Response of Red-Backed Voles to Recent Patch Cutting in Subalpine Forest

We examined the response of southern red-backed voles ( Clethrionomys gapperi ) to patch cutting in a forested landscape in which 23% of the forest cover had been removed by timber harvest. We live trapped voles in and around 18 patchcuts in one watershed of southern Wyoming. Although we found a significant difference in capture rates between patchcut interior and forest habitats in 1 of 2 years, voles did not strongly avoid the interior of patchcuts. This result contrasts with results from most studies of voles in clearcut ecosystems, which report that red-backed voles are generally rare or absent from clearcuts. Capture rates were highest on both sides of the patchcut edge, which also contrasts with studies of voles at the edges of forest remnants. The use of patchcut interior and edge habitats could not be explained as a consequence of juvenile voles dispersing to those habitats or males moving through the habitat in search of mates. We suggest that, despite similar physiognomy in patchcut and clearcut sites, the differences in landscape structure in perforated versus fragmented landscapes lead to very different patterns of vole movement. Understanding the scales at which voles perceive landscapes as coarse- or fine-grained will be key to developing predictive models to aid managers in designing timber sales that maintain high vole populations. Our results emphasize the importance of the spatial pattern and scale of disturbance in determining the response of vertebrates to landscape change and the need for more refined investigations of the consequences of deforestation.     

Evaluating the Effectiveness of Corridors: a Genetic Approach

Abstract: The effectiveness of corridors in maintaining dispersal in fragmented landscapes is a question of considerable conservation and ecological importance. We tested the efficacy of corridors as residual landscape structures in maintaining population structure in the red-backed vole ( Clethrionomys gapperi ), a closed-canopy specialist, and the deer mouse (   Peromyscus maniculatus ), a habitat generalist. In coniferous forests managed for timber production in northeastern Washington, we sampled pairs of populations in three landscape classes: (1) contiguous landscapes, in which sites were located completely within a matrix of closed-canopy forest; (2) corridor landscapes, in which sites were connected by a corridor of closed-canopy forest; and (3) isolated landscapes, in which sites were separated from one another by clearcut or young regeneration stands. For each species, we used four microsatellite loci to quantify genetic distance between population pairs. Nei's genetic distance (   D _s  ) increased from smallest to largest in the order of contiguous, corridor, and isolated landscapes for C. gapperi. For P. maniculatus, genetic distances across landscape configurations were not significantly different. The differences between the two species indicate that they respond differently to the presence of forest corridors. In managed forests, corridors between unlogged habitats appear to maintain higher population connectivity for C. gapperi than landscapes without corridors.     

Persistence of Forest Birds in the Costa Rican Agricultural Countryside

Abstract:  Understanding the persistence mechanisms of tropical forest species in human-dominated landscapes is a fundamental challenge of tropical ecology and conservation. Many species, including more than half of Costa Rica's native land birds, use mostly deforested agricultural countryside, but how they do so is poorly known. Do they commute regularly to forest or can some species survive in this human-dominated landscape year-round? Using radiotelemetry, we detailed the habitat use, movement, foraging, and nesting patterns of three bird species, Catharus aurantiirostris , Tangara icterocephala , and Turdus assimilis , by obtaining 8101 locations from 156 individuals. We chose forest birds that varied in their vulnerability to deforestation and were representative of the species found both in forest and human-dominated landscapes. Our study species did not commute from extensive forest; rather, they fed and bred in the agricultural countryside. Nevertheless, T. icterocephala and T. assimilis , which are more habitat sensitive, were highly dependent on the remaining trees. Although trees constituted only 11% of land cover, these birds spent 69% to 85% of their time in them. Breeding success of C. aurantiirostris and T. icterocephala in deforested habitats was not different than in forest remnants, where T. assimilis experienced reduced breeding success. Although this suggests an ecological trap for T. assimilis , higher fledgling survival in forest remnants may make up for lower productivity. Tropical countryside has high potential conservation value, which can be enhanced with even modest increases in tree cover. Our findings have applicability to many human-dominated tropical areas that have the potential to conserve substantial biodiversity if appropriate restoration measures are taken.     

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.     

Extraterritorial Movements of a Forest Songbird in a Fragmented Landscape

Abstract: Forest isolation resulting from fragmentation is thought to impede the movement of forest songbirds. Because of the difficulty of tracking birds continuously, however, few data exist documenting the influence of isolation and landscape features on avian movements. During the breeding season, male Hooded Warblers (    Wilsonia citrina ) leave their small (<2.5 ha), isolated forest patches to travel between forest fragments. We documented a total of 106 forays (  n = 20 males) and found that individuals traveled up to 2.5 km away from their resident forest patch, primarily to solicit covert extra-pair copulations. Forays occurred despite the absence of forested corridors connecting fragments; even when corridors were present, males most often chose to fly directly across open fields. Resident patch size and distance to forests visited were not correlated with the frequency of forays. The maximum distance males flew over open fields did not exceed 465 m, and longer distances likely inhibit males from traveling outside their woodlots. If territorial establishment depends on the availability of extra-pair partners, then higher degrees of isolation between forests could explain why some species avoid settling in extremely fragmented landscapes. Conservation efforts should limit isolation between forest stands, thereby preserving the ability of animals to move within fragmented landscapes during the breeding season.     

The Effect of Fragment Shape and Species' Sensitivity to Habitat Edges on Animal Population Size

Abstract:  Habitat fragmentation causes extinction of local animal populations by decreasing the amount of viable "core" habitat area and increasing edge effects. It is widely accepted that larger fragments make better nature reserves because core-dwelling species have a larger amount of suitable habitat. Nevertheless, fragments in real landscapes have complex, irregular shapes. We modeled the population sizes of species that have a representative range of preferences for or aversions to habitat edges at five spatial scales (within 10, 32, 100, 320, and 1000 m of an edge) in a nation-wide analysis of forest remnants in New Zealand. We hypothesized that the irregular shapes of fragments in real landscapes should generate statistically significant correlations between population density and fragment area, purely as a "geometric" effect of varying species responses to the distribution of edge habitat. Irregularly shaped fragments consistently reduced the population size of core-dwelling species by 10–100%, depending on the scale over which species responded to habitat edges. Moreover, core populations within individual fragments were spatially discontinuous, containing multiple, disjunct populations that inhabited small spatial areas and had reduced population size. The geometric effect was highly nonlinear and depended on the range of fragment sizes sampled and the scale at which species responded to habitat edges. Fragment shape played a strong role in determining population size in fragmented landscapes; thus, habitat restoration efforts may be more effective if they focus on connecting disjunct cores rather than isolated fragments.     

Nesting Success of Neotropical Migrant Songbirds in a Highly Fragmented Landscape

Abstract: Small forests in agricultural landscapes are generally thought to be population sinks where the nesting success of Neotropical migrant songbirds is too low to sustain populations. In 1996 and 1997, we assessed the nesting success and productivity of Wood Thrushes (  Hylocichla mustelina ) and Rose-breasted Grosbeaks (  Pheucticus ludovicianus ) in the Region of Waterloo, a highly fragmented agricultural landscape with 14% forest cover in southwestern Ontario. We located 154 Wood Thrush nests and 63 Rose-breasted Grosbeak nests in 26 small forest fragments (3–14 ha) and 15 large fragments (26–140 ha). We used the Mayfield method to determine nesting success. Across all sites nesting success was 51% for Wood Thrushes and 46% for Rose-breasted Grosbeaks. Nesting success and productivity rates were used in a simple population growth model that suggested that the Wood Thrushes but not the Rose-breasted Grosbeaks were a self-sustaining population. Forest size and distance of nests to the nearest forest edge (measured in five distance classes ranging from 0 to over 100 m) did not significantly affect the nesting success or productivity of either species. Brown-headed Cowbirds (   Molothrus ater ) parasitized 47% of all Wood Thrush nests and 10% of all Rose-breasted Grosbeak nests; parasitism significantly reduced Wood Thrush productivity but not that of Rose-breasted Grosbeaks. Our findings challenge the prevailing notion that small woodlots in a farmed landscape are invariably habitat sinks and offer additional incentive for habitat protection in settled landscapes where small forest fragments are often all that remain for conservation purposes.     

Spillover of Agriculturally Subsidized Predators as a Potential Threat to Native Insect Herbivores in Fragmented Landscapes

Abstract:  Habitat loss and fragmentation can have strong negative impacts on populations of some native species. Spillover of generalist natural enemies from the surrounding landscape matrix is one mechanism potentially generating such effects, yet this has been rarely studied in insects. We examined the influence of habitat conversion to agriculture on the abundance and potential effects of predatory coccinellid beetles on native insect herbivores within 12 grassland remnants in central Nebraska (U.S.A.). Results of sweep sampling revealed that coccinellids were three to six times more abundant at native grassland sites embedded within cropland-dominated landscapes compared with control sites in grassland-dominated landscapes over the 3 years of the study. Exclusion experiments further demonstrated that predation intensity was strongly related to coccinellid abundances across sites and that coccinellids can dramatically reduce densities of a native aphid herbivore. In contrast to studies of specialized insect parasitoids, which have generally found reduced enemy pressure in fragmented landscapes, our results suggest that native herbivores may in some cases experience increased consumer pressure in landscapes with increasing habitat loss because of spillover of generalist predators from surrounding cropland habitats.     

Managing Boreal Forest Landscapes for Flying Squirrels

Abstract: Flying squirrel (Pteromys volans) populations have declined severely during the past few decades, and the species has become a focal species in forest management and the conservation debate in Finland. We compared landscape structure around known flying squirrel home ranges with randomly chosen forest sites to determine which landscape patterns characterize the areas occupied by the species in northern Finland. We sought to identify the key characteristics of the landscape that support the remaining flying squirrel populations. We analyzed landscape structure within circular areas with 1- and 3-km radii around 63 forest sites occupied by flying squirrels, and around 96 random sites. We applied stepwise analysis of the landscape structure where landscapes were built up step-by-step by adding patch types in order of their suitability for the flying squirrel. The land-use and forest-resource data for the analysis were derived from multisource national forest inventory and imported to a geographical information system. Landscape patch types were divided into three suitability categories: breeding habitat (mixed spruce-deciduous forests); dispersal habitat ( pine and young forests); and unsuitable habitat ( young sapling stands, open habitats, water). Flying squirrel landscapes contained more suitable breeding habitat patches and were better connected by dispersal habitats than random landscapes. Our results suggest that for the persistence of the flying squirrel, forest managers should 1) maintain a deciduous mixture, particularly in spruce-dominated forests; 2) maintain physical connectivity between optimal breeding habitats; and 3) impose coarse-grained structures on northeastern Finnish landscapes at current levels of habitat availability.     

Demographic Limitations of the Ability of Habitat Restoration to Rescue Declining Populations

Abstract:  Habitat restoration is often recommended in conservation without first evaluating whether populations are in fact habitat limited and thus whether declining populations can be stabilized or recovered through habitat restoration. We used a spatially structured demographic model coupled with a dynamic neutral landscape model to evaluate whether habitat restoration could rescue populations of several generic migratory songbirds that differed in their sensitivity to habitat fragmentation (i.e., severity of edge effects on nesting success). Simulating a best-case scenario, landscapes were instantly restored to 100% habitat before, at, or after habitat loss exceeded the species' vulnerability threshold. The vulnerability threshold is a measure of extinction risk, in which the change in population growth rate ( δλ ) scaled to the rate of habitat loss ( δh ) falls below −1% ( δλ/δh ≤ −0.01). Habitat restoration was most effective for species with low-to-moderate edge sensitivities and in landscapes that had not previously experienced extensive fragmentation. To stabilize populations of species that were highly edge sensitive or any species in heavily fragmented landscapes, restoration needed to be initiated long before the vulnerability threshold was reached. In practice, habitat restoration is generally not initiated until a population is at risk of extinction, but our model results demonstrate that some populations cannot be recovered at this point through habitat restoration alone. At this stage, habitat loss and fragmentation have seriously eroded the species' demographic potential such that halting population declines is limited more by demographic factors than the amount of available habitat. Evidence that populations decline in response to habitat loss is thus not sufficient to conclude that habitat restoration will be sufficient to rescue declining populations.     

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.     

Gap Crossing Decisions by Forest Songbirds during the Post-Fledging Period

Gaps in forest cover, created by agriculture, forestry, and other anthropogenic activities, are assumed to impede the movements of many forest songbirds. Little is known, however, about the reluctance of different species of birds to cross habitat gaps. We studied this by inducing birds in the post-fledging period to cross gaps of varying widths and to choose between routes through woodland or across open areas by attracting them to a recording of mobbing calls by Chickadees (  Parus atricapillus). In 278 experiments conducted in boreal forest and agricultural landscapes near Québec city, 157 birds or flocks of birds of five species were attracted. Overall, birds were twice as likely to travel through 50 m of woodland than through 50 m in the open to reach the recording. When given a choice of traveling through woodland or across a gap, the majority of respondents preferred woodland routes, even when they were three times longer than shortcuts in the open. However, species differed greatly in their response to gaps. Our results show that woodland links significantly facilitate movements of birds across fragmented landscapes.     

Land Reform and Biodiversity Conservation in Brazil in the 1990s: Conflict and the Articulation of Mutual Interests

Abstract:  Land-reform and environmental movements, revitalized by the democratization of civil society in Brazil in the 1990s, found their objectives in conflict over forested parcels that settlers want for conversion to agriculture but that are important for wildlife conservation. In the Atlantic Forest, where 95% of the forest is gone, we reviewed three cases of Brazilian nongovernmental organization (NGOs) engagement with the land-reform movement with respect to forest remnants neighboring protected areas that have insufficient habitat for the long-term survival of unique endangered species. In the Pontal do Paranapanema (São Paulo), Poço das Antas (Rio de Janeiro), and southern Bahia, environmental NGOs have supported agricultural alternatives that improve livelihood options and provide incentives for habitat conservation planning. Where land-reform groups were better organized, technical cooperation on settlement agriculture permitted the exploration of mutual interests in conciliating the productive landscape with conservation objectives. Processes of regular consultation among NGOs, environmental agencies, and the private sector revealed that there was less zero-sum conflict over the same lands than commonly perceived. In both groups, technicians found forested lands less suitable for small-scale agriculture, and leaders took risks to justify and support claims to alternative existing agricultural lands. Based on the cases we examined, the construction of landscapes with both forest stewardship and poverty-reducing agrarian reform faces continued obstacles from contradictory agrarian and environmental sector policies and inadequate economic incentives for forest stewardship on private lands.     

Synergistic interactions between edge and area effects in a heavily fragmented landscape

Both area and edge effects have a strong influence on ecological processes in fragmented landscapes, but there is little understanding of how these two factors might interact to exacerbate local species declines. To test for synergistic interactions between area and edge effects, we sampled a diverse beetle community in a heavily fragmented landscape in New Zealand. More than 35,000 beetles of approximately 900 species were sampled over large gradients in habitat area (10(-2) 10(6) ha) and distance from patch edge (2(0)-2(10) m from the forest edge into both the forest and adjacent matrix). Using a new approach to partition variance following an ordination analysis, we found that a synergistic interaction between habitat area and distance to edge was a more important determinant of patterns in beetle community composition than direct edge or area effects alone. The strength of edge effects in beetle-species composition increased nonlinearly with increasing fragment area. One important consequence of the synergy is that the slopes of species area (SA) curves constructed from habitat islands depend sensitively on the distance from edge at which sampling is conducted. Surprisingly, we found negative SA curves for communities sampled at intermediate distances from habitat edges, caused by differential edge responses of matrix- vs. forest-specialist species in fragments of increasing area. Our data indicate that distance to habitat edge has a consistently greater impact on beetle community composition than habitat area and that variation in the strength of edge effects may underlie many patterns that are superficially related to habitat area.     

Landscape Context Moderates Edge Effects: Nesting Success of Wood Thrushes in Central New York

Abstract:  Despite two decades of research into the effects of habitat fragmentation and edges on nesting birds, critical information about how edges affect the success of natural nests of Neotropical migratory songbirds breeding in heterogeneous landscapes is still missing. We studied abundance and nesting success in Wood Thrushes ( Hylocichla mustelina ) breeding across a heterogeneous landscape in central New York from 1998 to 2000 to test the hypothesis that edge effects on nesting passerines are stronger in fragmented than contiguous landscapes. We monitored nests to estimate nesting success in edge and interior habitats in both fragmented and contiguously forested landscapes. In contiguous landscapes, daily survival rate did not differ between edge nests (0.963) and interior nests (0.968) (χ²= 0.19, p = 0.66). In contrast, in fragmented landscapes, daily survival estimates were higher in interior (0.971) than edge (0.953) nests (χ²= 3.1, p = 0.08). Our study supports the hypothesis that landscape composition moderates edge effects on actual nests of birds but does not determine the mechanisms causing these patterns.     

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.     

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.     

Modeling Dynamics of Habitat-Interior Bird Populations in Fragmented Landscapes

Abstract: A stochastic computer model was used to examine the effects of varying degrees of habitat fragmentation on the dynamics of a hypothetical population of forest-interior bid. The primary demographic parameter that influenced the population's dynamics was fecundity, which varied as a function of how far a birds territory was from an ecological edge. As our model landscape became more fragmented the proportion of forest habitat that was near edges increased geometrically, and the population's overall fecundity dropped as a result. The model demonstrates that impaired reproduction in a fragmented landscape is, by itself a sufficient disruption of the population's dynamics to generate population declines and shifts in distribution similar to those observed in the fragmented forests of southern Wisconsin. Without immigration of recruits from other regions where reproduction is better, habitat-interior populations in a severely fragmented landscape can become locally extinct.     

Wood Thrush Populations Are Not All Sinks in the Agricultural Midwestern United States

Abstract: Conservation strategies for forest-breeding, Neotropical migratory birds focus on the identification and preservation of source habitats. In sources, populations maintain relatively high levels of reproductive success, whereas in sinks, reproductive success is consistently below net replacement rate. In agricultural regions of the midwestern United States, forest patches of various sizes appear to act as sinks for Neotropical migrants. In 1994–1996 I monitored Wood Thrush (   Hylocichla mustelina ) nests in four forest patches in an agricultural landscape in northern Indiana. Given the prevailing estimates of adult and juvenile survival, I report productivity data that show substantial annual variation and that are sufficient to suggest the maintenance of temporal sources. Indeed, some Wood Thrush populations may act as source habitats in regions with relatively high levels of Brown-headed Cowbird (   Molothrys ater ) parasitism and nest predation because pairs produce multiple clutches and have the ability to fledge cowbirds and their own young from parasitized nests. My results and those from other studies of Wood Thrushes in landscapes with fragmented forests illustrate the variable ways in which source-sink dynamics are affected by habitat fragmentation. In some agricultural regions, populations in small forest patches may act as sources for Wood Thrushes and may be a legitimate focus of conservation efforts aimed at some migratory songbirds.     

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.