Does land use change influence predation of bird nests?

Worldwide, many areas of agricultural land which were once covered with native vegetation have been converted to tree plantations. Such landscape transformation can influence the dynamics of wildlife populations through, for example, altering rates of predation (e.g. predation of nests of birds). Nest predation can influence reproductive success and, in turn, may alter populations by affecting juvenile recruitment. We quantified predation of bird nests in woodland remnants surrounded by two types of land use, grazing farmland and exotic Radiata pine (Pinus radiata) plantation. We also examined differences in predation rates between artificial and natural nests. We found both artificial and natural nests were more susceptible to nest predation in woodland remnants surrounded by a pine plantation than in woodland remnants located within farmland. Our study suggests that higher levels of nest predation may reduce occupancy of woodland remnants by small‐bodied birds over time, including species of conservation concern. This may have been occurred as a result of the conversion of semi‐cleared grazing land to exotic pine plantation.     

Compounding effects of habitat fragmentation and predation on bird nests

Habitat fragmentation and invasive species are two of the greatest threats to species diversity worldwide. This is particularly relevant for oceanic islands with vulnerable endemics. Here, we examine how habitat fragmentation influences nest predation by Rattus spp. on cup‐nesting birds in Samoan forests. We determined models for predicting predation rates by Rattus on artificial nests at two scales: (i) the position of the bird's nest within the landscape (e.g. proximity to mixed crop plantations, distance to forest edge); and (ii) the microhabitat in the immediate vicinity of the nest (e.g. nest height, ground cover, slope). Nest cameras showed only one mammal predator, the black rat (Rattus rattus), predating artificial nests. The optimal model predicting nest predation rates by black rats included a landscape variable, proximity to plantations and a local nest site variable, the percentage of low (<15 cm) ground cover surrounding the nest tree. Predation rates were 22 ± 13% higher for nests in forest edges near mixed crop plantations than in edges without plantations. In contrast, predation rates did not vary significantly between edge habitat where the matrix did not contain plantations, and interior forest sites (>1 km from the edge). As ground cover reduced, nest predation rates increased. Waxtags containing either coconut or peanut butter were used as a second method for assessing nest predation. The rates at which these were chewed followed patterns similar to the predation of the artificial nests. Rural development in Samoa will increase the proportion of forest edge near plantations. Our results suggest that this will increase the proportion of forest birds that experience nest predation from black rats. Further research is required to determine if rat control is needed to maintain even interior forest sites populations of predator‐sensitive bird species on South Pacific islands.     

Different nest predator faunas and nest predation risk on ground and shrub nests at forest ecotones: an experiment and a review

This study examined predator faunas of artificial ground and shrub nests and whether nest predation risk was influenced by nest site, proximity to forest edge, and habitat structure in 38 grassland plots in south-central Sweden. There was a clear separation of predator faunas between shrub and ground nests as identified from marks in plasticine eggs. Corvids accounted for almost all predation on shrub nests whereas mammals mainly depredated ground nests. Nest predation risk was significantly greater for shrub than for ground nests at all distances (i.e. 0, 15 and 30 m) from the forest edge. However, nest predation risk was not significantly related to distance to forest edge, but significantly increased with decreasing distance to the nearest tree. Different corvid species robbed nests at different distances from the forest edge, with jays robbing nests closest to edges. We conclude that the relationship between the predation risk of grassland bird nests and distance to the forest edge mainly depends on the relative importance of different nest predator species and on the structure of the forest edge zone. A review of published articles on artificial shrub and ground nest predation in the temperate zone corroborated the results of our own study, namely that shrub nests experienced higher rates of depredation in open habitats close to the forest edge and that avian predators predominantly robbed shrub nests. Furthermore, the review results showed that predation rates on nests in general are highest <50 m inside the forest and lower in open as well as forest interior habitats (≥50 m from the edge). Received: 16 March 1998 / Accepted: 30 July 1998     

Nest Predation Deviates from Nest Predator Abundance in an Ecologically Trapped Bird

In human-modified environments, ecological traps may result from a preference for low-quality habitat where survival or reproductive success is lower than in high-quality habitat. It has often been shown that low reproductive success for birds in preferred habitat types was due to higher nest predator abundance. However, between-habitat differences in nest predation may only weakly correlate with differences in nest predator abundance. An ecological trap is at work in a farmland bird (Lanius collurio) that recently expanded its breeding habitat into open areas in plantation forests. This passerine bird shows a strong preference for forest habitat, but it has a higher nest success in farmland. We tested whether higher abundance of nest predators in the preferred habitat or, alternatively, a decoupling of nest predator abundance and nest predation explained this observed pattern of maladaptive habitat selection. More than 90% of brood failures were attributed to nest predation. Nest predator abundance was more than 50% higher in farmland, but nest predation was 17% higher in forest. Differences between nest predation on actual shrike nests and on artificial nests suggested that parent shrikes may facilitate nest disclosure for predators in forest more than they do in farmland. The level of caution by parent shrikes when visiting their nest during a simulated nest predator intrusion was the same in the two habitats, but nest concealment was considerably lower in forest, which contributes to explaining the higher nest predation in this habitat. We conclude that a decoupling of nest predator abundance and nest predation may create ecological traps in human-modified environments.     

Habitat edges affect patterns of artificial nest predation along a wetland-meadow boundary

Wetland habitats are among the most endangered ecosystems in the world. However, little is known about factors affecting the nesting success of birds in pristine grass-dominated wetlands. During three breeding periods we conducted an experiment with artificial ground nests to test two basic mechanisms (the matrix and ecotonal effects) that may result in edge effects on nest predation in grass-dominated wetland habitats. Whereas the matrix effect model supposes that predator penetrate from habitat of higher predator density to habitat of lower predator density, thus causing an edge effect in the latter, according to the ecotonal effect model predators preferentially use edge habitats over habitat interiors. In addition, we tested the edge effect in a wetland habitat using artificial shrub nests that simulated the real nests of small open-cup nesting passerines. In our study area, the lowest predation rates on ground nests were found in wetland interiors and were substantially higher along the edges of both wetland and meadow habitat. However, predation was not significantly different between meadow and wetland interiors, indicating that both mechanisms can be responsible for the edge effect in wetland edges. An increased predation rate along wetland edges was also observed for shrub nests, and resembled the predation pattern of real shrub nests in the same study area. Though we are not able to distinguish between the two mechanisms of the edge effect found, our results demonstrate that species nesting in wetland edges bordering arable land may be exposed to higher predation. Therefore, an increase in the size of wetland patches that would lead to a reduced proportion of edge areas might be a suitable management practice to protect wetland bird species in cultural European landscapes.     

Nest Predation Risk on Ground and Shrub Nests in Forest Margin Areas of Sulawesi, Indonesia

Forest loss and fragmentation in Indonesia may seriously affect the survivorship of forest birds and lead to local extinction of bird populations. We used 786 artificial nests baited with quail eggs to examine the effect of habitat alteration on nest predation in Lore Lindu National Park, Sulawesi. Natural forest and four habitats of forest margin areas: forest edge, forest gardens, coffee plantations, and secondary forest, were studied. Two types of artificial nests, ground and shrub nests were placed in these habitats at two different locations for a period of 8 days. In addition, we used automatic cameras and cage-traps to identify the predators. Nests in shrubs experienced significantly higher predation rates in forest margin areas than in natural forest. Predation on ground nests did not differ significantly between these habitat types, but was significantly higher than that on shrub nests in each habitat except forest edge. Rodents were the most common predators of both nests, but shrub nests were also susceptible to Dwarf cuscus (Strigocuscus celebensis), squirrels, and tree snakes. The nest predation rates we found were among the highest found in tropical rainforests, probably a consequence of the unique predator assemblages of Sulawesi. These results suggest that egg survival is negatively affected by human intervention and that human-induced habitats might have only limited importance for the conservation of Sulawesi's largely endemic understorey avifauna. These considerations might be important since forest margins comprise significant proportions of protected areas on Sulawesi and play an important role in future Park zoning concepts as well as in conservation-oriented land use management.     

Forest fragmentation and avian nest predation in forested landscapes

Summary The size of forest fragments, the use of land bordering fragments, and the distance of nests from an edge all affect the frequency of predation upon bird nests in Maine (USA), an area where the forest has been fragmented by roads, but not significantly reduced in area. We placed artificial nests containing quail eggs in forests of different sizes and at various distances from the edge to test which of these factors was most important in describing predation. Predation was greatest in small tracts surrounded completely by land. Large areas and those bordered on at least one side by a large water body had lower predation rates. This suggests that influx of predators from nearby habitats may be responsible for much of the nest predation in forest fragments.     

Nest predation in Reed Buntings Emberiza schoeniclus: an experimental study

Predation is a major cause of nest failure in many bird species. High levels of nest loss may be a consequence of habitat fragmentation, leading to increased amounts of edge habitat. Yet the evidence for generally high nest predation rates along edges in fragmented landscapes is ambiguous. Using real nests of Reed Buntings Emberiza schoeniclus in which artificial Reed Bunting and real Japanese Quail Coturnix japonica eggs were placed, we experimentally tested for edge effects on nest predation in highly fragmented reed Phragmites sp. habitats in the Swiss lowlands. We also examined seasonal patterns of predation and the impacts of nest visits by observers. We found evidence for an edge effect at the water-sided reed edge, with nests located closer to the water being more likely to be predated than those further away. Predation probability increased from early to late season, suggesting that nest predation may be density dependent. Probability of nest predation was only weakly influenced by whether or not a nest was visited. Our results suggest that the intensive reed management currently applied in Swiss nature reserves may result in unnaturally high levels of nest losses in the Reed Bunting, because reed bands are not wide enough to allow nest placement at a safe distance from reed edges.     

Herbaceous ground cover reduces nest predation in olive groves

Capsule Bare ground increases artificial nest predation in olive groves.

Aims To assess the effect of different soil management regimes on nest predation rates in olive groves.

Methods We performed nest predation experiments with artificial nests during the breeding season in 2013, in two areas of southern Spain. Each artificial nest (n?=?300) contained three quail Coturnix eggs, two of which were unmanipulated and the third one was emptied and injected with plaster. Predators were identified by marks on eggs filled with plaster.

Results Ground nests were significantly more depredated, irrespective of the presence of ground cover; tree nests were less depredated in fields with ground cover. There was a clear difference in nest predators of ground and tree nests. Rodents were the most frequent predators of tree nests.

Conclusion Lower predation rates of tree nests in orchards with ground cover are probably linked to a change in the foraging behaviour of rodents, which in these more complex habitats might be restricted by rodents' own risk of predation. This study underscores the important role of agricultural practices in preserving farmland bird communities, particularly tree-nesting species, suggesting that for this group, implementation of ground cover in olive groves might enhance breeding success by reducing nest predation rates.     

Protective nesting association between the Barred Warbler Sylvia nisoria and the Red-backed Shrike Lanius collurio: an experiment using artificial and natural nests

The pressure of predators may significantly affects the distribution pattern of nesting birds. Some individuals may reduce the risk of predation by nesting near other species with an aggressive nest defence. In the present study I tested the predator protection hypothesis using experimental (artificial nests) and observational (real nests) approaches on two ecologically similar passerine birds–the Barred Warbler Sylvia nisoria and the Red-backed Shrike Lanius collurio. Studies have been conducted in eastern Poland in two types of habitat: river valley and farmland. The main predators of natural and artificial nests were birds, and to a lesser extent, also mammals. I found wide variation level of predation of both types of nests in different years. Nest survival rate of artificial nests was significantly lower in the farmland than in the river valley and in natural nests I observed reverse pattern. According to the predictions of the predator protection hypothesis the survival rates of the natural and artificial nests were significantly higher in territories of individuals breeding in the protective nesting association. This type of interspecific positive interaction between two associate species can be classified as facultative mutualism.     

Egg predation in forest bird communities on islands and mainland

Summary To examine if differences in egg predation rates could explain differences in bird community composition, egg predation was studied in two years on small islands in a South Swedish lake and on the nearby mainland using both natural and artificial nests.In plots with similar vegetation, the combined density of ground- and tree-nesting bird species did not differ between the islands and the mainland. Egg predation rates were similar on islands and the mainland for natural Turdus nests in two years, and for artificial Turdus and Phylloscopus nests. Unmarked and unvisited experimental nests suffered similar rate of egg predation as marked and visited nests. Egg predation rates were higher on natural nests when artificial nests were also put out, increasing the total nest density. Initial egg predation rates in artificial nests were also higher than later when nest density had decreased by 75%.The egg predators involved differed for artificial Phylloscopus nests between the islands and the mainland. Small mammals were apparently responsible for 29% of the predation on the mainland, but none on the islands. Artificial Turdus nests near crow nests suffered from a higher egg predation rate than nests further away from crow nests. Daily survival rates of Turdus nests increased from the laying to the incubation and further to the fledging state.Egg predation can not explain differences in bird community composition between islands and mainland in the present case.     

千岛湖栖息地片段化效应对鸟类巢捕食风险的影响

为了解栖息地片段化对鸟类巢捕食风险的影响,我们于2010年4-8月,在千岛湖地区选取16个岛屿,分别在岛屿边缘和内部区域用放置鸡蛋(大型卵)和鹌鹑(Coturnix japonica)蛋(小型卵)的方法进行人工模拟地面巢试验,研究不同体积大小鸟卵的巢捕食率及其差异,分析巢捕食率与岛屿面积、隔离度、形状指数和植物物种丰富...     

Tropical forest fragmentation and nest predation – an experimental study in an Eastern Arc montane forest, Tanzania

When a habitat becomes fragmented and surrounded by another habitat this generally causes an increase in predation pressure at habitat transitions, often referred to as an edge effect. Edge effect in the form of enhanced nest predation intensities is one of the most cited explanations for bird population declines in fragmented landscapes. Here, we report results from a nest predation experiment conducted in a tropical montane forest landscape in the Uzungwa Mts., Tanzania. Using artificial nests with chicken eggs, we determined predation rates across a fragmentation gradient. The proportion of indigenous forest in four landscapes used in the study were 0.29, 0.58, 0.75 to 1.0. Nest predation intensities on artificial nests were about 19% higher inside intact forest than at edges in fragmented forest landscapes. Furthermore, predation intensities were relatively constant across a forest fragmentation gradient. Our results thus challenge the applicability and generality of the edge effect, derived from studies almost exclusively conducted in temperate regions rather than tropical forest ecosystems. Nest predation levels differences between tropical montane forest and that reported in other forest ecosystems are discussed.     

An ecological paradox: More woodland predators and less artificial nest predation in landscapes colonized by noisy miners

Many passerine bird populations, particularly those that have open‐cup nests, are in decline in agricultural landscapes. Current theory suggests that an increase in habitat generalist predators in response to landscape change is partially responsible for these declines. However, empirical tests have failed to reach a consensus on how and through what mechanisms landscape change affects nest predation. We tested one hypothesis, the Additive Predation Model, with an artificial nest experiment in fragmented landscapes in southern Queensland, Australia. We employed structural equation modelling of the influence of the relative density of woodland and habitat generalist predators and landscape features at the nest, site, patch and landscape scales on the probability of nest predation. We found little support for the Additive Predation Model, with no significant influence of the density of woodland predators on the probability of nest predation, although landscape features at different spatial scales were important. Within woodlands fragmented by agriculture in eastern Australia, the presence of noisy miner colonies appears to influence ecological processes important for nest predation such that the Additive Predation Model does not hold. In the absence of colonies of the aggressive native bird, the noisy miner, the influence of woodland predators on the risk of artificial nest predation was low compared with that of habitat generalist predators. Outside noisy miner colonies, we found significant edge effects with greater predation rates for artificial nests within woodland patches located closer to the agricultural matrix. Furthermore, the density of habitat generalist predators increased with the extent of irrigated land‐use, suggesting that in the absence of noisy miner colonies, nest predation increases with land‐use intensity at the landscape scale.     

Conservation potential of semi-natural habitats for birds in intensively-used agricultural landscapes

Agricultural intensification resulted in substantial loss of farmland biodiversity. Semi-natural habitats may be viewed as potential buffers of these adverse impacts, but a rigorous assessment of their capacity for supporting farmland biodiversity is lacking. In this study, we explored conservation potential of two different types of semi-natural habitats for birds in intensively-used agricultural landscapes – farmland hedges (i.e., linear strips of shrubby and tree vegetation) and open scrubland (i.e., scattered shrubs and abandoned orchards). Specifically, we tested whether the abundance and species richness of birds differ between these habitats considering various species traits, such as habitat affinity (i.e., forest, farmland and urban species), diet specialization (i.e., animal eaters, plant eaters, and omnivores) and conservation status (Species of European Conservation Concern). We found that open scrubland hosted on average 37.9 bird species and 122.6 individuals per 1 km² of the transect, whereas farmland hedges hosted only 19 species and 61.8 individuals per 1 km² of the transect. However, results have substantially changed if we considered the area of suitable habitat into account. More specifically, open scrubland hosted more bird species and individuals when we considered open habitat species and the area of open habitats, whereas farmland hedges had higher species diversity and individuals of woodland bird species when we considered the area of woodland habitats. Similarly, analyses of habitat affiliations of individual species corresponded to the whole-community patterns; and revealed that several woodland bird species were mainly associated with farmland hedges (e.g., Chaffinch Fringilla coelebs, Common Nightingale Luscinia megarhynchos and Blackcap Sylvia atricapilla), whereas the open scrubland was preferred by open habitat bird species (e.g., Corn Bunting Emberiza calandra, Quail Coturnix coturnix and Skylark Alauda arvensis). These results demonstrate that semi-natural habitats, both open scrubland and farmland hedges, have large potential for promotion and conservation of bird communities within intensively used agricultural landscapes, as both may have represented suitable habitats for species with different ecological requirements. Therefore, management measures focused on the enlargement of the area of these habitats, in combination with suitable management (e.g., regulating the progress of natural succession in open scrubland; increasing structural diversity of existing farmland hedges), may substantially contribute to bird conservation within agricultural landscapes.     

Synergistic effects of habitat configuration and land-use intensity shape the structure of bird assemblages in human-modified landscapes across three major neotropical biomes

Maximizing biodiversity persistence in heterogeneous human-modified landscapes is hindered by the complex interactions between habitat quality and configuration of native and non-native habitats. Here we examined these complex interactions considering avian diversity across 26 sampling sites, each of which comprised of three sampling points located across a gradient of disturbance: core native habitat fragment, fragment edge, and non-native adjacent matrix. The 78 sampling points were further nested within three neotropical biomes—Amazonia, Cerrado and Pantanal—in central-western Brazil. Matrix type consisted of cattle pastures in the Amazon and teak plantations in the Pantanal and Cerrado. We considered the interactive effects of (1) disturbance-context: fragment core, edge and adjacent matrix, (2) matrix type: tree plantation or cattle pastures, both subject to varying land-use intensity, and (3) native habitat configuration (fragment size, shape and isolation) on bird species richness, abundance and composition. Based on point-count surveys, we recorded 210 bird species. Bird species richness and abundance declined across the disturbance gradient, while genus composition only differed within the adjacent matrix, particularly cattle-pastures. The effect of native habitat area was positive but only detected at fragment edges. Overall bird diversity increased at sites characterized by higher availability of either relict trees within pasture landscapes or old-growth trees within teak plantation landscapes. The core of native fragments played a primary role in ensuring the persistence of bird diversity, regardless of fragment size. In contrast to pastures, tree plantations likely harbour a higher proportion of forest-dependent species while bird diversity can be further enhanced by reduced management intensity in both matrix types. Strategies to maximize avian persistence should not only include retaining native habitats, but also maximizing the size of core native habitats. Likewise, more structurally complex matrix types should be encouraged while maintaining low levels of land-use intensity.

     

Effects of Habitat Structure and Adjacent Habitats on Birds in Tropical Rainforest Fragments and Shaded Plantations in the Western Ghats, India

As large nature reserves occupy only a fraction of the earth’s land surface, conservation biologists are critically examining the role of private lands, habitat fragments, and plantations for conservation. This study in a biodiversity hotspot and endemic bird area, the Western Ghats mountains of India, examined the effects of habitat structure, floristics, and adjacent habitats on bird communities in shade-coffee and cardamom plantations and tropical rainforest fragments. Habitat and birds were sampled in 13 sites: six fragments (three relatively isolated and three with canopy connectivity with adjoining shade-coffee plantations and forests), six plantations differing in canopy tree species composition (five coffee and one cardamom), and one undisturbed primary rainforest control site in the Anamalai hills. Around 3300 detections of 6000 individual birds belonging to 106 species were obtained. The coffee plantations were poorer than rainforest in rainforest bird species, particularly endemic species, but the rustic cardamom plantation with diverse, native rainforest shade trees, had bird species richness and abundance comparable to primary rainforest. Plantations and fragments that adjoined habitats providing greater tree canopy connectivity supported more rainforest and fewer open-forest bird species and individuals than sites that lacked such connectivity. These effects were mediated by strong positive effects of vegetation structure, particularly woody plant variables, cane, and bamboo, on bird community structure. Bird community composition was however positively correlated only to floristic (tree species) composition of sites. The maintenance or restoration of habitat structure and (shade) tree species composition in shade-coffee and cardamom plantations and rainforest fragments can aid in rainforest bird conservation in the regional landscape.     

Prevalence of nest predators in a sub-Arctic ecosystem

High nest loss is an important driver of gallinaceous bird population dynamics. Identifying factors determining the spatial distribution of potential nest predators and thereby indirectly risk of nest losses is therefore essential. The aim of this 1-year study was to estimate relative predation rates on artificial ground nests in willow ptarmigan (Lagopus lagopus) habitats, along replicate altitudinal gradients (transects, n?=?60) spanning from sub-Arctic birch forest to the low-alpine tundra in three locations in northern Norway. In each transect, one artificial nest (track board) was placed in three different habitats: (1) birch forest, (2) edge between birch forest and low-alpine tundra and (3) low-alpine tundra. Total predation rates over all habitats within locations ranged from 47.4% to 77.5% and did not vary systematically in space and time. The average predation rate by avian predators was consistently high (58%), and mammalian predation rate was consistently low (5.6%). The consistently high level of predation inflicted by birds was mainly due to omnipresent corvids, especially the hooded crow (Corvus cornix). Analysis of species-specific predation rates showed that habitat and location effects were insignificant for all species, except for raven (Corvus corax) that showed clearly higher predation in one of the locations. The results indicate that from the perspective of the spatial distribution of potential nest predators in sub-Arctic birch forest, ground nesting birds like willow ptarmigan should not be expected to be selective with respect to nesting habitat in the ecotone between birch forest and the low-alpine tundra.     

Predation on artificial avian nests is higher in forests bordering small anthropogenic openings

Habitat edges alter the diversity of avian communities and are often associated with higher rates of nest predation. However, most previous studies on habitat edges have been conducted along long linear corridors or at the transition between large field and forest patches in agricultural systems. Less is known about predation rates when the habitat edge is the result of a small interior forest opening. We assessed predation rates on artificial nests mimicking ground and shrub nesters in Northern Michigan forests perforated by small clearings used previously for oil and gas extraction. Nests were placed at varying distances from the edges of these clearings, and in similar spatial arrangements within unfragmented interior forest plots. Predation rates increased in forests near edges, but significant impacts were limited to shrub nests. Markings on predated clay eggs indicated that the type of predation also differed. Scratch marks were the most prevalent egg indentation, but eggs with poked holes were twice as common near the forest edge. The increase in the number of poked eggs suggests that a higher density of avian predators occurred in forests near an edge. Predation rates at forest edges did not vary by distance from the forest edge. Surveys of the avian community revealed differences between edge and interior forests: American Crows Corvus brachyrhynchos and Blue Jays Cyanocitta cristata, two species known to predate bird nests, were more common near edges. Our results suggest that small forest openings alter the avian community and may adversely impact reproductive output in some species. If the alteration of these processes results in population‐level impacts, small forest perforations should be avoided when possible and reforestation of abandoned well‐pads should be encouraged.     

Edge effect on bird nest predation in the fragmented caldén (<Emphasis Type="Italic">Prosopis caldenia</Emphasis>) forest of central Argentina: an experimental analysis

Clearing of caldén (Prosopis caldenia) forests for agriculture and cattle raising in east-central La Pampa Province, central Argentina, has created a highly fragmented landscape, a condition that has resulted in adverse effects on birds in other forests, mainly through increased predation rates near forest edges. We evaluated bird nest predation rates using artificial nests, assessing the effects of forest fragment size, distance to the edge and nest height. We measured survival rate of 570 artificial nests located in trees, in bushes and on the ground, at different distances from the edge, in six forest fragments ranging in size from 2.1 to 117.6 ha, during two consecutive breeding seasons. Nest predation rates were significantly related with the number of days of exposition of the nest, nest height and distance to the edge, whereas fragment size and year of the experiment were not associated with predation rates. Ground nests were less likely to be predated than those located in bushes and trees. Predation rates decreased with the distance to the edge, showing a pattern consistent with the existence of an edge effect.