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.     

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

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

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.     

Effectiveness of Corridors Relative to Enlargement of Habitat Patches

Abstract:  The establishment of biological corridors between two otherwise isolated habitat patches is a common yet contentious strategy for conserving populations in fragmented landscapes. We compared the effectiveness of corridors with the effectiveness of an alternate conservation strategy, the enlargement of existing habitat patches. We used a spatially explicit population model that simulated population size in two kinds of patches. One patch had a corridor that connected it to a larger "source" patch and the other patch was unconnected and enlarged at the periphery by an area the same size as the corridor. Patch isolation, corridor width, patch size, and the probability that individuals would cross the border from habitat to matrix were varied independently. In general, population size was greater in enlarged patches than in connected patches when patches were relatively large and isolated. Corridor width and the probability of crossing the border from habitat to matrix did not affect the relative benefit of corridors versus patch enlargement. Although biological corridors may mitigate potential effects of inbreeding depression at long time scales, our results suggest that they are not always the best method of conserving fragmented populations.     

Gap-Crossing Decisions by the Red Squirrel, a Forest-Dependent Small Mammal

Abstract:  Forest-associated species in fragmented landscapes must traverse potentially inhospitable gaps to move between habitat patches. Although conservation biologists advocate connecting patches with corridors or improving the matrix to make it suitable for movement, little is known about the factors influencing gap-crossing decisions for most species. We investigated gap crossing by the red squirrel ( Tamiasciurus hudsonicus ) in logged landscapes in southeastern Alaska, where the species avoids microhabitats associated with gaps created by clearcutting. We released individuals across clearcuts and determined the routes they took home with tracking spools and radio telemetry. Of 36 adult red squirrels translocated across six clearcuts, 14 crossed clearcuts to reach home. Squirrels were more likely to cross clearcuts if the detour efficiency (distance to home crossing gap divided by distance of forested detour) was low, indicating an ability to compare distances along alternate routes and travel costs or risks in different habitats. No other landscape metrics, such as gap size or crossing distance, predicted crossing behavior. Red squirrels of low body mass were more likely to cross clearcuts, where the probability of encountering conspecifics is low. Distance predicted route choice for squirrels detouring around clearcuts. Indirect evidence suggests that perceived predation risk, energetic costs, or both are higher in clearcuts. Detour efficiency reportedly influences the gap-crossing decisions of some forest-associated birds, but this is the first demonstration of its role in gap-crossing decisions by a mammal.     

Effects of matrix characteristics and interpatch distance on functional connectivity in fragmented temperate rainforests

The connectivity of remnant patches of habitat may affect the persistence of species in fragmented landscapes. We evaluated the effects of the structural connectivity of forest patches (i.e., distance between patches) and matrix class (land-cover type) on the functional connectivity of 3 bird species (the White-crested Elaenia [Elaenia albiceps], the Green-backed Firecrown Hummingbird [Sephanoides sephaniodes], and the Austral Thrush [Turdus falklandii]). We measured functional connectivity as the rate at which each species crossed from one patch to another. We also evaluated whether greater functional connectivity translated into greater ecological connectivity (dispersal of fruit and pollen) by comparing among forest patches fruit set of a plant pollinated by hummingbirds and abundance of seedlings and adults of 2 plants with bird- and wind-dispersed seeds. Interpatch distance was strongly associated with functional connectivity, but its effect was not independent of matrix class. For one of the bird-dispersed plants, greater functional connectivity for White-crested Elaenias and Austral Thrushes (both frugivores) was associated with higher densities of this plant. The lack of a similar association for the wind-dispersed species suggests this effect is linked to the dispersal vector. The abundance of the hummingbird-pollinated species was not related to the presence of hummingbirds. Interpatch distance and matrix class affect animal movement in fragmented landscapes and may have a cascading effect on the distribution of some animal-dispersed species. On the basis of our results, we believe effort should be invested in optimizing patch configuration and modifying the matrix so as to mitigate the effects of patch isolation in fragmented landscapes.     

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.     

Comparison of Two Types of Metapopulation Models in Real and Artificial Landscapes

Abstract: Application of metapopulation models is becoming increasingly widespread in the conservation of species in fragmented landscapes. We provide one of the first detailed comparisons of two of the most common modeling techniques, incidence function models and stage-based matrix models, and test their accuracy in predicting patch occupancy for a real metapopulation. We measured patch occupancies and demographic rates for regional populations of the Florida scrub lizard (   Sceloporus woodi ) and compared the observed occupancies with those predicted by each model. Both modeling strategies predicted patch occupancies with good accuracy ( 77–80%) and gave similar results when we compared hypothetical management scenarios involving removal of key habitat patches and degradation of habitat quality. To compare the two modeling approaches over a broader set of conditions, we simulated metapopulation dynamics for 150 artificial landscapes composed of equal-sized patches (2–1024 ha) spaced at equal distances (50–750 m). Differences in predicted patch occupancy were small to moderate (<20%) for about 74% of all simulations, but 22% of the landscapes had differences openface> 50%. Incidence function models and stage-based matrix models differ in their approaches, assumptions, and requirements for empirical data, and our findings provide evidence that the two models can produce different results. We encourage researchers to use both techniques and further examine potential differences in model output. The feasibility of obtaining data for population modeling varies widely among species and limits the modeling approaches appropriate for each species. Understanding different modeling approaches will become increasingly important as conservation programs undertake the challenge of managing for multiple species in a landscape context.     

Habitat patch shape, not corridors, determines herbivory and fruit production of an annual plant

Habitat corridors confer many conservation benefits by increasing movement of organisms between habitat patches, but the benefits for some species may exact costs for others. For example, corridors may increase the abundance of consumers in a habitat to the detriment of the species they consume. In this study we assessed the impact of corridors on insect herbivory of a native plant, Solanum americanum, in large-scale, experimentally fragmented landscapes. We quantified leaf herbivory and assessed fruit production as a proxy for plant fitness. We also conducted field surveys of grasshoppers (Orthoptera), a group of abundant, generalist herbivores that feed on S. americanum, and we used exclosure cages to explicitly link grasshopper herbivory to fruit production of individual S. americanum. The presence of corridors did not increase herbivory or decrease plant fruit production. Likewise, corridors did not increase grasshopper abundance. Instead, patches in our landscapes with the least amount of edge habitat and the greatest amount of warmer "core" area had the highest levels of herbivory, the largest cost to plant fruit production as a result of herbivory, and the most grasshoppers. Thus habitat quality, governed by patch shape, can be more important than connectivity for determining levels of herbivory and the impact of herbivory on plant fitness in fragmented landscapes.     

Determinants of local extinction and turnover rates in urban bird communities.

Studying the effects of urbanization on the dynamics of communities has become a priority for biodiversity conservation. The consequences of urbanization are mainly an increased fragmentation of the original landscapes associated with a decrease in the amount of favorable habitats and an increased pressure of human activities on the remaining patches suitable for wildlife. Patterns of bird species richness have been studied at different levels of urbanization, but little is known about the temporal dynamics of animal communities in urban landscapes. In particular, urbanization is expected to have stronger negative effects on migratory breeding bird communities than on sedentary ones, which should lead to different patterns of change in composition. Using an estimation method accounting for heterogeneity in species detection probability and data collected between 2001 and 2003 within a suburban area near the city of Paris, France, we tested whether these communities differ in their local extinction and turnover rates. We considered the potential effects of patch size and distance to Paris' center as a measure of the degree of urbanization around the patches. As expected, local rates of extinction and turnover were higher for migratory than for sedentary species, and they were negatively related to patch size for migratory species. Mean species richness of the sedentary species increased during the study period and their local turnover rate was negatively related to the distance to the urban core, showing a trend to colonize the most urban patches. These results highlight the very dynamic nature of the composition of some local bird communities in fragmented habitats and help to identify factors affecting colonization and extinction.     

Relative Importance of the Area and Shape of Patches to the Diversity of Multiple Taxa

Abstract: Although enhancing reserve shape has been suggested as an alternative to enlarging nature reserves, the importance of reserve shape relative to reserve area remains unclear. Here we examined the relative importance of area and shape of forest patches to species richness, species composition, and species abundance (abundance of each species) for 3 taxa (33 birds, 41 butterflies, and 91 forest‐floor plants) in a fragmented landscape in central Hokkaido, northern Japan. We grouped the species according to their potential edge responses (interior‐, neutral‐, and edge‐species groups for birds and forest‐floor plants, woodland‐ and open‐land‐species groups for butterflies) and analyzed them separately. We used a shape index that was independent of area as an index of shape circularization. Hierarchical partitioning and variation partitioning revealed that patch area was generally more important than patch shape for species richness and species composition of birds and butterflies. For forest‐floor plants, effects of patch area and shape were small, whereas effects of local forest structure were large. Patch area and circularization generally increased abundances of interior species of birds and forest‐floor plants and woodland species of butterflies. Nevertheless, only patch circularization increased abundances of 1 woodland species of butterfly and 2 and 6 interior species of birds and forest‐floor plants, respectively. We did not find any significant interaction effects between patch area and shape. Our results suggest that although reserves generally should be large and circular, there is a trade‐off between patch area and shape, which should be taken into consideration when managing reserves.     

Desert-dwelling Mountain Sheep: Conservation Implications of a Naturally Fragmented Distribution

Abstract: Mountain sheep (Ovis canadensis) are closely associated with steep, mountainous, open terrain. Their habitat consequently occurs in a naturally fragmented pattern, often with substantial expanses of unsuitable habitat between suitable patches; the sheep have been noted to be slow colonizers of vacant suitable habitat. As a result, resource managers have focused on (1) conserving "traditional" mountainous habitats, and (2) forced colonization through reintroduction. Telemetry studies in desert habitats have recorded more intermountain movement by desert sheep than was previously thought to OCCUT. Given the heretofore unrecognized vagility of mountain sheep, we argue that existing corridors of "nontraditional" habitat connecting mountain ranges be given adequate conservation consideration. Additionally, small areas of mountainous habitat that an? not permanently occupied but that may serve as "stepping stones" within such corridors must be recognized for their potential importance to relatively isolated populations of mountain sheep. We discuss the potential importance of such corridors to other large, vagile species.     

A Meta‐Analytic Review of Corridor Effectiveness

Abstract: Using corridors for conservation is increasing despite a lack of consensus on their efficacy. Specifically, whether corridors increase movement of plants and animals between habitat fragments has been addressed on a case‐by‐case basis with mixed results. Because of the growing number of well‐designed experiments that have addressed this question, we conducted a meta‐analysis to determine whether corridors increase movement; whether corridor effectiveness differs among taxa; how recent changes in experimental design have influenced findings; and whether corridor effectiveness differs between manipulative and natural experiments. To conduct our meta‐analysis, we analyzed 78 experiments from 35 studies using a conservative hierarchical Bayesian model that accounts for hierarchical and sampling dependence. We found a highly significant result that corridors increase movement between habitat patches by approximately 50% compared to patches that are not connected with corridors. We found that corridors were more important for the movement of invertebrates, nonavian vertebrates, and plants than they were for birds. Recent methodological advances in corridor experiments, such as controlling for the area added by corridors, did not influence whether corridors increased movement, whereas controlling for the distance between source and connected or unconnected recipient patches decreased movement through corridors. After controlling for taxa differences and whether studies controlled for distance in experimental design, we found that natural corridors (those existing in landscapes prior to the study) showed more movement than manipulated corridors (those created and maintained for the study). Our results suggest that existing corridors increase species movement in fragmented landscapes and that efforts spent on maintaining and creating corridors are worthwhile.     

Geometrical Factors and Metapopulation Dynamics of the Bush Cricket, Metrioptera bicolor Philippi (Orthoptera: Tettigoniidae)

This paper presents a metapopulation study of the bush cricket, Metrioptera bicolor , living in a recently fragmented landscape. The species inhabits grass and heathland patches of varying area and isolation. Analyses are made of how these geometrical factors affect local population size and density, distribution pattern, and the probability of local extinction and colonization. The proportion of available patches occupied varied between 72 and 79% during 1985–1990. Unoccupied patches were smaller and more isolated than those that were occupied. Patches where populations became extinct during this period were smaller than those with persisting populations. Since local population size was well correlated with patch area, it was clear that stochastic extinctions only occurred in small populations. Critical patch size for population extinction was approximately half a hectare. Colonized patches were less isolated than those that had not been colonized. Critical inter-patch distance for colonization was about 100 meters. The turnover was restricted to an identifiable share of the available patches. Only 33% of the patches were so small that extinction due to stochastic causes could be considered highly probable. This metapopulation will therefore most likely persist over a considerable period in its present spatial structure. There are apparent threats of further fragmentation, however, and nothing is known about the likelihood of large-scale extinctions resulting from extremely unfavorable weather conditions. Nevertheless, our results show that it is appropriate to include geometrical factors in metapopulation models.     

Forest Fragmentation Increases Nest Predation in the Eurasian Treecreeper

Abstract:  We used long-term breeding data to monitor the influences of fragmentation and habitat composition at different spatial scales on the reproductive success of Eurasian Treecreepers ( Certhia familiaris ) breeding in nest boxes. We collected data from the same forest patches (2.7–65.1 ha in size) during seven breeding seasons. Nest predation varied considerably over the years and was the primary cause of nesting failure (mean annual rate of 21.6 ± 12.8%). Nest predation explained most of the variation in fledgling production during the study period. Landscape-level fragmentation (radius of 500 m from territory center) affected nest predation more than did fragmentation on the territory scale (radius of 200 m from territory center). In general, nest loss due to predation in fragmented landscapes (32.4%) was almost threefold that of less fragmented (12.0%) landscapes. Of the habitat variables, predation rate correlated positively with the density of edges between forest and open land and with the proportion of sapling stands on the spatial scale of 500 m around a nest. In the core area of a territory (radius of 30 m from territory center), a high density of trees increased the frequency of nest predation. Further, a high proportion of agricultural land close to a nest site increased nest losses of treecreepers, probably because of a high degree of mustelid predation. Our results showed that the spatial scale on which we examined nest predation is important and that even within moderately fragmented landscapes it is possible to detect fragmentation-related nest predation.     

A Comparison of Corridors and Intrinsic Connectivity to Promote Dispersal in Transient Successional Landscapes

Low-vagility organisms that specialize on transitory successional habitats may be especially dependent upon habitat connectivity to maintain population viability. We analyzed the theoretical intrinsic connectivity of successional landscapes (i.e., the natural juxtaposition of similar habitats that allows dispersal) as a function of patch geometry coupled with the disperser's habitat specificity. Habitat specialists living in poorly connected landscapes (approximating hexagonal patches) have only a 26.5% chance of colonizing a new site when their resident patch becomes unsuitable. In contrast, generalists living in well connected landscapes can virtually always colonize a new site when needed. We infer from our simulation that for some habitat specialists, such as the rare, endemic Florida scrub lizard (Sceloporus woodi), anthropogenic control of successional dynamics for commercial logging may significantly reduce intrinsic connectivity. Lizard population viability may now depend upon the extrinsic connectivity provided by artificial corridors. However, the use of corridors will not serve as a general solution to the problem of anthropogenically reduced intrinsic connectivity until key logistical design problems have been resolved. Moreover, efforts to enhance intrinsic connectivity by modifying patch geometry may produce undesirable edge effects and conflict with old-growth preservation. Future research should focus on developing spatially explicit corridor models, documenting natural levels of intrinsic connectivity, quantifying anthropogenic disruption of natural connectivity, and describing species-specific mechanisms of inter-patch dispersal.     

城市景观生态建设──以沈阳市为例

本文对城市进行了景观生态学分析，认为城市景观嵌块体小，数量多，景观高度破碎，道路廊道多，而缺少绿色廊道及绿色嵌块体，结合沈阳市，分析了景观质量及绿地现状，从而提出在城市景观生态建设中应增加绿色嵌块体及绿色廊道的数量。     

Value of Small Patches in the Conservation of Plant-Species Diversity in Highly Fragmented Rainforest

Abstract:  We evaluated the importance of small (<5 ha) forest patches for the conservation of regional plant diversity in the tropical rainforest of Los Tuxtlas, Mexico. We analyzed the density of plant species (number of species per 0.1 ha) in 45 forest patches of different sizes (1–700 ha) in 3 landscapes with different deforestation levels (4, 11, and 24% forest cover). Most of the 364 species sampled (360 species, 99%) were native to the region, and only 4 (1%) were human-introduced species. Species density in the smallest patches was high and variable; the highest (84 species) and lowest (23 species) number of species were recorded in patches of up to 1.8 ha. Despite the small size of these patches, they contained diverse communities of native plants, including endangered and economically important species. The relationship between species density and area was significantly different among the landscapes, with a significant positive slope only in the landscape with the highest deforestation level. This indicates that species density in a patch of a given size may vary among landscapes that have different deforestation levels. Therefore, the conservation value of a patch depends on the total forest cover remaining in the landscape. Our findings revealed, however, that a great portion of regional plant diversity was located in very small forest patches (<5 ha), most of the species were restricted to only a few patches (41% of the species sampled were distributed in only 1–2 patches, and almost 70% were distributed in 5 patches) and each landscape conserved a unique plant assemblage. The conservation and restoration of small patches is therefore necessary to effectively preserve the plant diversity of this strongly deforested and unique Neotropical region.     

Patch Isolation, Corridor Effects, and Colonization by a Resident Sparrow in a Managed Pine Woodland

The isolation of habitat patches is often cited as having a major impact on the dynamics of small populations occupying patches in a complex landscape. Few studies, however, have provided field data demonstrating that isolation has an identifiable effect on specific populations independent of other factors such as local habitat quality or that landscape factors such as corridors can alleviate such effects. We conducted field surveys of Bachman's Sparrow ( Aimophila aestivalis ) populations in regions, which we call linear landscapes, where suitable habitat patches were isolated to varying degrees from potential sources of dispersing birds. In these linear landscapes, isolated patches of habitat were less likely to be colonized than were nonisolated patches. We also found that corridor configurations of habitat patches improved the ability of sparrows to find and settle in newly created patches. These results suggest that, for species that do not disperse easily through inhospitable landscapes, habitat occupancy at a regional scale can be enhanced by careful landscape design and planning.