all habitat patches are equivalent in size and quality and (2) all local populations (in the patches) are equally accessible by dispersers. Models that gloss over spatial details of landscape structure can be useful for theoretical developments but will almost always be misleading when applied to real-world conservation problems. We show that local extinctions of fragmented populations are common. From this it follows that recolonization of local extinctions is critical for regional survival of fragmented populations. The probability of recolonization depends on (1) spatial relationships among landscape elements used by the population, including habitat patches for breeding and elements of the inter-patch matrix through which dispersers move, (2) dispersal characteristics of the organism of interest, and (3) temporal changes in the landscape structure. For endangered species, which are typically restricted in their dispersal range and in the kinds of habitat through which they can disperse, these factors are of primary importance and must be explicitly considered in management decisions.     

Estimating Extinction Risk with Metapopulation Models of Large‐Scale Fragmentation

Habitat loss is the principal threat to species. How much habitat remains—and how quickly it is shrinking—are implicitly included in the way the International Union for Conservation of Nature determines a species’ risk of extinction. Many endangered species have habitats that are also fragmented to different extents. Thus, ideally, fragmentation should be quantified in a standard way in risk assessments. Although mapping fragmentation from satellite imagery is easy, efficient techniques for relating maps of remaining habitat to extinction risk are few. Purely spatial metrics from landscape ecology are hard to interpret and do not address extinction directly. Spatially explicit metapopulation models link fragmentation to extinction risk, but standard models work only at small scales. Counterintuitively, these models predict that a species in a large, contiguous habitat will fare worse than one in 2 tiny patches. This occurs because although the species in the large, contiguous habitat has a low probability of extinction, recolonization cannot occur if there are no other patches to provide colonists for a rescue effect. For 4 ecologically comparable bird species of the North Central American highland forests, we devised metapopulation models with area‐weighted self‐colonization terms; this reflected repopulation of a patch from a remnant of individuals that survived an adverse event. Use of this term gives extra weight to a patch in its own rescue effect. Species assigned least risk status were comparable in long‐term extinction risk with those ranked as threatened. This finding suggests that fragmentation has had a substantial negative effect on them that is not accounted for in their Red List category. Estimación del Riesgo de Extinción Mediante Modelos Metapoblacionales de Fragmentación a Gran Escala     

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.     

Inferring Metapopulation Dynamics from Patch-Level Incidence of Florida Scrub Plants

Spatial structure and dynamics of multiple populations may explain species distribution patterns in patchy communities with heterogeneous disturbance regimes, especially when species have poor dispersal. The endemic-rich Florida (U.S.A.) rosemary scrub occupies about 4% of the west portion of Archbold Biological Station and occurs scattered within a matrix of less xeric vegetation. Longer fire-return times and higher frequency of open patches in rosemary scrub provide favorable habitat for many plant species. Occupancy of 123 species of vascular plants and ground lichens in 89 patches was determined by repeated site surveys. About two-thirds of the species occurring at more than 14 patches had a significant logistic regression of presence on time-since-fire, patch size, patch isolation, or their interactions. Species with presence related to the interaction between patch isolation and patch size were primarily herbs and small shrubs specializing in rosemary scrub. These results suggest the importance of spatial characteristics of the landscape for population turnover of these species. An incidence-based metapopulation model was used to predict extinction and colonization probabilities of those species with presence in rosemary scrub patches related to the studied spatial variables. This is the first attempt to apply incidence-based metapopulation models to plants. The results showed stronger effects of patch size and patch isolation on extinction probabilities of herbs than on those of woody species. Because of their effect on spatial heterogeneity and habitat availability, fire suppression and habitat destruction may decrease persistence probabilities for these rosemary scrub specialists, many of which are endangered species.     

Validating graph-based connectivity models with independent presence–absence and genetic data sets

Habitat connectivity is a key objective of current conservation policies and is commonly modeled by landscape graphs (i.e., sets of habitat patches [nodes] connected by potential dispersal paths [links]). These graphs are often built based on expert opinion or species distribution models (SDMs) and therefore lack empirical validation from data more closely reflecting functional connectivity. Accordingly, we tested whether landscape graphs reflect how habitat connectivity influences gene flow, which is one of the main ecoevolutionary processes. To that purpose, we modeled the habitat network of a forest bird (plumbeous warbler [Setophaga plumbea]) on Guadeloupe with graphs based on expert opinion, Jacobs’ specialization indices, and an SDM. We used genetic data (712 birds from 27 populations) to compute local genetic indices and pairwise genetic distances. Finally, we assessed the relationships between genetic distances or indices and cost distances or connectivity metrics with maximum-likelihood population-effects distance models and Spearman correlations between metrics. Overall, the landscape graphs reliably reflected the influence of connectivity on population genetic structure; validation R² was up to 0.30 and correlation coefficients were up to 0.71. Yet, the relationship among graph ecological relevance, data requirements, and construction and analysis methods was not straightforward because the graph based on the most complex construction method (species distribution modeling) sometimes had less ecological relevance than the others. Cross-validation methods and sensitivity analyzes allowed us to make the advantages and limitations of each construction method spatially explicit. We confirmed the relevance of landscape graphs for conservation modeling but recommend a case-specific consideration of the cost-effectiveness of their construction methods. We hope the replication of independent validation approaches across species and landscapes will strengthen the ecological relevance of connectivity models.     

Effects of the spatial pattern of disturbance on the patch-occupancy dynamics of juniper–pine open woodland

Typically, studies of the disturbance effect on metapopulation dynamics are limited to understanding the effect of habitat loss although, recently, the spatial pattern of the disturbance has been shown to influence dynamics. In this study, we used a stochastic patch-dynamic model to investigate the effects of spatial disturbance patterns on the persistence of an open woodland community of Juniperus spp. and Pinus spp. First, we estimated patch-occupancy dynamics by using the coefficients that best predicted the occupancy observed in 1998 based on occupancy data from 1957. Next, we evaluated the effects of the rate and pattern of the disturbance on the extinction probability. In modeling the disturbance, we considered (1) the degree of disturbance produced by scenarios of complete destruction or degradation (with the potential for recolonization), (2) the overall rate of disturbance, and (3) the spatial autocorrelation of habitat destruction. Twenty 40-year simulations predicted a 25% increase in the number of patches, and when 50% of the habitat was removed, the impact was more pronounced after complete destruction than it was after degradation of the area. Predictions based on scenarios of complete destruction, including random, contiguous, Brownian, and autoregressive noise, demonstrated that the impact of disturbance depends upon the spatial structure of the disturbance regimen. The autocorrelated structure of the disturbance regimen had the greatest impact on patch persistence. Patch-occupancy was higher after 20 40-year simulations when habitat loss was randomly distributed than when it followed an autocorrelated patch destruction, which was simulated using autoregressive noise to produce 50% habitat destruction. In addition, while habitat loss was negatively linearly correlated with patch persistence when habitat destruction was randomly distributed, a dramatic transition shift occurred when habitat destruction was simulated following an autoregressive spatial distribution after a certain threshold of habitat destruction (40% of the actual open woodland habitat). Our study suggests that the spatial patterns of the disturbance should be considered when predicting the consequences of fragmentation and improving management strategies.     

Metapopulation Extinction Risk under Spatially Autocorrelated Disturbance

Abstract:  Recent extinction models generally show that spatial aggregation of habitat reduces overall extinction risk because sites emptied by local extinction are more rapidly recolonized. We extended such an investigation to include spatial structure in the disturbance regime. A spatially explicit metapopulation model was developed with a wide range of dispersal distances. The degree of aggregation of both habitat and disturbance pattern could be varied from a random distribution, through the intermediate case of a fractal distribution, all the way to complete aggregation (single block). Increasing spatial aggregation of disturbance generally increased extinction risk. The relative risk faced by populations in different landscapes varied greatly, depending on the disturbance regime. With random disturbance, the spatial aggregation of habitat reduced extinction risk, as in earlier studies. Where disturbance was spatially autocorrelated, however, this advantage was eliminated or reversed because populations in aggregated habitats are at risk of mass extinction from coarse-scale disturbance events. The effects of spatial patterns on extinction risk tended to be reduced by long-distance dispersal. Given the high levels of spatial correlation in natural and anthropogenic disturbance processes, population vulnerability may be greatly underestimated both by classical (nonspatial) models and by those that consider spatial structure in habitat alone.     

Animal dispersal modelling: Handling landscape features and related animal choices

Animal dispersal in a fragmented landscape depends on the complex interaction between landscape structure and animal behavior. To better understand how individuals disperse, it is important to explicitly represent the properties of organisms and the landscape in which they move. A common approach to modelling dispersal includes representing the landscape as a grid of equal sized cells and then simulating individual movement as a correlated random walk. This approach uses a priori scale of resolution, which limits the representation of all landscape features and how different dispersal abilities are modelled.We develop a vector-based landscape model coupled with an object-oriented model for animal dispersal. In this spatially explicit dispersal model, landscape features are defined based on their geographic and thematic properties and dispersal is modelled through consideration of an organism's behavior, movement rules and searching strategies (such as visual cues). We present the model's underlying concepts, its ability to adequately represent landscape features and provide simulation of dispersal according to different dispersal abilities. We demonstrate the potential of the model by simulating two virtual species in a real Swiss landscape. This illustrates the model's ability to simulate complex dispersal processes and provides information about dispersal such as colonization probability and spatial distribution of the organism's path.     

Use of linkage mapping and centrality analysis across habitat gradients to conserve connectivity of gray wolf populations in western North America

Centrality metrics evaluate paths between all possible pairwise combinations of sites on a landscape to rank the contribution of each site to facilitating ecological flows across the network of sites. Computational advances now allow application of centrality metrics to landscapes represented as continuous gradients of habitat quality. This avoids the binary classification of landscapes into patch and matrix required by patch-based graph analyses of connectivity. It also avoids the focus on delineating paths between individual pairs of core areas characteristic of most corridor- or linkage-mapping methods of connectivity analysis. Conservation of regional habitat connectivity has the potential to facilitate recovery of the gray wolf (Canis lupus), a species currently recolonizing portions of its historic range in the western United States. We applied 3 contrasting linkage-mapping methods (shortest path, current flow, and minimum-cost-maximum-flow) to spatial data representing wolf habitat to analyze connectivity between wolf populations in central Idaho and Yellowstone National Park (Wyoming). We then applied 3 analogous betweenness centrality metrics to analyze connectivity of wolf habitat throughout the northwestern United States and southwestern Canada to determine where it might be possible to facilitate range expansion and interpopulation dispersal. We developed software to facilitate application of centrality metrics. Shortest-path betweenness centrality identified a minimal network of linkages analogous to those identified by least-cost-path corridor mapping. Current flow and minimum-cost-maximum-flow betweenness centrality identified diffuse networks that included alternative linkages, which will allow greater flexibility in planning. Minimum-cost-maximum-flow betweenness centrality, by integrating both land cost and habitat capacity, allows connectivity to be considered within planning processes that seek to maximize species protection at minimum cost. Centrality analysis is relevant to conservation and landscape genetics at a range of spatial extents, but it may be most broadly applicable within single- and multispecies planning efforts to conserve regional habitat connectivity.     

Modeling the spatio-temporal dynamics and interactions of households,landscapes, and giant panda habitat

Human modification of land-cover has been a leading cause of floral and faunal species extirpation and loss of local and global biodiversity. As natural areas are impacted, habitat and populations can become fragmented and isolated. This is particularly evident in the mountainous areas of southwestern China that support the remaining populations of giant pandas (Ailuropoda melanoleuca). Giant panda populations have been restricted to remnants of habitat from extensive past land use and land-cover change. Households are a basic socio-economic unit that continues to impact the remaining habitat through activities such as fuelwood consumption and new household creation. Therefore, we developed a spatio-temporal model of human activities and their impacts by directly integrating households into the landscape. The integrated model allows us to examine the landscape factors influencing the spatial distribution of household activities and household impacts on habitat. As an example application, we modeled household activities in a giant panda reserve in China and examined the spatio-temporal dynamics of households, the landscape, and their mutual interactions. Human impacts are projected to result in the loss of up to 16% of all existing habitat within the reserve over the next 30 years. In addition, we found that accessibility largely controls the spatial distribution of household activities and considerable changes in management and household activities will be required to maintain the current level of habitat within the reserve.     

Appreciating Ecological Complexity: Habitat Contours as a Conceptual Landscape Model

Abstract:  Organisms respond to their surroundings at multiple spatial scales, and different organisms respond differently to the same environment. Existing landscape models, such as the "fragmentation model" (or patch-matrix-corridor model) and the "variegation model," can be limited in their ability to explain complex patterns for different species and across multiple scales. An alternative approach is to conceptualize landscapes as overlaid species-specific habitat contour maps. Key characteristics of this approach are that different species may respond differently to the same environmental conditions and at different spatial scales. Although similar approaches are being used in ecological modeling, there is much room for habitat contours as a useful conceptual tool. By providing an alternative view of landscapes, a contour model may stimulate more field investigations stratified on the basis of ecological variables other than human-defined patches and patch boundaries. A conceptual model of habitat contours may also help to communicate ecological complexity to land managers. Finally, by incorporating additional ecological complexity, a conceptual model based on habitat contours may help to bridge the perceived gap between pattern and process in landscape ecology. Habitat contours do not preclude the use of existing landscape models and should be seen as a complementary approach most suited to heterogeneous human-modified landscapes.     

Modeling Species' Distributions to Improve Conservation in Semiurban Landscapes: Koala Case Study

Abstract:  Models of species' distributions are commonly used to inform landscape and conservation planning. In urban and semiurban landscapes, the distributions of species are determined by a combination of natural habitat and anthropogenic impacts. Understanding the spatial influence of these two processes is crucial for making spatially explicit decisions about conservation actions. We present a logistic regression model for the distribution of koalas (  Phascolarctos cinereus ) in a semiurban landscape in eastern Australia that explicitly separates the effect of natural habitat quality and anthropogenic impacts on koala distributions. We achieved this by comparing the predicted distributions from the model with what the predicted distributions would have been if anthropogenic variables were at their mean values. Similar approaches have relied on making predictions assuming anthropogenic variables are zero, which will be unreliable if the training data set does not include anthropogenic variables close to zero. Our approach is novel because it can be applied to landscapes where anthropogenic variables are never close to zero. Our model showed that, averaged across the study area, natural habitat was the main determinant of koala presence. At a local scale, however, anthropogenic impacts could be more important, with consequent implications for conservation planning. We demonstrated that this modeling approach, combined with the visual presentation of predictions as a map, provides important information for making decisions on how different conservation actions should be spatially allocated. This method is particularly useful for areas where wildlife and human populations exist in close proximity.     

Interactive effects of natural and human disturbances on vegetation dynamics across landscapes.

Accurate measures of human effects on landscape processes require consideration of both the direct impacts from human activities and the indirect consequences of the interactions between humans and the landscape. This is particularly evident in systems experiencing regular natural disturbances such as in the mountainous areas of southwestern China, where the remaining population of giant pandas (Ailuropoda melanoleuca) is supported. Here the spatiotemporal patterns of human impacts, forests, and bamboo episodic die-offs combine to determine the distribution of panda habitat. To study the complex interactions of humans and landscapes, we developed an integrated spatiotemporally explicit model of household activities, natural vegetation dynamics, and their impacts on panda habitat. Using this model we examined the direct consequences of local fuelwood collection and household creation on areas of critical giant panda habitat and the indirect impacts when coupled with vegetation dynamics. Through simulations, we found that over the next 30 years household impacts would result in the loss of up to 30% of the habitat relied on by pandas during past bamboo die-offs. The accumulation and spatial distribution of household impacts would also have a considerable indirect influence on the spatial distribution of understory bamboo. While human impacts influence both bamboo die-off and regeneration, over 19% of pre-existing low-elevation bamboo habitat may be lost following an episodic die-off depending on the severity of the impacts and timing of the die-offs. Our study showed not only the importance of the spatial distribution of direct household impacts on habitat, but also the far-reaching effects of the indirect interactions between humans and the landscapes they are modifying.     

Extinction, Colonization, and Metapopulations: Environmental Tracking by Rare Species

Extinction and metapopulation theories emphasize that stochastic fluctuations in local populations cause extinction and that local extinctions generate empty habitat patches that are then available for recolonization. Metapopulation persistence depends on the balance of extinction and colonization in a static environment. For many rare and declining species, I argue (1) that extinction is usually the deterministic consequence of the local environment becoming unsuitable (through habitat loss or modification, introduction of a predator, etc.); (2) that the local environment usually remains unsuitable following local extinction, so extinctions only rarely generate empty patches of suitable habitat; and (3) that colonization usually follows improvement of the local environment for a particular species (or long-distance transfer by humans). Thus, persistence depends predominantly on whether organisms are able to track the shifting spatial mosaic of suitable environmental conditions or on maintainance of good conditions locally.     

Landscape bioacoustics allow detection of the effects of habitat patchiness on population structure

Landscape structure may affect individual dispersal abilities, thus influencing the genotypic and phenotypic composition of populations. We analyzed the interplay among landscape, behavior, and evolutionary processes by correlating habitat patchiness to the variability in vocalizations of Dupont's Lark Chersophilus duponti, one of the most habitat-selective and rare European songbirds. We tape-recorded males throughout the species distribution in Spain, analyzed the spatial patterns of territorial call variation at different scales (individuals, populations, and broad geographic areas), and related acoustic variability to patterns of isolation by geographic distance and by landscape unsuitability (calculated by building a predictive model of habitat suitability). The differentiation of spectro-temporal call features resulted from both isolation by distance and isolation by landscape unsuitability mechanisms. Landscape connectivity was often a better determinant of call differentiation than simple straight-line distance between individuals, providing the first evidence that call transmission can be limited by the presence and distribution of patches of adequate habitat, which likely mediates bird dispersal. Landscape patchiness resulted in a reduction of acoustic diversity (repertoire size) within populations, and a parallel increase in differentiation among populations. Landscape bioacoustics can represent a promising tool for estimating population structure, although the study of animal communication cannot be viewed as an alternative, but a source of complementary information to genetics, given that it provides evidence of male-male transmission and social and cultural phenomena that are currently undetectable from molecular data.     

A Framework for Conceptualizing Human Effects on Landscapes and Its Relevance to Management and Research Models

Abstract: The concept of habitat fragmentation is limited in its ability to describe the range of possible landscape configurations created by a variety of disturbances. This limitation is especially problematic in landscapes where human use of the habitat matrix occurs at multiple levels and where habitat modification may be a more important consideration than a simple binary classification of habitat versus nonhabitat. We propose a synthesizing scheme that places intact, variegated, fragmented, and relictual landscape states on a continuum, depending on the degree of habitat destruction. At a second level, the scheme considers the patterns of habitat modification that are imposed on remaining habitats. Management for conservation involves halting and sometimes reversing the trends of habitat destruction and modification. Conservation strategies will differ according to the state of alteration of the landscape, but all strategies include some consideration of the degree of modification of the matrix in determining habitat viability. It is convenient for biologists to assess landscape alteration state in terms of the persistence of large structural elements such as trees. Because animal species use habitats differently, however, they also experience the landscape differently. A landscape considered structurally fragmented by humans may be functionally variegated to other species. Therefore, it is necessary to consider the extent to which the entire landscape, including the matrix, is accessible and utilized by organisms with different spatial scales of resource use.     

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.