Vegetation Responses along Edge-to-Interior Gradients in the Mixed Hardwood Forests of the Roanoke River Basin, North Carolina

Compared with forest interiors, forest edges typically have a different plant species composition and community structure, a phenomenon known as "edge effect." Edge effects make the functional interior area of a forest smaller than its actual area. The objective of this study was to estimate how far the effects of agriculturally maintained edges penetrate the mixed hardwood forests of the Roanoke River Basin, North Carolina. I determined percentage cover for all vascular plant species in 10-by-100-meter belt transects on north-facing or south-facing edges of four relatively undisturbed forests. Changes in the percentage cover of individual species, the relative cover of exotic species, and species richness all indicated that edge effects penetrate deeper on south-facing edges (to 60 meters) than on north-facing edges (to 20 meters). Analyses of species responses to the edge showed a number of species to be edge oriented, but no species was found to be interior oriented. The results of multivariate analyses (ordination and cluster analysis) suggested that edge effects could be detected to 50 meters on south-facing edges and 10–30 meters on north-facing edges. These results allow us to better understand the difference between a forest's actual area and its functional interior area.     

Parasitism at the Landscape Scale: Cowbirds Prefer Forests

Landscape-scale examination of parasitism patterns of Brown-headed Cowbirds ( Molothrus ater ) revealed heterogeneous parasitism rates across the mosaic of a forest and associated old-field communities. In a two year study in Dutchess County, New York, we found a significantly higher parasitism rate in the forest-interior community ( n = 301 nests; 17 species) than on the species in the adjacent and nearby old field and edge ( n = 328 nests; 15 species; 32.3% versus 6.5%; p < 0.0001). Cowbirds invaded a mature 1300-ha forest stand even when their traditional host species were available in adjacent old-field and edge habitats. The forest and old-field study areas were located in a 38,000-ha township with 55% forest cover and contained numerous agriculture, dairy, and horse farms that provided favorable habitat for cowbirds. Within-forest examination of parasitism patterns revealed four aspects of cowbird parasitism that contrasted with patterns described in other regions: (1) parasitism was concentrated significantly more often on ground- and low-nesting (nests ≤ 1 m) forest species than on medium- and high-nesting species (nests> 1 m; 35.01% versus 29.93%; p = 0.0393); (2) parasitism was not significantly greater on Neotropical migrant species than on short-distance migrants and residents; (3) the parasitism rate was not higher in nests close to edges; and (4) the parasitism level was low on certain forest species (such as Wood Thrush) that have experienced high parasitism levels in the Midwest. From a management perspective these data suggest that cowbirds exhibit regional differences in host and habitat use; the target host community of a particular cowbird population is unpredictable at the landscape scale; and a landscape scale should be used in designing cowbird studies to accurately assess local population dynamics.     

Conservation Value of Remnant Riparian Forest Corridors of Varying Quality for Amazonian Birds and Mammals

Abstract:  Forest corridors are often considered the main instrument with which to offset the effects of habitat loss and fragmentation. Brazilian forestry legislation requires that all riparian zones on private landholdings be maintained as permanent reserves and sets fixed minimum widths of riparian forest buffers to be retained alongside rivers and perennial streams. We investigated the effects of corridor width and degradation status of 37 riparian forest sites (including 24 corridors connected to large source-forest patches, 8 unconnected forest corridors, and 5 control riparian zones embedded within continuous forest patches) on bird and mammal species richness in a hyper-fragmented forest landscape surrounding Alta Floresta, Mato Grosso, Brazil. We used point-count and track-sampling methodology, coupled with an intensive forest-quality assessment that combined satellite imagery and ground truthed data. Vertebrate use of corridors was highly species-specific, but broad trends emerged depending on species life histories and their sensitivity to disturbance. Narrow and/or highly disturbed riparian corridors retained only a depauperate vertebrate assemblage that was typical of deforested habitats, whereas wide, well-preserved corridors retained a nearly complete species assemblage. Restriction of livestock movement along riparian buffers and their exclusion from key areas alongside deforested streams would permit corridor regeneration and facilitate restoration of connectivity .     

Effects of Fragmentation of Araucarian Vine Forest on Small Mammal Communities

Abstract: Habitat loss and fragmentation are major threats to the survival of forest-dependent fauna. We examined the abundance of small mammal species in forests, corridors, remnants of araucarian vine forest, and Araucaria cunninghamii plantations and pastures. None of the forest mammal species persisted following conversion of forest to pasture. Plantations supported lowered abundances of a subset of forest species that were mainly habitat generalists with respect to their occurrence in different floristic types of undisturbed native forest. Within plantations, an increased subcanopy cover was associated with a more forest-like small mammal assemblage. Species' responses to habitat fragmentation varied. The floristic habitat generalists were largely tolerant of habitat fragmentation, their abundance being similar in forests, corridors, and remnants, and were capable of persisting in remnants a few hectares in area. Floristic habitat specialists were vulnerable to habitat fragmentation and thus were abundant in continuous forest, were less abundant in corridors, and were generally absent from remnants. Species that avoid the corridor matrix and are therefore constrained to the corridor may be disadvantaged by the linearity of the habitat, consistent with the predictions of central-place foraging theory. Although small remnants and corridors provide habitat for some species, those that are more specialized in their use of undisturbed habitat types require the retention or reestablishment of large intact areas.     

Effects of Exurban Development and Temperature on Bird Species in the Southern Appalachians

Land‐use dynamics and climatic gradients have large effects on many terrestrial systems. Exurban development, one of the fastest growing forms of land use in the United States, may affect wildlife through habitat fragmentation and building presence may alter habitat quality. We studied the effects of residential development and temperature gradients on bird species occurrence at 140 study sites in the southern Appalachian Mountains (North Carolina, U.S.A.) that varied with respect to building density and elevation. We used occupancy models to determine 36 bird species’ associations with building density, forest canopy cover, average daily mean temperature, and an interaction between building density and mean temperature. Responses varied with habitat requirement, breeding range, and migration distance. Building density and mean temperature were both included in the top occupancy models for 19 of 36 species and a building density by temperature interaction was included in models for 8 bird species. As exurban development expands in the southern Appalachians, interior forest species and Neotropical migrants are likely to decline, but shrubland or edge species are not likely to benefit. Overall, effects of building density were greater than those of forest canopy cover. Exurban development had a greater effect on birds at high elevations due to a greater abundance of sensitive forest‐interior species and Neotropical migrants. A warming climate may exacerbate these negative effects. Efectos del Desarrollo Exurbano y de la Temperatura sobre Especies de Aves en las Apalaches del Sur     

Edge Effects on the Understory Bird Community in a Logged Forest in Uganda

Abstract: Understanding how the fauna of logged tropical rainforests responds to fragmentation and the creation of edges is vital to ensure conservation of biodiversity. We studied the composition of the understory bird community from the edge of a 15-ha clearing toward the interior of the forest in a part of Budongo Forest Reserve, Uganda, that was selectively logged about 45 years ago. Mist netting was conducted along five transects from the edge and 500 m into the interior. The total number of individuals captured did not change with distance from the edge, but there was a significant increase in the number of species. We sampled fewer, but more common species near the edge, whereas the interior of the forest had more, and less common species. Guild composition also changed with distance from the edge. Frugivore-insectivores and nectarivores were most common close to the edge. Among insectivores, ground foragers, bark-gleaners, and leaf-gleaners were most common in the interior of the forest, whereas sallying insectivores favored the edge. Graminivores were unaffected by the edge. Analysis of common species showed that Ispidina picta , Andropadus curvirostris , A. latirostris , Camaroptera brachyura , Terpsiphone rufiventer , and Nectarinia olivacea were associated with the edge, but no species showed significant avoidance of the edge. This finding may be explained by the generally low sample sizes of interior species. Our results show that even bird communities in logged forests respond to edges. Estimates of edge effects suggested that changes in bird densities may have occurred several hundred meters from the edge. In conclusion, logged forests provide habitat for bird species avoiding forest edges, and this should be considered in the management of such forests for conservation.     

Empirical selection between least-cost and current-flow designs for establishing wildlife corridors in Gabon

Corridors are intended to increase species survival by abating landscape fragmentation resulting from the conversion of natural habitats into human-dominated matrices. Conservation scientists often rely on 1 type of corridor model, typically the least-cost model or current-flow model, to construct a linkage design, and their choice is not usually based on theory or empirical evidence. We developed a method to empirically confirm whether corridors produced by these 2 models are used by target species under current landscape conditions. We applied this method in the Gamba landscape between 2 national parks in southwestern Gabon. We collected signs of presence of African forest elephant (Loxodonta cyclotis), forest buffalo (Syncerus caffer nanus), and 2 apes, western lowland gorilla (Gorilla gorilla gorilla) and central chimpanzee (Pan troglodytes troglodytes), on transects. We used patch-occupancy models to identify least-cost and current-flow corridors for these 4 species. On average, 28.7% of current-flow corridors overlapped with least-cost corridors, confirming that the choice of corridor model can affect the location of the resulting linkage design. We validated these corridors by monitoring signs and examining camera detections on new transects within and outside modeled corridors. Current-flow corridors performed better than least-cost corridors for elephants, whereas the opposite was found for buffalo and apes. Locations of the highest priority corridors for the 3 taxa did not overlap, and only 18.3% of their combined surface was common among 2 species. We used centrality metrics to calculate the average contribution of corridor pixels to landscape connectivity and derived an index that can be used to prioritize corridors. As a result, we recommend protecting at least 17.4% of the land surface area around Gamba town to preserve the preferred travel routes of the target species.     

The Use of Corridors by Mammals in Fragmented Australian Eucalypt Forests

We used a replicated sampling program to examine the use of roadside corridors as habitat by native mammals. Our procedure compared the abundance and diversity of mammals in remnant forest, pasture, and two types of roadside corridor. Fixed transects were established in these four habitat types at six replicate sites. Spotlight, live-trap, and daytime observation surveys were used as census techniques. Few mammals were detected in pasture, and spotlighting revealed a higher total density of mammals in corridors than in forests, indicating corridors provide important habitat. Nevertheless, the number of species using corridors distant to forest was less than that in the corridors close to forest and the forest patches. Different species did not utilize corridors in the same way. We also found intraspecific differences in habitat use by one species of small mammal (Antechinus stuartii), which may have implications for the value of corridors to this species. Specifically, there was a higher proportion of males, and individuals of both sexes had lower body weight in corridors than in forests. Our study demonstrates that corridors can provide useful habitat for mammalian assemblages, but may not provide a complete solution to the problem of landscape fragmentation.     

Effects of Roads on Movements by Understory Birds in Mixed-Species Flocks in Central Amazonian Brazil

Abstract: Roads through tropical forest create linear disturbances that have unknown consequences for forest birds. We studied how a narrow, rarely used road through otherwise undisturbed Amazonian forest affected the movements and area requirements of understory birds that form mixed-species flocks. Differences in road maintenance led to two distinct treatments along the same road. Trees along the "closed" road formed a partial canopy connecting the two sides of the road, although the roadway itself was kept open. The "open" road was regularly maintained, making a complete opening 10–30 m wide.We followed 15 flocks, 5 each in interior forest, along the open road, and along the closed road. These flocks were led by Thamnomanes antshrikes, and each flock had a discreet, permanent territory. Flock territory size (mean = 8.5 ha) did not differ among the three locations. The open road formed the territorial boundary for all five flocks, although birds moved within a few meters of the edge of the road. The closed road was less of a barrier: 2 of 5 flocks used both sides of the closed road. Playback experiments showed that flocks readily crossed the closed road to approach agonistic vocalizations. Along the open road, even though birds responded to playback by becoming agitated and moving to the extreme edge of the roadside vegetation, they were less likely to cross the road and did so only after a longer duration of playback. Our results suggest that flocks respond to a road as they would to a long linear gap. They use the vegetation along the edges of the road, but because they are unwilling to cross the open area, it becomes a flock territory boundary. Similarly, as in forest gaps, successional change along the closed road produced suitable habitat for flocks. Although this suggests that roads are a trivial problem, we caution that this result applies only to narrow roads that are not accompanied by deforestation or other disturbance.     

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.     

Linear Strips of Rain Forest Vegetation as Potential Dispersal Corridors for Rain Forest Insects

The potential of linear strips of vegetation to act as corridors to facilitate dispersal is examined for three taxa of insects in lowland rain forest in northeastern Australia. The taxa selected were ants, butterflies, and dung beetles, all of which are taxonomically well known and could be considered bioindicator groups. The sampling design encompassed four habitats, namely rain-forest interior, rain-forest edge, rain-forest linear strip (corridor), and arable land. Ants and dung beetles were sampled using baited pitfall traps, and visual surveys were used to census butterflies. Potential increase in dispersal was examined by first identifying those species that specialized on the rain-forest interior habitat and then determining whether these species were present in the linear strips as opposed to the surrounding arable land. Two species of butterfly and two species of dung beetle were identified as rain-forest interior specialists, and two of these species were found in the linear strips but not in the arable habitat. This result supports the concept that the presence of corridors can increase the potential for dispersal of these species. But the remaining rain-forest interior species did not occur in the linear strips, which suggests that corridors will not increase dispersal for these species.     

Effects of Residential Development on Forest-Dwelling Neotropical Migrant Songbirds

Many bird species are in local or regional decline because of habitat loss or degradation. We attempted to disentangle the effects of residential development from the effects of forest size on forest-bird communities, with particular emphasis on Neotropical migrant species. Two variables were examined for their influence on avian diversity and abundance: forest size and the number of houses outside a forest within 100 m of the edge. We found that Neotropical migrants consistently increased in number and abundance as forest size increased. Of greater interest, we found that the number of houses surrounding a forest severely undermined its suitability for Neotropical migrants. Neotropical migrants consistently decreased in diversity and abundance as the level of adjacent development increased, regardless of forest size. The effects of development were striking: 4-ha woodlots without any nearby houses had on average a richer, more abundant Neotropical community than did 25-ha urban woodlots. No predictable pattern of change concerning development or forest size was observed for short-distance migrants or permanent residents. Current planning regulations generally permit housing right up to forest edges. This practice may prevent protection of ecological features within the forest. Threshold distances for housing developments around forests need to be determined to prevent or minimize adverse effects on features and functions within the forests.     

Effects of Forest Roads on Macroinvertebrate Soil Fauna of the Southern Appalachian Mountains

Abstract: Many forested landscapes are fragmented by roads, but our understanding of the effects of these roads on the function and diversity of the surrounding forest is in its infancy. I investigated the effect of roads in otherwise continuous forests on the macroinvertebrate fauna of the soil. I took soil samples along transects leading away from the edges of unpaved roads in the Cherokee National Forest in the Southern Appalachian mountains of the United States. Roads significantly depressed both the abundance and the richness of the macroinvertebrate soil fauna. Roads also significantly reduced the depth of the leaf-litter layer. These effects persisted up to 100 m into the forest. Wider roads and roads with more open canopies tended to produce steeper declines in abundance, richness, and leaf-litter depth, but these effects were significant only for canopy cover and litter depth. The macroinvertebrate fauna of the leaf litter plays a pivotal role in the ability of the soil to process energy and nutrients. These macroinvertebrates also provide prey for vertebrate species such as salamanders and ground-foraging birds. The effect of roads on the surrounding forest is compounded by the sprawling nature of the road system in this and many other forests. My data suggest that even relatively narrow roads through forests can produce marked edge effects that may have negative consequences for the function and diversity of the forest ecosystem.     

Role of Light Availability and Dispersal in Exotic Plant Invasion along Roads and Streams in the H. J. Andrews Experimental Forest, Oregon

Abstract: We examined the roles of dispersal mechanism, a biological barrier; light availability, an environmental barrier; and level of disturbance, a physical barrier, in explaining the spatial patterns of exotic plant species along road and stream segments in a forest landscape in the western Cascade Range of Oregon (U.S.A). The presence or absence of 21 selected exotic plant species and light levels were observed along 0.3- to 1.0-km transects within four habitat types. Each habitat represented a different level of disturbance: high-use roads, low-use roads, abandoned roads, and streams in the H. J. Andrews Experimental Forest. Nearly 300 50 × 2–m sampling units were surveyed along five transects in each habitat type. We used ordination (nonmetric multidimensional scaling) and logistic regression to analyze data. All of the nearly 200 sampling units along roads with high and low levels of vehicle traffic contained at least one exotic plant species, and some contained as many as 14. Streams that were most recently disturbed by floods 20–30 years ago and abandoned spur roads with no traffic for 20–40 years also had numerous exotic species. Roads and streams apparently serve multiple functions that enhance exotic species invasion in this landscape: they act as corridors or agents for dispersal, provide suitable habitat, and contain reservoirs of propagules for future episodes of invasion. Species-specific dispersal mechanisms, habitat characteristics, and disturbance history each explain some, but not all, of the patterns of exotic species invasion observed in this study.     

Distribution of Carabid Beetles (Coleoptera, Carabidae) across a Boreal Forest–Clearcut Ecotone

Abstract: We studied the occurrence of carabid beetles (Coleoptera, Carabidae) in boreal forest fragments, their edges, and adjacent clearcuts in central Finland. Beetles were collected with pitfall traps along transects extending 60 m from the edge into clearcuts and 60 m into forest interior. Our main findings were that (1) species richness was significantly higher in the clearcut than in the forest fragments, (2) clearcuts hosted many open-habitat species, which increased overall species richness in these sites, (3) carabid assemblages in the edges were more similar to forest assemblages than to those found in the clearcuts, (4) no edge specialists were found, and (5) open-habitat species did not penetrate into the forest fragments from the clearcut. Because forest specialists occurred all the way to the edge on the forest side, it seems that edge effects per se do not adversely affect these species, at least in the short term. In the long term, however, habitat conditions in the edges may deteriorate for interior species because of trees falling over in strong winds, thereby reducing the size of the fragments and widening the edge zone.     

Tropical Forest Fragmentation and the Local Extinction of Understory Birds in the Eastern Usambara Mountains, Tanzania

Abstract. Small tropical forest fragments in the Eastern Usambara Mountains have lost understory bird species following forest fragmentation. The local extinction of these species is documented through a comparison of species number on nine smaller fores t fragments with species number on a tenth larger control forest site. The number of species captured in the control forest site is significantly larger than the number captured in eight of the nine smaller forest fragments under an equivalent capture effort. Forest-dependent understory species vary greatly in their vulnerability to forest fragmentation. Relatively rare species and forest interior species are the avifauna most adversely affected by forest fragmentation. A concerted effort should be made to protect the existing corridors of native forest that link established forest reserves in the Eastern Usambara Mountains.     

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.     

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.     

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.     

Potential Negative Ecological Effects of Corridors

Despite many studies showing that landscape corridors increase dispersal and species richness for disparate taxa, concerns persist that corridors can have unintended negative effects. In particular, some of the same mechanisms that underlie positive effects of corridors on species of conservation interest may also increase the spread and impact of antagonistic species (e.g., predators and pathogens), foster negative effects of edges, increase invasion by exotic species, increase the spread of unwanted disturbances such as fire, or increase population synchrony and thus reduce persistence. We conducted a literature review and meta‐analysis to evaluate the prevalence of each of these negative effects. We found no evidence that corridors increase unwanted disturbance or non‐native species invasion; however, these have not been well‐studied concerns (1 and 6 studies, respectively). Other effects of corridors were more often studied and yielded inconsistent results; mean effect sizes were indistinguishable from zero. The effect of edges on abundances of target species was as likely to be positive as negative. Corridors were as likely to have no effect on antagonists or population synchrony as they were to increase those negative effects. We found 3 deficiencies in the literature. First, despite studies on how corridors affect predators, there are few studies of related consequences for prey population size and persistence. Second, properly designed studies of negative corridor effects are needed in natural corridors at scales larger than those achievable in experimental systems. Third, studies are needed to test more targeted hypotheses about when corridor‐mediated effects on invasive species or disturbance may be negative for species of management concern. Overall, we found no overarching support for concerns that construction and maintenance of habitat corridors may result in unintended negative consequences. Negative edge effects may be mitigated by widening corridors or softening edges between corridors and the matrix. Other negative effects are relatively small and manageable compared with the large positive effects of facilitating dispersal and increasing diversity of native species. Efectos Negativos Potenciales de los Corredores