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.     

Nesting Success of a Neotropical Migrant in a Multiple-Use, Forested Landscape

Abstract: We studied the nesting success of an individually marked population of Kentucky Warblers ( Oporornis formosus ), a species that nests in disturbed and undisturbed forests, in a heterogeneous, managed forest site in the Shawnee National Forest in southern Illinois from 1992 to 1995. We examined the effects of forest stand type (clearcuts of various ages, tree plantations, and older forest) and distance from habitat edges on rates of nest predation and brood parasitism by Brown-headed Cowbirds (   Molothrus ater ). Brood parasitism levels gradually decreased from 60% to 3% ( nests) over a distance of 2 km from an agricultural edge proximal to a known cowbird foraging site (a pig feedlot), but they did not vary with distance from any other kinds of edges or with forest stand type. Rates of nest predation ( nests) did not vary with distance from any edges, but they were significantly lower in older forest than within even-aged clearcuts, a tree plantation, and in successional vegetation adjacent to a residential facility. These results suggest that, even in fragmented landscapes with high overall levels of parasitism and nest predation, management practices within and immediately adjacent to forest tracts can affect the nesting success of some species, but not necessarily as a simple function of distance from edge. For the Kentucky Warbler, our results suggest that a management strategy that avoids even-age silviculture and leaves core stands of older forest far from cowbird feeding areas can increase nesting success to levels similar to those measured in more forested landscapes.     

Changes in Wildlife Communities Near Edges

Abstract: Wildlife managers and land managers have traditionally considered edges as beneficial to wildlife because species diversity generally increases near habitat edges. Explanations for this edge effect include greater vegetative complexity at edges or the simultaneous availability of more than one landscape element. However, edges can have negative consequences for wildlife by modifying distribution and dispersal and by increasing incidence of nest predation and parasitism Edges also may be detrimental to species requiring large undisturbed areas because increases in edge generally result in concommitant reductions in size and possible isolation of patches and corridots. Thus, both wildlife and land managers should be cautious when describing the benefits of edges to wildlife: particularly when dealing with species that require forest interiors.
Changes in wildlife communities associated with habitat edges are not easily assessed because defining edge species and measuring edge dimensions can be difficult in field studies Also, there is no general consensus as to how edge effect is best measured. Well-designed long-term studies of edges in various landscapes are needed (1) to better understand the positive and negative impacts of edges on wildlife communities, guilds, or key species, and (2) to effectively quantify edge effect and thereby develop management recommendations to improve the quality of edges for wildlife. Additional studies of edge effect are timely because greater amounts of edge will continue to be created in future landscapes due to extensive agriculture and other land-use-practices, and because developing knowledge in conservation biology and landscape ecology will facilitate multidisciplinary approaches to edge and landscape management for the benefit of wildlife.     

Fragmention by agriculture influences reproductive success of birds in a shrubsteppe landscape.

Shrubsteppe communities are among the most imperiled ecosystems in North America as a result of conversion to agriculture and other anthropogenic changes. In the Intermountain West of the United States, these communities support a unique avifauna, including several species that are declining and numerous others that are of conservation concern. Extensive research in the eastern and central United States and in Scandinavia suggests that fragmentation of formerly continuous forests and grasslands adversely affects reproductive success of birds, yet little is known of the potential effects on avian communities in Western shrublands. I used multi-model inference to evaluate the potential effects of local and landscape variables on nest predation and brood parasitism, and behavioral observations of color-banded birds to evaluate the potential effects of habitat fragmentation on seasonal reproductive success of passerines in the shrubsteppe of eastern Washington State, USA. Reproductive success of shrubsteppe-obligate passerines was lower in landscapes fragmented by agriculture than in continuous shrubsteppe landscapes. Daily survival rates for nests of Brewer's Sparrows (Spizella breweri; n=496) and Sage Thrashers (Oreoscoptes montanus; n=128) were lower in fragmented landscapes, and seasonal reproductive success (percentage of pairs fledging young) of Sage Sparrows (Amphispiza belli; n=146) and Brewer's Sparrows (n=59) was lower in fragmented landscapes. Rates of parasitism by Brown-headed Cowbirds (Molothrus ater) overall were low (4%) but were significantly greater in fragmented landscapes for Brewer's Sparrows, and parasitism resulted in fewer young fledged from successful nests. Simple models of population growth using landscape-specific fecundity and estimates of adult survival derived from return rates of banded male Sage Sparrows and Brewer's Sparrows suggest that fragmented shrubsteppe in Washington may be acting as a population sink for some species. Immediate conservation needs include halting further fragmentation of shrubsteppe, restoring low-productivity agricultural lands and annual grasslands to shrubsteppe where possible, and convincing the public of the intrinsic value of these imperiled ecosystems.     

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

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

Habitat Split as a Cause of Local Population Declines of Amphibians with Aquatic Larvae

Abstract:  Most amphibian species have biphasic life histories and undergo an ontogenetic shift from aquatic to terrestrial habitats. In deforested landscapes, streams and forest fragments are frequently disjunct, jeopardizing the life cycle of forest-associated amphibians with aquatic larvae. We tested the impact of habitat split—defined as human-induced disconnection between habitats used by different life-history stages of a species—on four forest-associated amphibian species in a severely fragmented landscape of the Brazilian Atlantic Forest. We surveyed amphibians in forest fragments with and without streams (referred to as wet and dry fragments, respectively), including the adjacent grass-field matrix. Our comparison of capture rates in dry fragments and nearby streams in the matrix allowed us to evaluate the number of individuals that engaged in high-risk migrations through nonforested habitats. Adult amphibians moved from dry fragments to matrix streams at the beginning of the rainy season, reproduced, and returned at the end of the breeding period. Juveniles of the year moved to dry fragments along with adults. These risky reproductive migrations through nonforested habitats that expose individuals to dehydration, predation, and other hazards may cause population declines in dry fragments. Indeed, capture rates were significantly lower in dry fragments compared with wet fragments. Declining amphibians would strongly benefit from investments in the conservation and restoration of riparian vegetation and corridors linking breeding and nonbreeding areas.     

How Corridors Reduce Indigo Bunting Nest Success

Abstract:  Corridors are a popular strategy to conserve biodiversity and promote gene flow in fragmented landscapes. Corridor effectiveness has been bolstered by the fact that no empirical field studies have shown negative effects on populations or communities. I tested the hypothesis that corridors increase nest predation in connected habitat fragments relative to unconnected fragments. I evaluated this hypothesis in a large-scale experimental system of open-habitat fragments that varied in shape and connectivity. Corridors increased nest predation rates in connected fragments relative to unconnected fragments with lower edge:area ratios. Nest predation rates were similar between connected and unconnected fragments with higher edge:area ratios. These results suggest that the increase in predator activity is largely attributable to edge effects incurred through the addition of a corridor. This is the first field study to demonstrate that corridors can negatively impact animal populations occupying connected fragments.     

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.     

Tests of landscape influence: nest predation and brood parasitism in fragmented ecosystems

The effects of landscape fragmentation on nest predation and brood parasitism, the two primary causes of avian reproductive failure, have been difficult to generalize across landscapes, yet few studies have clearly considered the context and spatial scale of fragmentation. Working in two river systems fragmented by agricultural and rural-housing development, we tracked nesting success and brood parasitism in > 2500 bird nests in 38 patches of deciduous riparian woodland. Patches on both river systems were embedded in one of two local contexts (buffered from agriculture by coniferous forest, or adjacent to agriculture), but the abundance of agriculture and human habitation within 1 km of each patch was highly variable. We examined evidence for three models of landscape effects on nest predation based on (1) the relative importance of generalist agricultural nest predators, (2) predators associated with the natural habitats typically removed by agricultural development, or (3) an additive combination of these two predator communities. We found strong support for an additive predation model in which landscape features affect nest predation differently at different spatial scales. Riparian habitat with forest buffers had higher nest predation rates than sites adjacent to agriculture, but nest predation also increased with increasing agriculture in the larger landscape surrounding each site. These results suggest that predators living in remnant woodland buffers, as well as generalist nest predators associated with agriculture, affect nest predation rates, but they appear to respond at different spatial scales. Brood parasitism, in contrast, was unrelated to agricultural abundance on the landscape, but showed a strong nonlinear relationship with farm and house density, indicating a critical point at which increased human habitat causes increased brood parasitism. Accurate predictions regarding landscape effects on nest predation and brood parasitism will require an increased appreciation of the multiple scales at which landscape components influence predator and parasite behavior.     

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.     

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.     

Effects of Pioneer Tree Species Hyperabundance on Forest Fragments in Northeastern Brazil

Abstract: Despite many studies on fragmentation of tropical forests, the extent to which plant and animal communities are altered in small, isolated forest fragments remains obscure if not controversial. We examined the hypothesis that fragmentation alters the relative abundance of tree species with different vegetative and reproductive traits. In a fragmented landscape (670 km²) of the Atlantic Forest of northeastern Brazil, we categorized 4056 trees of 182 species by leafing pattern, reproductive phenology, and morphology of seeds and fruit. We calculated relative abundance of traits in 50 1‐ha plots in three types of forest configurations: forest edges, small forest fragments (3.4–83.6 ha), and interior of the largest forest fragment (3500 ha, old growth). Although evergreen species were the most abundant across all configurations, forest edges and small fragments had more deciduous and semideciduous species than interior forest. Edges lacked supra‐annual flowering and fruiting species and had more species and stems with drupes and small seeds than small forest fragments and forest interior areas. In an ordination of species similarity and life‐history traits, the three types of configurations formed clearly segregated clusters. Furthermore, the differences in the taxonomic and functional (i.e., trait‐based) composition of tree assemblages we documented were driven primarily by the higher abundance of pioneer species in the forest edge and small forest fragments. Our work provides strong evidence that long‐term transitions in phenology and seed and fruit morphology of tree functional groups are occurring in fragmented tropical forests. Our results also suggest that edge‐induced shifts in tree assemblages of tropical forests can be larger than previously documented.     

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.     

Effects of Clearcutting on Habitat Use and Reproductive Success of the Ovenbird in Forested Landscapes

We studied Ovenbirds ( Seiurus aurocapillus ) in northern New Hampshire during 1992 and 1993 to determine whether edge-related changes in habitat use and reproductive success reported in fragmented landscapes exist in predominantly forested landscapes. Six study plots were placed adjacent to four recent clearcuts (2.1–5 ha) and extended 400 m into the forest interior. Nests, territories, and territorial males obtaining mates were equally distributed in edge (0–200 m) and interior (201–400 m) areas. Nest survival was higher in the forest interior in 1992 and for 1992 and 1993 combined. The proportion of pairs fledging ≥ 1 young, fledgling weight, and fledgling wing-chord did not differ between edge and interior in either year. Number of young fledged per pair was slightly lower in edge areas, but these differences were not significant. We conclude that clearcutting in extensively forested landscapes can affect Ovenbird reproductive success. Nevertheless, the effect on Ovenbird populations is moderated by the abundance of mature forest cover in the region and by the tendency of Ovenbirds to renest after initial nest failure.     

Potential Misuse of Avian Density as a Conservation Metric

Abstract: Effective conservation metrics are needed to evaluate the success of management in a rapidly changing world. Reproductive rates and densities of breeding birds (as a surrogate for reproductive rate) have been used to indicate the quality of avian breeding habitat, but the underlying assumptions of these metrics rarely have been examined. When birds are attracted to breeding areas in part by the presence of conspecifics and when breeding in groups influences predation rates, the effectiveness of density and reproductive rate as indicators of habitat quality is reduced. It is beneficial to clearly distinguish between individual‐ and population‐level processes when evaluating habitat quality. We use the term reproductive rate to refer to both levels and further distinguish among levels by using the terms per capita fecundity (number of female offspring per female per year, individual level) and population growth rate (the product of density and per capita fecundity, population level). We predicted how density and reproductive rate interact over time under density‐independent and density‐dependent scenarios, assuming the ideal free distribution model of how birds settle in breeding habitats. We predicted population density of small populations would be correlated positively with both per capita fecundity and population growth rate due to the Allee effect. For populations in the density‐dependent growth phase, we predicted no relation between density and per capita fecundity (because individuals in all patches will equilibrate to the same success rate) and a positive relation between density and population growth rate. Several ecological theories collectively suggest that positive correlations between density and per capita fecundity would be difficult to detect. We constructed a decision tree to guide interpretation of positive, neutral, nonlinear, and negative relations between density and reproductive rates at individual and population levels.     

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.     

Effects of Restoring Oak Savannas on Bird Communities and Populations

Abstract:  Efforts to restore and maintain oak savannas in North America, with emphasis on the use of prescribed fire, have become common. Little is known, however, about how restoration affects animal populations, especially those of birds. I compared the breeding densities, community structure, and reproductive success of birds in oak savannas maintained by prescribed fire (12 sites) with those in closed-canopy forests (13 sites). All sampling was conducted in Illinois (U.S.A.). Of the 31 bird species analyzed, 12 were more common in savannas, 14 were not affected by habitat structure, and 5 were more common in forest habitat. The species favored by disturbance and restoration included Northern Bobwhites ( Colinus virginianus ), Mourning Doves (  Zenaida macroura ), Red-headed Woodpeckers (  Melanerpes erythrocephalus ), Indigo Buntings (  Passerina cyanea ), and Baltimore Orioles ( Icterus galbula ). Those more common in closed-canopy forest included Ovenbirds ( Seiurus aurocapilla ) and Wood Thrushes (  Hylocichla mustelina ). Few species were unique to one type of habitat, but overall avian community structure in oak savannas and closed-canopy forests was generally distinctive. Estimates of nesting success (derived from 785 nests) revealed that 6 of the 13 species considered experienced greater productivity in the savanna habitat. Rates of brood parasitism were unaffected by restoration and habitat structure. Within savannas, tract size had little effect on breeding abundances and reproductive success. My results illustrate that restoration techniques can significantly affect the ecology of constituent animal populations and communities and have key implications regarding avian conservation and the management of forest habitat in fragmented landscapes. Small patches of forest habitat that regularly function as population sinks may offer far better prospects for birds if they are subjected to disturbance and ecosystem restoration.     

Possible causes and consequences of philopatry and breeding dispersal in red-backed shrikes <Emphasis Type="BoldItalic">Lanius collurio</Emphasis>

Studies of animal breeding dispersal have often focused on possible causes, whereas its adaptive significance has received less attention. Using an information-theoretic approach, we assessed predictions of four hypotheses relating to causes and consequences of breeding dispersal in a migratory passerine, the red-backed shrike Lanius collurio. As predicted by the reproductive performance hypothesis, probability of breeding dispersal in females (though not in males) decreased with increasing annual average number of fledglings produced in the past year, but there was no association with conspecific reproductive performance in either sex. The site choice hypothesis, stating that individuals disperse to improve breeding site quality, received support in males only, as dispersal probability was positively associated to a measure indicating low territory quality. The social constraints hypothesis, referring to dispersal in relation to intraspecific interactions, received little support in either sex. The predation risk hypothesis was hardly supported either. Consequences of dispersal were marginal in both sexes because neither fledgling production in females, nor territory quality in males improved after dispersal. In addition, males settled on territories closer to the forest edge than those occupied predispersal, which is opposite to the prediction of the predation risk hypothesis. We conclude that own reproductive success was the major factor determining dispersal behavior in females, whereas territory quality and possibly predation risk were most important in males. Overall, breeding dispersal appeared not to be adaptive in this dense population inhabiting an optimal habitat.     

The Early Development of Forest Fragmentation Effects on Birds

The early development of forest fragmentation effects on forest organisms is poorly understood partly because most fragmentation studies have been done in agricultural or suburban landscapes, long after the onset of fragmentation. We develop a temporal model of forest fragmentation effects on densities of forest-breeding birds and provide data from an active industrial forest landscape to test the model. The model and our empirical data indicate that densities of several forest-dwelling bird species can increase within a forest stand soon after the onset of fragmentation as a result of displaced individuals packing into remaining habitat. Along with higher densities in the newly formed fragments, pairing success in one species, the Ovenbird ( Seiurus aurocapillus ), was lower in fragments than nonfragments, possibly due to behavioral dysfunction resulting from high densities. Thus, density was inversely related to productivity. The duration and extent of increased densities following onset of fragmentation depends on many factors, including the sensitivity of a species to edge and area effects, the duration and rate of habitat loss and fragmentation, and the proximity of a forest stand to the disturbance. Incipient forest fragmentation may affect populations differently from later stages of fragmentation when the geometry of the landscape has reached a more stable configuration. Our model and data indicate, for reasons unrelated to traditional edge effects, that large tracts of forest can be important because they are relatively free from the variety of plant and animal population dynamics that might take place near new edges, including the encroachment of individuals displaced by habitat loss.