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.     

Disentangling the drivers of reduced long-distance seed dispersal by birds in an experimentally fragmented landscape

Seed dispersal is a crucial component of plant population dynamics. Human landscape modifications, such as habitat destruction and fragmentation, can alter the abundance of fruiting plants and animal dispersers, foraging rates, vector movement, and the composition of the disperser community, all of which can singly or in concert affect seed dispersal. Here, we quantify and tease apart the effects of landscape configuration, namely, fragmentation of primary forest and the composition of the surrounding forest matrix, on individual components of seed dispersal of Heliconia acuminata, an Amazonian understory herb. First we identified the effects of landscape configuration on the abundance of fruiting plants and six bird disperser species. Although highly variable in space and time, densities of fruiting plants were similar in continuous forest and fragments. However, the two largest-bodied avian dispersers were less common or absent in small fragments. Second, we determined whether fragmentation affected foraging rates. Fruit removal rates were similar and very high across the landscape, suggesting that Heliconia fruits are a key resource for small frugivores in this landscape. Third, we used radiotelemetry and statistical models to quantify how landscape configuration influences vector movement patterns. Bird dispersers flew farther and faster, and perched longer in primary relative to secondary forests. One species also altered its movement direction in response to habitat boundaries between primary and secondary forests. Finally, we parameterized a simulation model linking data on fruit density and disperser abundance and behavior with empirical estimates of seed retention times to generate seed dispersal patterns in two hypothetical landscapes. Despite clear changes in bird movement in response to landscape configuration, our simulations demonstrate that these differences had negligible effects on dispersal distances. However, small fragments had reduced densities of Turdus albicollis, the largest-bodied disperser and the only one to both regurgitate and defecate seeds. This change in Turdus abundance acted together with lower numbers of fruiting plants in small fragments to decrease the probability of long-distance dispersal events from small patches. These findings emphasize the importance of foraging style for seed dispersal and highlight the primacy of habitat size relative to spatial configuration in preserving biotic interactions.     

Fragmentation drives tropical forest fragments to early successional states: A modelling study for Brazilian Atlantic forests

Land use leads to massive habitat destruction and fragmentation in tropical forests. Despite its global dimensions the effects of fragmentation on ecosystem dynamics are not well understood due to the complexity of the problem. We present a simulation analysis performed by the individual-based model FORMIND. The model was applied to the Brazilian Atlantic Forest, one of the world's biodiversity hot spots, at the Plateau of São Paulo. This study investigates the long-term effects of fragmentation processes on structure and dynamics of different sized remnant tropical forest fragments (1-100 ha) at community and plant functional type (PFT) level. We disentangle the interplay of single effects of different key fragmentation processes (edge mortality, increased mortality of large trees, local seed loss and external seed rain) using simulation experiments in a full factorial design.Our analysis reveals that particularly small forest fragments below 25 ha suffer substantial structural changes, biomass and biodiversity loss in the long term. At community level biomass is reduced up to 60%. Two thirds of the mid- and late-successional species groups, especially shade-tolerant (late successional climax) species groups are prone of extinction in small fragments. The shade-tolerant species groups were most strongly affected; its tree number was reduced more than 60% mainly by increased edge mortality. This process proved to be the most powerful of those investigated, explaining alone more than 80% of the changes observed for this group. External seed rain was able to compensate approximately 30% of the observed fragmentation effects for shade-tolerant species.Our results suggest that tropical forest fragments will suffer strong structural changes in the long term, leading to tree species impoverishment. They may reach a new equilibrium with a substantially reduced subset of the initial species pool, and are driven towards an earlier successional state. The natural regeneration potential of a landscape scattered with forest fragments appears to be limited, as external seed rain is not able to fully compensate for the observed fragmentation-induced changes. Our findings suggest basic recommendations for the management of fragmented tropical forest landscapes.     

Fragmentation effects on an endangered species across a gradient from the interior to edge of its range

Understanding how habitat fragmentation affects individual species is complicated by challenges associated with quantifying species-specific habitat and spatial variability in fragmentation effects within a species’ range. We aggregated a 29-year breeding survey data set for the endangered marbled murrelet (Brachyramphus marmoratus) from >42,000 forest sites throughout the Pacific Northwest (Oregon, Washington, and northern California) of the United States. We built a species distribution model (SDM) in which occupied sites were linked with Landsat imagery to quantify murrelet-specific habitat and then used occupancy models to test the hypotheses that fragmentation negatively affects murrelet breeding distribution and that these effects are amplified with distance from the marine foraging habitat toward the edge of the species’ nesting range. Murrelet habitat declined in the Pacific Northwest by 20% since 1988, whereas the proportion of habitat comprising edges increased by 17%, indicating increased fragmentation. Furthermore, fragmentation of murrelet habitat at landscape scales (within 2 km of survey stations) negatively affected occupancy of potential breeding sites, and these effects were amplified near the range edge. On the coast, the odds of occupancy decreased by 37% (95% confidence interval [CI] –54 to 12) for each 10% increase in edge habitat (i.e., fragmentation), but at the range edge (88 km inland) these odds decreased by 99% (95% CI 98 to 99). Conversely, odds of murrelet occupancy increased by 31% (95% CI 14 to 52) for each 10% increase in local edge habitat (within 100 m of survey stations). Avoidance of fragmentation at broad scales but use of locally fragmented habitat with reduced quality may help explain the lack of murrelet population recovery. Further, our results emphasize that fragmentation effects can be nuanced, scale dependent, and geographically variable. Awareness of these nuances is critical for developing landscape-level conservation strategies for species experiencing broad-scale habitat loss and fragmentation.     

Mediation of area and edge effects in forest fragments by adjacent land use

Habitat loss, fragmentation, and degradation have pervasive detrimental effects on tropical forest biodiversity, but the role of the surrounding land use (i.e., matrix) in determining the severity of these impacts remains poorly understood. We surveyed bird species across an interior-edge-matrix gradient to assess the effects of matrix type on biodiversity at 49 different sites with varying levels of landscape fragmentation in the Brazilian Atlantic Forest—a highly threatened biodiversity hotspot. Both area and edge effects were more pronounced in forest patches bordering pasture matrix, whereas patches bordering Eucalyptus plantation maintained compositionally similar bird communities between the edge and the interior and exhibited reduced effects of patch size. These results suggest the type of matrix in which forest fragments are situated can explain a substantial amount of the widely reported variability in biodiversity responses to forest loss and fragmentation.     

Landscape history, fragmentation, and patch occupancy: models for a forest bird with limited dispersal.

We developed and tested patch occupancy models for an endemic understory bird with limited dispersal ability, the Chucao Tapaculo (Scelorchilus rubecula), in two South American temperate rain forest landscapes that differed in levels and duration of forest loss. We assessed cover changes since 1961 in each landscape and surveyed patches for Chucao Tapaculo occupancy. We then developed incidence-based predictive models independently for each landscape and tested each model reciprocally in the alternative study area. We thereby assessed the domain of model applicability and identified those predictor variables with general effects and those that varied between the two landscapes. The two models were consistent regarding variable selection, and predictive accuracy of each model was high in the landscape where training data were collected. However, the models differed substantially in the magnitudes of effects related to patch size, with larger unoccupied patches observed in the landscape with the more advanced stage of fragmentation. Due to this discrepancy, each model performed poorly when applied to the alternative landscape, potentially reflecting the contrasting stages of habitat loss. Although it was impossible to dissociate effects of level and duration of forest loss, we viewed the landscapes as representing two extremes along a continuum of fragmentation, providing insights into potential trajectories for portions of the biome where deforestation is occurring. Further, our data suggest that static equilibrium models developed from distribution patterns in recently fragmented landscapes may overestimate persistence when used as a forecasting tool, or when extrapolated to alternative landscapes where fragmentation is more advanced. Thus, we suggest that landscapes used as standards for model building should be selected with caution. We recommend that distribution patterns be obtained from landscapes where fragmentation is advanced, preferably with histories of fragmentation long enough that time-delayed extinctions already would have occurred.     

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.     

Rain forest fragmentation and the proliferation of successional trees

The effects of habitat fragmentation on diverse tropical tree communities are poorly understood. Over a 20-year period we monitored the density of 52 tree species in nine predominantly successional genera (Annona, Bellucia, Cecropia, Croton, Goupia, Jacaranda, Miconia, Pourouma, Vismia) in fragmented and continuous Amazonian forests. We also evaluated the relative importance of soil, topographic, forest dynamic, and landscape variables in explaining the abundance and species composition of successional trees. Data were collected within 66 permanent 1-ha plots within a large (approximately 1000 km2) experimental landscape, with forest fragments ranging from 1 to 100 ha in area. Prior to forest fragmentation, successional trees were uncommon, typically comprising 2-3% of all trees (> or =10 cm diameter at breast height [1.3 m above the ground surface]) in each plot. Following fragmentation, the density and basal area of successional trees increased rapidly. By 13-17 years after fragmentation, successional trees had tripled in abundance in fragment and edge plots and constituted more than a quarter of all trees in some plots. Fragment age had strong, positive effects on the density and basal area of successional trees, with no indication of a plateau in these variables, suggesting that successional species could become even more abundant in fragments over time. Nonetheless, the 52 species differed greatly in their responses to fragmentation and forest edges. Some disturbance-favoring pioneers (e.g., Cecropia sciadophylla, Vismia guianensis, V. amazonica, V. bemerguii, Miconia cf. crassinervia) increased by >1000% in density on edge plots, whereas over a third (19 of 52) of all species remained constant or declined in numbers. Species responses to fragmentation were effectively predicted by their median growth rate in nearby intact forest, suggesting that faster-growing species have a strong advantage in forest fragments. An ordination analysis revealed three main gradients in successional-species composition across our study area. Species gradients were most strongly influenced by the standlevel rate of tree mortality on each plot and by the number of nearby forest edges. Species-composition also varied significantly among different cattle ranches, which differed in their surrounding matrices and disturbance histories. These same variables were also the best predictors of total successional-tree abundance and species richness. Successional-tree assemblages in fragment interior plots (>150 m from edge), which are subjected to fragment area effects but not edge effects, did not differ significantly from those in intact forest, indicating that area effects per se had little influence on successional trees. Soils and topography also had little discernable effect on these species. Collectively, our results indicate that successional-tree species proliferate rapidly in fragmented Amazonian forests, largely as a result of chronically elevated tree mortality near forest edges and possibly an increased seed rain from successional plants growing in nearby degraded habitats. The proliferation of fast-growing successional trees and correlated decline of old-growth trees will have important effects on species composition, forest dynamics, carbon storage, and nutrient cycling in fragmented forests.     

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.     

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.     

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.     

Influence of Forest Fragmentation on Community Structure of Frogs and Lizards in Northeastern Costa Rica

Abstract:  To better understand responses of reptiles and amphibians to forest fragmentation in the lowland Neotropics, we examined community and population structure of frogs and lizards in the fragmented landscape surrounding La Selva Biological Station in the Sarapiquí region of northeastern Costa Rica. We used diurnal quadrats and nocturnal transects to sample frogs and lizards in nine forest fragments (1–7 ha each) and La Selva (1100 ha). Species richness in all fragments combined was 85% of that found in La Selva with comparable sampling effort. Richness varied from 10 to 24 species among forest fragments, compared with 36 species at La Selva. Lizard density was higher and frog density was lower in forest fragments than in La Selva. Community composition varied among sites and by fragment size class, and species occurrence was nested with respect to fragment area. Isolation and habitat variables did not significantly affect species richness, composition, or nestedness. We classified 34% of species as fragmentation sensitive because they were absent or occurred at low densities in fragments. Nevertheless, the relatively high diversity observed in the entire set of fragments indicates that preserving a network of small forest patches may be of considerable conservation value to the amphibians and reptiles of this region.     

Local and Landscape Correlates of Primate Distribution and Persistence in the Remnant Lowland Rainforests of the Upper Brahmaputra Valley,Northeastern India

Habitat fragmentation affects species distribution and abundance, and drives extinctions. Escalated tropical deforestation and fragmentation have confined many species populations to habitat remnants. How worthwhile is it to invest scarce resources in conserving habitat remnants within densely settled production landscapes? Are these fragments fated to lose species anyway? If not, do other ecological, anthropogenic, and species‐related factors mitigate the effect of fragmentation and offer conservation opportunities? We evaluated, using generalized linear models in an information‐theoretic framework, the effect of local‐ and landscape‐scale factors on the richness, abundance, distribution, and local extinction of 6 primate species in 42 lowland tropical rainforest fragments of the Upper Brahmaputra Valley, northeastern India. On average, the forest fragments lost at least one species in the last 30 years but retained half their original species complement. Species richness declined as proportion of habitat lost increased but was not significantly affected by fragment size and isolation. The occurrence of western hoolock gibbon (Hoolock hoolock) and capped langur (Trachypithecus pileatus) in fragments was inversely related to their isolation and loss of habitat, respectively. Fragment area determined stump‐tailed (Macaca arctoides) and northern pig‐tailed macaque occurrence (Macaca leonina). Assamese macaque (Macaca assamensis) distribution was affected negatively by illegal tree felling, and rhesus macaque (Macaca mulatta) abundance increased as habitat heterogeneity increased. Primate extinction in a fragment was primarily governed by the extent of divergence in its food tree species richness from that in contiguous forests. We suggest the conservation value of these fragments is high because collectively they retained the entire original species pool and individually retained half of it, even a century after fragmentation. Given the extensive habitat and species loss, however, these fragments urgently require protection and active ecological restoration to sustain this rich primate assemblage. Correlaciones Locales y de Paisaje de la Distribución y Persistencia de Primates en los Bosques Lluviosos Remanentes en el Valle del Alto Brahmaputra, Noreste de India     

Are shrubland birds edge specialists?

In studies of forest fragmentation, birds of scrubby, early-successional habitats are considered edge specialists. Because these birds are assumed to thrive in fragmented, edge-dominated areas, their landscape ecology has received little attention from ecologists. With populations of shrubland birds declining throughout the eastern United States, the question of whether or not these birds really prefer edge habitats has important conservation implications. We used a meta-analysis to test how edges affect the abundance of shrubland birds in early-successional habitats. We analyzed data for 17 species from seven studies that compared the abundances of birds in the interiors and edges of regenerating clearcuts surrounded by mature forest. The meta-analysis clearly showed that shrubland birds avoid edges. All 17 species tested had higher abundances in patch centers than along edges, and edge effects were significant for 8 of 17 species. The key implication of this result is that small or irregular patches, dominated by edge, are unlikely to provide suitable habitat for shrubland birds. Thus, management for these declining species should involve providing large patches and minimizing edges. These findings demonstrate the importance of testing widely accepted ecological classifications and the need to view landscape ecology from the perspective of non-forest wildlife.     

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.     

Reduced Feather Growth Rates of Two Common Birds Inhabiting Central Amazonian Forest Fragments

Abstract: Forest fragmentation may negatively affect populations typically found within continuous forest tracts. Some effects, such as absence from small fragments, are obvious, but other effects may be subtle and easily overlooked. We evaluated the hypothesis that forest birds dwelling in fragments, where microclimatic conditions have been shown to be hotter and drier than in continuous forest, may be in poorer physiological condition than those in the forest interior. We studied two bird species, the Wedge-billed Woodcreeper (  Glyphorynchus spirurus ) and the White-crowned Manakin (    Pipra pipra ), common to the fragmented landscape north of Manaus, Brazil. We analyzed feather growth rates in Pipra and Glyphorynchus captured in 1-, 10-, and 100-ha forest fragments and continuous forest. Mean daily feather growth rates of the outer right rectrix of birds captured in fragments were significantly slower than feather growth rates of birds captured in continuous forest. Based on recapture data, Wedge-billed Woodcreepers probably grew their feathers in sites where they were first captured. White-crowned Manakins, however, were highly mobile and were recaptured rarely. Although we cannot conclusively show that fragmentation caused birds to be in poorer physiological condition, the data indicate that birds in poorer physiological condition were more likely to be captured in fragments than in continuous forest. Thus, our data suggest that forest fragmentation may have subtle but important effects on species that are relatively common after landscape alteration.     

Responses of Seed‐Dispersing Birds to Amount of Rainforest in the Landscape around Fragments

Habitat loss and fragmentation alter the composition of bird assemblages in rainforest. Because birds are major seed dispersers in rainforests, fragmentation‐induced changes to frugivorous bird assemblages are also likely to alter the ecological processes of seed dispersal and forest regeneration, but the specific nature of these changes is poorly understood. We assessed the influence of fragment size and landscape forest cover on the abundance, species composition, and functional properties of the avian seed disperser community in an extensively cleared, former rainforest landscape of subtropical Australia. Bird surveys of fixed time and area in 25 rainforest fragments (1–139 ha in size across a 1800 km² region) provided bird assemblage data which were coupled with prior knowledge of bird species’ particular roles in seed dispersal to give measurements of seven different attributes of the seed disperser assemblage. We used multimodel regression to assess how patch size and surrounding forest cover (within 200 m, 1000 m, and 5000 m radii) influenced variation in the abundance of individual bird species and of functional groups based on bird species’ responses to fragmentation and their roles in seed dispersal. Surrounding forest cover, specifically rainforest cover, generally had a greater effect on frugivorous bird assemblages than fragment size. Amount of rainforest cover within 200 m of fragments was the main factor positively associated with abundances of frugivorous birds that are both fragmentation sensitive and important seed dispersers. Our results suggest a high proportion of local rainforest cover is required for the persistence of seed‐dispersing birds and the maintenance of seed dispersal processes. Thus, even small rainforest fragments can function as important parts of habitat networks for seed‐dispersing birds, whether or not they are physically connected by vegetation. Respuestas de Aves Dispersoras de Semillas al Incremento de Selvas en el Paisaje Alrededor de Fragmentos     

Associations of Forest Cover,Fragment Area,and Connectivity with Neotropical Understory Bird Species Richness and Abundance

Theoretical and empirical studies demonstrate that the total amount of forest and the size and connectivity of fragments have nonlinear effects on species survival. We tested how habitat amount and configuration affect understory bird species richness and abundance. We used mist nets (almost 34,000 net hours) to sample birds in 53 Atlantic Forest fragments in southeastern Brazil. Fragments were distributed among 3 10,800‐ha landscapes. The remaining forest in these landscapes was below (10% forest cover), similar to (30%), and above (50%) the theoretical fragmentation threshold (approximately 30%) below which the effects of fragmentation should be intensified. Species‐richness estimates were significantly higher (F= 3715, p = 0.00) where 50% of the forest remained, which suggests a species occurrence threshold of 30–50% forest, which is higher than usually occurs (<30%). Relations between forest cover and species richness differed depending on species sensitivity to forest conversion and fragmentation. For less sensitive species, species richness decreased as forest cover increased, whereas for highly sensitive species the opposite occurred. For sensitive species, species richness and the amount of forest cover were positively related, particularly when forest cover was 30–50%. Fragment size and connectivity were related to species richness and abundance in all landscapes, not just below the 30% threshold. Where 10% of the forest remained, fragment size was more related to species richness and abundance than connectivity. However, the relation between connectivity and species richness and abundance was stronger where 30% of the landscape was forested. Where 50% of the landscape was forested, fragment size and connectivity were both related to species richness and abundance. Our results demonstrated a rapid loss of species at relatively high levels of forest cover (30–50%). Highly sensitive species were 3‐4 times more common above the 30–50% threshold than below it; however, our results do not support a unique fragmentation threshold. Asociaciones de la Cobertura Forestal, Superficie del Fragmento y Conectividad con la Riqueza y Abundancia de Aves Neotropicales de Sotobosque     

Conservation of Epiphyte Diversity in an Andean Landscape Transformed by Human Land Use

Abstract: Epiphytes are diverse and important elements of tropical forests, but as canopy‐dwelling organisms, they are highly vulnerable to deforestation. To assess the effect of deforestation on epiphyte diversity and the potential for epiphyte conservation in anthropogenically transformed habitats, we surveyed the epiphytic vegetation of an Ecuadorian cloud forest reserve and its surroundings. Our study was located on the western slopes of the Andes, a global center of biodiversity. We sampled vascular epiphytes of 110 study plots in a continuous primary forest; 14 primary forest fragments; isolated remnant trees in young, middle‐aged, and old pastures; and young and old secondary forests. It is the first study to include all relevant types of habitat transformation at a single study site and to compare epiphyte diversity at different temporal stages of fragmentation. Epiphyte diversity was highest in continuous primary forest, followed by forest fragments and isolated remnant trees, and lowest in young secondary forests. Spatial parameters of habitat transformation, such as fragment area, distance to the continuous primary forest, or distance to the forest edge from inside the forest, had no significant effect on epiphyte diversity. Hence, the influence of dispersal limitations appeared to be negligible or appeared to operate only over very short distances, whereas microclimatic edge effects acted only in the case of completely isolated trees, but not in larger forest fragments. Epiphyte diversity increased considerably with age of secondary forests, but species assemblages on isolated remnant trees were impoverished distinctly with time since isolation. Thus, isolated trees may serve for recolonization of secondary forests, but only for a relatively short time. We therefore suggest that the conservation of even small patches of primary forest within agricultural landscape matrices is essential for the long‐term maintenance of the high epiphyte diversity in tropical cloud forests.     

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.