Corridor Length and Patch Colonization by a Butterfly, Junonia coenia

Abstract: Corridors have been proposed to reduce isolation and increase population persistence in fragmented landscapes, yet little research has evaluated the types of landscapes in which corridors will be most effective. I tested the hypothesis that corridors increase patch colonization by a butterfly, Junonia coenia , regardless of the butterfly's initial distance from a patch. I chose J. coenia because it has been shown to move between patches preferentially through corridors. Individuals were released 16–192 m away from open experimental patches into adjacent open corridors or forest. Neither corridors nor distance had a significant effect on patch colonization, but there was a significant interaction between the presence or absence of corridors and distance. At small distances (16–64 m), J. coenia was more likely to colonize open patches when released within forest than within open corridors, most likely because J. coenia used corridors as habitat. Nevertheless, patch colonization by butterflies released within forest decreased rapidly as distance from patches increased, as predicted by a null model of random movement. Colonization did not change with distance in the corridor, and at long distances (128–192 m), butterflies released in corridors were twice as likely to colonize open patches as those released in forest. These results suggest that one critical factor, interpatch distance, may determine the relative effectiveness of corridors and other landscape configurations, such as stepping stones, in reducing isolation in fragmented landscapes. When distances between patches are short compared to an animal's movement ability, a stepping stone approach may most effectively promote dispersal. Alternatively, the conservation value of corridors is highest relative to other habitat configurations when longer distances separate patches in fragmented landscapes.     

The relative influence of habitat loss and fragmentation: do tropical mammals meet the temperate paradigm?

The relative influence of habitat loss vs. habitat fragmentation per se (the breaking apart of habitat) on species distribution and abundance is a topic of debate. Although some theoretical studies predict a strong negative effect of fragmentation, consensus from empirical studies is that habitat fragmentation has weak effects compared with habitat loss and that these effects are as likely to be positive as negative. However, few empirical investigations of this issue have been conducted on tropical or wide-ranging species that may be strongly influenced by changes in patch size and edge that occur with increasing fragmentation. We tested the relative influence of habitat loss and fragmentation by examining occupancy of forest patches by 20 mid- and large-sized Neotropical mammal species in a fragmented landscape of northern Guatemala. We related patch occupancy of mammals to measures of habitat loss and fragmentation and compared the influence of these two factors while controlling for patch-level variables. Species responded strongly to both fragmentation and loss, and response to fragmentation generally was negative. Our findings support previous assumptions that conservation of large mammals in the tropics will require conservation strategies that go beyond prevention of habitat loss to also consider forest cohesion or other aspects of landscape configuration.     

Modelling dispersal with diffusion and habitat selection: Analytical results for highly fragmented landscapes

Quantifying dispersal is crucial both for understanding ecological population dynamics, and for gaining insight into factors that affect the genetic structure of populations. The role of dispersal becomes pronounced in highly fragmented landscapes inhabited by spatially structured populations. We consider a landscape consisting of a set of habitat patches surrounded by unsuitable matrix, and model dispersal by assuming that the individuals follow a random walk with parameters that may be specific to the habitat type. We allow for spatial variation in patch quality, and account for edge-mediated behavior, the latter meaning that the individuals bias their movement towards the patches when close to an edge between a patch and the matrix. We employ a diffusion approximation of the random walk model to derive analytical expressions for various characteristics of the dispersal process. For example, we derive formulae for the time that an individual is expected to spend in its current patch i, and for the time that it will spend in the matrix, both conditional on the individual hitting next a given patch j before hitting any of the other patches or dying. The analytical formulae are based on the assumptions that the landscape is infinitely large, that the patches are circularly shaped, and that the patches are small compared to interpatch distances. We evaluate the effect of these assumptions by comparing the analytical results to numerical results in a real patch network that violates all of the three assumptions. We then consider a landscape that fulfills the assumptions, and show that in this case the analytical results are in a very good agreement with the numerical results. The results obtained here allow the construction of computationally efficient dispersal models that can be used as components of metapopulation models.     

Ranking individual habitat patches as connectivity providers: Integrating network analysis and patch removal experiments

Here we propose an integrated framework for modeling connectivity that can help ecologists, conservation planners and managers to identify patches that, more than others, contribute to uphold species dispersal and other ecological flows in a landscape context. We elaborate, extend and partly integrate recent network-based approaches for modeling and supporting the management of fragmented landscapes. In doing so, experimental patch removal techniques and network analytical approaches are merged into one integrated modeling framework for assessing the role of individual patches as connectivity providers. In particular, we focus the analyses on the habitat availability metrics PC and IIC and on the network metric Betweenness Centrality. The combination and extension of these metrics jointly assess both the immediate connectivity impacts of the loss of a particular patch and the resulting increased vulnerability of the network to subsequent disruptions. In using the framework to analyze the connectivity of two real landscapes in Madagascar and Catalonia (NE Spain), we suggest a procedure that can be used to rank individual habitat patches and show that the combined metrics reveal relevant and non-redundant information valuable to assert and quantify distinctive connectivity aspects of any given patch in the landscape. Hence, we argue that the proposed framework could facilitate more ecologically informed decision-making in managing fragmented landscapes. Finally, we discuss and highlight some of the advantages, limitations and key differences between the considered metrics.     

Dispersal depression with habitat fragmentation in the bog fritillary butterfly

Habitat fragmentation is expected to impose strong selective pressures on dispersal rates. However, evolutionary responses of dispersal are not self-evident, since various selection pressures act in opposite directions. Here we disentangled the components of dispersal behavior in a metapopulation context using the Virtual Migration model, and we linked their variation to habitat fragmentation in the specialist butterfly Proclossiana eunomia. Our study provided a nearly unique opportunity to study how habitat fragmentation modifies dispersal at the landscape scale, as opposed to microlandscapes or simulation studies. Indeed, we studied the same species in four landscapes with various habitat fragmentation levels, in which large amounts of field data were collected and analyzed using similar methodologies. We showed the existence of quantitative variations in dispersal behavior correlated with increased fragmentation. Dispersal propensity from habitat patches (for a given patch size), and mortality during dispersal (for a given patch connectivity) were lower in more fragmented landscapes. We suggest that these were the consequences of two different evolutionary responses of dispersal behavior at the individual level: (1) when fragmentation increased, the reluctance of individuals to cross habitat patch boundaries also increased; (2) when individuals dispersed, they flew straighter in the matrix, which is the best strategy to improve dispersal success. Such evolutionary responses could generate complex nonlinear patterns of dispersal changes at the metapopulation level according to habitat fragmentation. Due to the small size and increased isolation of habitat patches in fragmented landscapes, overall emigration rate and mortality during dispersal remained high. As a consequence, successful dispersal at the metapopulation scale remained limited. Therefore, to what extent the selection of individuals with a lower dispersal propensity and a higher survival during dispersal is able to limit detrimental effects of habitat fragmentation on dispersal success is unknown, and any conclusion that metapopulations would compensate for them is flawed.     

Fish Responses to Experimental Fragmentation of Seagrass Habitat

Abstract: Understanding the consequences of habitat fragmentation has come mostly from comparisons of patchy and continuous habitats. Because fragmentation is a process, it is most accurately studied by actively fragmenting large patches into multiple smaller patches. We fragmented artificial seagrass habitats and evaluated the impacts of fragmentation on fish abundance and species richness over time (1 day, 1 week, 1 month). Fish assemblages were compared among 4 treatments: control (single, continuous 9‐m² patches); fragmented (single, continuous 9‐m² patches fragmented to 4 discrete 1‐m² patches); prefragmented/patchy (4 discrete 1‐m² patches with the same arrangement as fragmented); and disturbance control (fragmented then immediately restored to continuous 9‐m² patches). Patchy seagrass had lower species richness than actively fragmented seagrass (up to 39% fewer species after 1 week), but species richness in fragmented treatments was similar to controls. Total fish abundance did not vary among treatments and therefore was unaffected by fragmentation, patchiness, or disturbance caused during fragmentation. Patterns in species richness and abundance were consistent 1 day, 1 week, and 1 month after fragmentation. The expected decrease in fish abundance from reduced total seagrass area in fragmented and patchy seagrass appeared to be offset by greater fish density per unit area of seagrass. If fish prefer to live at edges, then the effects of seagrass habitat loss on fish abundance may have been offset by the increase (25%) in seagrass perimeter in fragmented and patchy treatments. Possibly there is some threshold of seagrass patch connectivity below which fish abundances cannot be maintained. The immediate responses of fish to experimental habitat fragmentation provided insights beyond those possible from comparisons of continuous and historically patchy habitat.     

An Experimental Test of Matrix Permeability and Corridor Use by an Endemic Understory Bird

Abstract:  Because of widespread habitat fragmentation, maintenance of landscape connectivity has become a major focus of conservation planning, but empirical tests of animal movement in fragmented landscapes remain scarce. We conducted a translocation experiment to test the relative permeability of three landscape elements (open habitat, shrubby secondary vegetation, and wooded corridors) to movement by the Chucao Tapaculo ( Scelorchilus rubecula ), a forest understory bird endemic to South American temperate rainforest. Forty-one radio-tagged subjects were translocated (individually) to three landscape treatments consisting of small release patches that were either entirely surrounded by open habitat (pasture), entirely surrounded by dense shrubs, or linked to other patches by wooded corridors that were otherwise surrounded by open matrix. The number of days subjects remained in release patches before dispersal (a measure of habitat resistance) was significantly longer for patches surrounded by open habitat than for patches adjoining corridors or surrounded by dense shrubs. These results indicate that open habitat significantly constrains Chucao dispersal, in accord with expectation, but dispersal occurs equally well through wooded corridors and shrub-dominated matrix. Thus, corridor protection or restoration and management of vegetation in the matrix (to encourage animal movement) may be equally feasible alternatives for maintaining connectivity.     

Contribution of Roads to Forest Fragmentation in the Rocky Mountains

The contribution of roads to forest fragmentation has not been adequately analyzed. We quantified fragmentation due to roads in a 30,213-ha section of the Medicine Bow-Routt National Forest in sout heastern Wyoming with several indices of landscape structure using a geographic information system. The number of patches, mean patch area, mean interior area, mean area of edge influence, mean patch perimeter, total perimeter, and mean patch shape identified patch- and edge-related landscape changes. Shannon-Wiener diversity, dominance, contagion, contrast, and angular second moment indicated effects on landscape diversity and texture. Roads added to forest fragmentation more than clearcuts by dissecting large patches into smaller pieces and by converting forest interior habitat into edge habitat. Edge habitat created by roads was 1.54–1.98 times the edge habitat created by clearcuts. The total landscape area affected by clearcuts and roads was 2.5–3.5 times the actual area occupied by these disturbances. Fragmentation due to roads could be minimized if road construction is minimized or rerouted so that its fragmentation effects are reduced. Geographic information system technology can be used to quantify the potential fragmentation effects of individual roads and the cumulative effects of a road network on landscape structure.     

Effects of Patch Size and Type of Coffee Matrix on Ithomiine Butterfly Diversity and Dispersal in Cloud-Forest Fragments

Abstract: Determining the permeability of different types of landscape matrices to animal movement is essential for conserving populations in fragmented landscapes. We evaluated the effects of habitat patch size and matrix type on diversity, isolation, and dispersal of ithomiine butterflies in forest fragments surrounded by coffee agroecosystems in the Colombian Andes. Because ithomiines prefer a shaded understory, we expected the highest diversity and abundance in large fragments surrounded by shade coffee and the lowest in small fragments surrounded by sun coffee. We also thought shade coffee would favor butterfly dispersal and immigration into forest patches. We marked 9675 butterflies of 39 species in 12 forest patches over a year. Microclimate conditions were more similar to the forest interior in the shade‐coffee matrix than in the sun‐coffee matrix, but patch size and matrix type did not affect species richness and abundance in forest fragments. Furthermore, age structure and temporal recruitment patterns of the butterfly community were similar in all fragments, independent of patch size or matrix type. There were no differences in the numbers of butterflies flying in the matrices at two distances from the forest patch, but their behavior differed. Flight in the sun‐coffee matrix was rapid and directional, whereas butterflies in shade‐coffee matrix flew slowly. Seven out of 130 recaptured butterflies immigrated into patches in the shade‐coffee matrix, and one immigrated into a patch surrounded by sun coffee. Although the shade‐coffee matrix facilitated movement in the landscape, sun‐coffee matrix was not impermeable to butterflies. Ithomiines exhibited behavioral plasticity in habitat use and high mobility. These traits favor their persistence in heterogeneous landscapes, opening opportunities for their conservation. Understanding the dynamics and resource requirements of different organisms in rural landscapes is critical for identifying management options that address both animals’ and farmers’ needs.     

Fragments as Islands: a Synthesis of Faunal Responses to Habitat Patchiness

Abstract:  Scientific interest in the impact of habitat fragmentation on biodiversity is increasing, but our understanding of fragmentation is clouded by a lack of appreciation for fundamental similarities and differences across studies representing a wide range of taxa and landscape types. In an effort to synthesize data describing ecological responses of animals to fragmentation across two classes of independent variables (taxonomic group and landscape), we sampled 148 studies of five major faunal groups from the primary literature and analyzed data on 13 variables extracted from those studies. We focused our analyses on three classes of dependent variables (effects of area and isolation on species richness, z values, and nestedness and species composition). Area ranged over more orders of magnitude than isolation and tended to explain more variation in species richness than isolation. There were few matrix or taxon effects on the patterns we investigated, although we did find that sky islands tended to manifest isolation effects on both species richness and nestedness more frequently than other patch types. Sky islands may offer insight into the future of habitat patches fragmented by contemporary habitat loss, and because they show a stronger effect of isolation than other patch types, we suggest that isolation will play an increasing role in the biology of habitat fragments. We use multiple lines of evidence to suggest that our understanding of the role of isolation on community assembly in fragmented landscapes is inadequate. Finally, our observation that consistent taxonomic differences in community patterns were minimal suggests that conservation actions intended to mitigate the negative effects of extinction may have far-reaching effects across taxonomic groups.     

Population viability of the Siberian Tiger in a changing landscape: Going, going and gone?

The Amur tiger (Panthera tigris altaica) is a flagship species of the boreal forest ecosystem in northeastern China and Russia Far East. During the past century, the tiger population has declined sharply from more than 3000 to fewer than 600 individuals, and its habitat has become much smaller and greatly fragmented. Poaching, habitat degradation, habitat loss, and habitat fragmentation have been widely recognized as the primary causes for the observed population decline. Using a population viability analysis tool (RAMAS/GIS), we simulated the effects of poaching, habitat degradation, habitat loss, and habitat fragmentation on the population dynamics and extinction risk of the Amur tiger, and then explored the relative effectiveness of three conservation strategies involving improving habitat quality and establishing movement corridors in China and Russia. A series of controlled simulation experiments were performed based on the current spatial distribution of habitat and field-observed vital rates. Our results showed that the Amur tiger population could be viable for the next 100 years if the current habitat area and quality were well-maintained, with poaching strictly prohibited of the tigers and their main prey species. Poaching and habitat degradation (mainly prey scarcity) had the largest negative impacts on the tiger population persistence. While the effect of habitat loss was also substantial, habitat fragmentation per se had less influence on the long-term fate of the tiger population. However, to sustain the subpopulations in both Russia and China would take much greater conservation efforts. The viability of the Chinese population of tigers would rely heavily on its connectivity with the largest patch on the other side of the border. Improving the habitat quality of small patches only or increasing habitat connectivity through movement corridors alone would not be enough to guarantee the long-term population persistence of the Amur tiger in both Russia and China. The only conservation strategy that allowed for long-term persistence of tigers in both countries required both the improvement of habitat quality and the establishment of a transnational reserve network. Our study provides new insights into the metapopulation dynamics and persistence of the Amur tiger, which should be useful in landscape and conservation planning for protecting the biggest cat species in the world.     

Connectivity in an Agricultural Landscape as Reflected by Interpond Movements of a Freshwater Turtle

Abstract:  Connectivity is a measure of how landscape features facilitate movement and thus is an important factor in species persistence in a fragmented landscape. The scarcity of empirical studies that directly quantify species movement and determine subsequent effects on population density have, however, limited the utility of connectivity measures in conservation planning. We undertook a 4-year study to calculate connectivity based on observed movement rates and movement probabilities for five age-sex classes of painted turtles ( Chrysemys picta ) inhabiting a pond complex in an agricultural landscape in northern Virginia (U.S.A.). We determined which variables influenced connectivity and the relationship between connectivity and subpopulation density. Interpatch distance and quality of habitat patches influenced connectivity but characteristics of the intervening matrix did not. Adult female turtles were more influenced by the habitat quality of recipient ponds than other age-sex classes. The importance of connectivity on spatial population dynamics was most apparent during a drought. Population density and connectivity were low for one pond in a wet year but dramatically increased as other ponds dried. Connectivity is an important component of species persistence in a heterogeneous landscape and is strongly dependent on the movement behavior of the species. Connectivity may reflect active selection or avoidance of particular habitat patches. The influence of habitat quality on connectivity has often been ignored, but our findings highlight its importance. Conservation planners seeking to incorporate connectivity measures into reserve design should not ignore behavior in favor of purely structural estimates of connectivity.     

Influence of habitat quality, population size, patch size, and connectivity on patch-occupancy dynamics of the middle spotted woodpecker

Despite extensive research on the effects of habitat fragmentation, the ecological mechanisms underlying colonization and extinction processes are poorly known, but knowledge of these mechanisms is essential to understanding the distribution and persistence of populations in fragmented habitats. We examined these mechanisms through multiseason occupancy models that elucidated patch-occupancy dynamics of Middle Spotted Woodpeckers (Dendrocopos medius) in northwestern Spain. The number of occupied patches was relatively stable from 2000 to 2010 (15-24% of 101 patches occupied every year) because extinction was balanced by recolonization. Larger and higher quality patches (i.e., higher density of oaks >37 cm dbh [diameter at breast height]) were more likely to be occupied. Habitat quality (i.e., density of large oaks) explained more variation in patch colonization and extinction than did patch size and connectivity, which were both weakly associated with probabilities of turnover. Patches of higher quality were more likely to be colonized than patches of lower quality. Populations in high-quality patches were less likely to become extinct. In addition, extinction in a patch was strongly associated with local population size but not with patch size, which means the latter may not be a good surrogate of population size in assessments of extinction probability. Our results suggest that habitat quality may be a primary driver of patch-occupancy dynamics and may increase the accuracy of models of population survival. We encourage comparisons of competing models that assess occupancy, colonization, and extinction probabilities in a single analytical framework (e.g., dynamic occupancy models) so as to shed light on the association of habitat quality and patch geometry with colonization and extinction processes in different settings and species.     

Effects of Habitat Fragmentation on Effective Dispersal of Florida Scrub‐Jays

Abstract: Studies comparing dispersal in fragmented versus unfragmented landscapes show that habitat fragmentation alters the dispersal behavior of many species. We used two complementary approaches to explore Florida Scrub‐Jay (Aphelocoma c?rulescens) dispersal in relation to landscape fragmentation. First, we compared dispersal distances of color‐marked individuals in intensively monitored continuous and fragmented landscapes. Second, we estimated effective dispersal relative to the degree of fragmentation (as inferred from two landscape indexes: proportion of study site covered with Florida Scrub‐Jay habitat and mean distance to nearest habitat patch within each study site) by comparing genetic isolation‐by‐distance regressions among 13 study sites having a range of landscape structures. Among color‐banded individuals, dispersal distances were greater in fragmented versus continuous landscapes, a result consistent with other studies. Nevertheless, genetic analyses revealed that effective dispersal decreases as the proportion of habitat in the landscape decreases. These results suggest that although individual Florida Scrub‐Jays may disperse farther as fragmentation increases, those that do so are less successful as breeders than those that disperse short distances. Our study highlights the importance of combining observational data with genetic inferences when evaluating the complex biological and life‐history implications of dispersal.     

The value of small size: loss of forest patches and ecological thresholds in southern Madagascar.

Many services generated by forest ecosystems provide essential support for human well-being. However, the vulnerability of these services to environmental change such as forest fragmentation are still poorly understood. We present spatial modeling of the generation of ecosystem services in a human-dominated landscape where forest habitat patches, protected by local taboos, are located in a matrix of cultivated land in southern Madagascar. Two ecosystem services dependent on the forest habitats were addressed: (1) crop pollination services by wild and semidomesticated bees (Apoidea), essential for local crop production of, for example, beans, and (2) seed dispersal services based on the presence of ring-tailed lemurs (Lemur catta). We studied the vulnerability of these ecosystem services to a plausible scenario of successive destruction of the smallest habitat patches. Our results indicate that, in spite of the fragmented nature of the landscape, the fraction of the landscape presently covered by both crop pollination and seed dispersal services is surprisingly high. It seems that the taboo system, though indirectly and unintentionally, contributes to upholding the generation of these services by protecting the forest patches. Both services are, however, predicted to be very vulnerable to the successive removal of small patches. For crop pollination, the rate of decrease in cover was significant even when only the smallest habitat patches were removed. The capacity for seed dispersal across the landscape displayed several thresholds with habitat patch removal. Our results suggest that, in order to maintain capacity for seed dispersal across the landscape and crop pollination cover in southern Androy, the geographical location of the remaining forest patches is more crucial than their size. We argue that in heavily fragmented production landscapes, small forest patches should increasingly be viewed as essential for maintaining ecosystem services, such as agricultural production, and also should be considered in the ongoing process of tripling the area of protected habitats in Madagascar.     

Using network centrality measures to manage landscape connectivity

We use a graph-theoretical landscape modeling approach to investigate how to identify central patches in the landscape as well as how these central patches influence (1) organism movement within the local neighborhood and (2) the dispersal of organisms beyond the local neighborhood. Organism movements were theoretically estimated based on the spatial configuration of the habitat patches in the studied landscape. We find that centrality depends on the way the graph-theoretical model of habitat patches is constructed, although even the simplest network representation, not taking strength and directionality of potential organisms flows into account, still provides a coarse-grained assessment of the most important patches according to their contribution to landscape connectivity. Moreover, we identify (at least) two general classes of centrality. One accounts for the local flow of organisms in the neighborhood of a patch, and the other accounts for the ability to maintain connectivity beyond the scale of the local neighborhood. Finally, we study how habitat patches with high scores on different network centrality measures are distributed in a fragmented agricultural landscape in Madagascar. Results show that patches with high degree and betweenness centrality are widely spread, while patches with high subgraph and closeness centrality are clumped together in dense clusters. This finding may enable multispecies analyses of single-species network models.     

Genes and song: genetic and social connections in fragmented habitat in a woodland bird with limited dispersal

Understanding the processes leading to population declines in fragmented landscapes is essential for successful conservation management. However, isolating the influence of disparate processes, and dispersal in particular, is challenging. The Grey Shrike-thrush, Colluricincla harmonica, is a sedentary woodland-dependent songbird, with learned vocalizations whose incidence in suitable habitat patches falls disproportionally with decline in tree cover in the landscape. Although it has been suggested that gaps in tree cover might act as barriers to its dispersal, the species remains in many remnants of native vegetation in agricultural landscapes, suggesting that it may have responded to habitat removal and fragmentation by maintaining or even increasing dispersal distances. We quantified population connectivity of the Grey Shrike-thrush in a system fragmented over more than 120 years using genetic (microsatellites) and acoustic (song types) data. First, we tested for population genetic and acoustic structure at regional and local scales in search of barriers to dispersal or gene flow and signals of local spatial structuring indicative of restricted dispersal or localized acoustic similarity. Then we tested for effects of habitat loss and fragmentation on genetic and acoustic connectivity by fitting alternative models of mobility (isolation-by-distance [the null model] and reduced and increased movement models) across treeless vs. treed areas. Birds within -5 km of each other had more similar genotypes and song types than those farther away, suggesting that dispersal and song matching are limited in the region. Despite restricted dispersal detected for females (but not males), populations appeared to be connected by gene flow and displayed some cultural (acoustic) connectivity across the region. Fragmentation did not appear to impact greatly the dispersal of the Grey Shrike-thrush: none of the mobility models fit the genetic distances of males, whereas for females, an isolation-by-distance model could not be rejected in favor of the models of reduced or increased movement through treeless gaps. However, dissimilarities of the song types were more consistent with the model of reduced cultural connectivity through treeless areas, suggesting that fragmentation impedes song type sharing in the Grey Shrike-thrush. Our paper demonstrates that habitat fragmentation hinders important population processes in an Australian woodland bird even though its dispersal is not detectably impacted.     

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.     

Thresholds in Songbird Occurrence in Relation to Landscape Structure

Abstract:  Theory predicts the occurrence of threshold levels of habitat in landscapes, below which ecological processes change abruptly. Simulation models indicate that below critical thresholds, fragmentation of habitat influences patch occupancy by decreasing colonization rates and increasing rates of local extinction. Uncovering such putative relationships is important for understanding the demography of species and in developing sound conservation strategies. Using segmented logistic regression, we tested for thresholds in occurrence of 15 bird species as a function of the amount of suitable habitat at multiple scales (150–2000-m radii). Suitable habitat was defined quantitatively based on previously derived, spatially explicit distribution models for each species. The occurrence of 10 out of 15 species was influenced by the amount of habitat at a landscape scale (≥500-m radius). Of these species all but one were best predicted by threshold models. Six out of nine species exhibited asymptotic thresholds; the effects of habitat loss intensified at low amounts of habitat in a landscape. Landscape thresholds ranged from 8.6% habitat to 28.7% (     = 18.5 ± 2.6%[95% CI]). For two species landscape thresholds coincided with sensitivity to fragmentation; both species were more likely to occur in large patches, but only when the amount of habitat in a landscape was low. This supports the fragmentation threshold hypothesis. Nevertheless, the occurrence of most species appeared to be unaffected by fragmentation, regardless of the amount of habitat present at landscape extents. The thresholds we identified may be useful to managers in establishing conservation targets. Our results indicate that findings of landscape-scale studies conducted in regions with relatively high proportions of habitat and low fragmentation may not be applicable in regions with low habitat proportions and high fragmentation.