Artificial nest predation rates in tropical and temperate forests: a review of the effects of edge and nest site

Nest predation rates are believed to be higher in tropical than in temperate forests. This notion is central in explaining different life history traits of tropical and temperate birds, but it is not known whether this assertion is true for all nest sites, such as ground and shrub nests, and at different distances from forest edge. I reviewed 22 studies using artificial nest experiments which concurrently contrasted predation rates of ground and shrub nests in temperate and tropical forests and found, contrary to the current dogma, no overall difference in predation rates between regions. However, there was a significant interaction between region and nest site. Ground nest predation rates were significantly higher in the tropical region, while predation rates on shrub nests were similar between regions. Within the tropical region, ground nests had significantly higher predation rates than shrub nests. Elevated nest predation rates at forest edges were found in both temperate and tropical forests. The results may have great implications for expected patterns of avian life histories and for the effects of forest fragmentation in temperate and tropical regions. First, if nest predation affects avian life histories, my results predict ground-nesting species in tropical forests to have shorter nestling periods, more broods and smaller clutch sizes than shrub-nesting species. Second, vulnerability of ground- and shrub-nesting guilds is suggested to differ between regions due to differences in forest vegetation structure, and the composition of predator faunas and their specific responses to forest fragmentation. Data to test these hypotheses are limited, but agree with the results of this review.     

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.     

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     

Predation on artificial nests in relation to forest type: contrasting the use of quail and plasticine eggs

Previous studies of avian nest predation have focused on how human-induced changes in the landscape influence the frequency of predation However, natural variation in the abundance of predators due to their choice of habitat can also influence predation rate To determine if predation on artificial nests was influenced by forest stand type, we placed ground and shrub nests containing quail and plasticine eggs in contiguous coniferous, mixedwood and deciduous stands in the southern boreal mixedwood forest of central Canada Nest predators were identified using remotely triggered cameras and marks left in plasticine eggs, while the relative abundance of nest predators such as squirrels and corvids were estimated using acoustic-visual surveys Using the fate of quail eggs to calculate predation rate, we found that predation was significantly higher in coniferous (67%) than in deciduous (17%) or mixedwood (25%) forest, with similar predation on ground (37%) and shrub (29%) nests Using plasticine eggs to calculate predation rate, nests in coniferous forest still suffered higher rates of predation, although predation rates were 15–20% higher, and ground nests suffered significantly higher rates of predation than shrub nests Quail eggs seemed to suffer lower rates of predation because small mammals were unable to penetrate the shell, but could leave marks on plasticine eggs The higher predation rate in coniferous forest was likely caused by higher abundance of red squirrels Tamiasciurus hudsonicus , the presence of fishers Martes pennanti and a simplified understory which may have made it easier for predators to find nests relative to the deciduous and mixedwood forest Plasucine eggs provide new insights into nest predation by identifying predation events by smaller predators such as mice that are missed when using quail eggs     

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.     

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.     

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.     

Forest fragmentation relaxes natural nest predation in an Afromontane forest

Nest predation is widely regarded as a major driver underlying the population dynamics of small forest birds. Following forest fragmentation and the subsequent invasion by species from non-forested landscape matrices, shifts in predator communities may increase nest predation near forest edges. However, effects of human-driven habitat change on nest predation have mainly been inferred from studies with artificial nests, despite being regarded as poor surrogates for natural ones. We studied variation in predation rates, and relationships with timing of breeding and characteristics of microhabitats and fragments, on natural white-starred robin Pogonocichla stellata nests during three consecutive breeding seasons (2004–2007) in a Kenyan fragmented cloud forest. More than 70% of all initiated nests were predated during each breeding season. Predation rates nearly quadrupled between the earliest and the latest nests within a single breeding season, increased with distance to the forest edge, and decreased with the edge-to-area ratio of forest fragments. These spatial relationships oppose the traditional perception of edge and fragmentation effects on nest predation, but are in line with results from artificial nest experiments in other East African forests. In case of inverse edge and fragmentation effects on nest predation, such as shown in this study, species that tolerate edges for breeding may be affected positively, rather than negatively, by forest fragmentation, while the opposite can be expected for species restricted to the forest interior. The possibility of inverse edge effects, and its conservation implications, should therefore be taken into account when drafting habitat restoration plans.     

Cropland edge,forest succession,and landscape affect shrubland bird nest predation

The effects of habitat edges on nest survival of shrubland birds, many of which have experienced significant declines in the eastern United States, have not been thoroughly studied. In 2007 and 2008, we collected data on nests of 5 shrubland passerine species in 12 early successional forest patches in North Carolina, USA. We used model selection methods to assess the effect of distance to cropland and mature forest edge on nest predation rates and additionally accounted for temporal trends, nest stage, vegetation structure, and landscape context. For nests of all species combined, nest predation decreased with increasing distance to cropland edge, by nearly 50% at 250 m from the cropland edge. Nest predation of all species combined also was higher in patches with taller saplings and less understory vegetation, especially in the second year of our study when trees were 4–6 m tall. Predation of field sparrow (Spizella pusilla) nests was lower in landscapes with higher agricultural landcover. Nest predation risk for shrubland birds appears to be greater near agricultural edges than mature forest edges, and natural forest succession may drive patterns of local extirpation of shrubland birds in early successional forest patches. Thus, we suggest that habitat patches managed for shrubland bird populations should be considerably large or wide (>250 m) when adjacent to crop fields and maintained in structurally diverse early seral stages. © 2011 The Wildlife Society.     

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.     

Effects of Edge Habitat and Nest Characteristics on Depredation of Artificial Nests in Fragmented Australian Tropical Rainforest

Variation in nest predation levels associated with rainforest fragmentation (edge effects) was assessed in Australia's Wet Tropics bioregion. Artificial nests were placed in the forest understorey at seven edge sites where continuous forest adjoined pasture, seven interiors (about 1 km from the edge), and six linear riparian forest remnants (50–100 m wide) that were connected to continuous forest. Four nest types were also compared, representing different combinations of two factors; height (ground, shrub) and shape (open, domed). At each site, four nests of each type, containing one quail egg and two model plasticine eggs, were interspersed about 15 m apart within a 160 m transect during September–October 2001. Predators were identified from marks on the plasticine eggs. The overall depredation rate was 66.5% of 320 nests' contents damaged over a three-day period. Large rodents, especially the rat Uromys caudimaculatus, and birds, especially the spotted catbird Ailuroedus melanotis, were the main predators. Mammals comprised 56.5% and birds 31.0% of predators, with 12.5% of unknown identity. The depredation rate did not vary among site-types, or between open and domed nests, and there were no statistically significant interactions. Nest height strongly affected depredation rates by particular types of predator; depredation rates by mammals were highest at ground nests, whereas attacks by birds were most frequent at shrub nests. These effects counterbalanced so that overall there was little net effect of nest height. Mammals accounted for 78.4% of depredated ground nests and birds for at least 47.4% of shrub nests (and possibly up to 70.1%). The main predators were species characteristic of rainforest, rather than habitat generalists, open-country or edge specialists. For birds that nest in the tropical rainforest understorey of the study region, it is unlikely that edges and linear remnants presently function as ecological population sinks due to mortality associated with increased nest predation.     

Predation on artificial nests in a forest dominated landscape – the effects of nest type, patch size and edge structure

We studied the effects of forest patch size and forest edge structure on nest predation in a boreal coniferous forest landscape. The following predictions were tested. Nest predation should be higher in small than in large stands, in edges than in interior areas of forest stands, and in barren forest/clear–cut edges created by forestry than in natural forest/open marsh edges. Four types of artificial above ground nests (total of 261) were used; open cup nests with reindeer Rangifer t. tarandus hair, open cup nests with domestic hen Gallus domesticus feathers, and unlined open cup and nest–box nests. Nests were baited with one Japanese quail Coturnix coturnix japonica egg. Nest–boxes were depredated significantly less than open cup nests of all types. No edge- or stand size–related nest predation was found. The predation rate, regardless of the nest type, did not differ relative to the edge type and vegetation characteristics. However, better horizontal visibility of open cup nests due to more open vegetation structure increased predation risk in man–made edges compared to inherent edges. The results suggest that edge–related nest predation is absent or weak in forest dominated landscapes. This may be due to predator types present in the landscape and/or predators habitat use in forest dominated areas. Therefore, it might be that findings documented in other areas, such as in agricultural dominated landscapes, cannot be directly applied to managed forest landscapes.     

Predation on artificial ground nests in relation to forest fragmentation, agricultural land and habitat structure

The impacts of forest fragmentation agricultural land and habitat structure on depredation of artificial ground nests were studied in the cultivated area in central Finland and in the forest dominated area in Finnish Lapland The overall predation rate did not differ between the regions The overall predation rate was also independent of landscape characteristics forest patch size and the distance to patch edge However, nest predation was clearly affected by the agricultural land since the robbing rate in forest edges was higher near farmlands than further away This effect was caused by avian predators which proportional importance in predation was higher in the agricultural landscape than in the forest landscape In both regions, depredation correlated positively with high numbers of pine and spruce This can be mainly explained by the preference of predators over coniferous forest habitat as a living or hunting area     

Nest position and type affect predation rates of artificial avian nests in the tropical lowland forest on Mount Cameroon

Nest predation is the leading cause of reproductive failure in birds and thus it shapes their life history strategies. Intensities of nest predation appear to differ among nest locations and types in both temperate and tropical regions. However, there is limited knowledge of factors influencing susceptibility of avian nests to predation in Africa. The aim of our study was to investigate artificial nest predation rates of different ground and shrub nests located at different heights in the rainforest undergrowth. We placed artificial avian nests within a homogeneous lowland forest interior with sparse forest undergrowth in the Mount Cameroon National Park, Cameroon. We exposed three sets of nests: 50 bare-ground, 50 cup-ground and 50 cup-shrub nests, for 10 d. Predation was higher for cup-ground nests compared to cup-shrub nests, and bare-ground nests were more depredated than cup-ground nests. We concluded that the presence of a cup as well as higher nest position significantly increased probability of artificial nest survival. The results of this study suggest a potential selection pressure on nest type and placement in lowland forest birds for a poorly known tropical region.     

Temporal variation in nest predation risk along habitat edges between grassland and secondary forest in Central Europe

Habitat fragmentation alters many ecological processes, including trophic cascades. For example, increased predation pressure along habitat edges has often been observed in fragmented landscapes. Here, we studied how nest predation risk varies along the transition zone between grassland and mixed forest in Central Europe. Using artificial nests, we tested the two mechanisms that are expected to underlie higher predation rates along edges: (1) the matrix effect model that supposes predator penetration from a habitat type with higher predator density to one with lower predator density and (2) the ecotonal effect model that assumes specific predator preferences for habitat edges. Although our results do not fully support either of these scenarios, our data show high temporal instability in nest predation along forest–grassland edges. Predation was higher in habitat interiors compared to edges during the first year, whereas the opposite pattern was observed during the subsequent year. In addition, dramatic between-year differences in the species composition of nest predators were observed. Therefore, we hypothesise that the effect of edges on nest predation is difficult to predict in landscapes with high predator diversity. In addition, our data indicate that a high abundance of wild boar considerably increases the risk of predation for ground-nesting birds.     

Species traits and the response of open‐habitat species to forest edge in landscape mosaics

Human driven changes in land‐use have increased the need to understand how landscape structure affects species distribution. We studied how forest edges affected the distribution of birds in grasslands recently encroached by forest patches. We investigated how species’ biological traits influenced their response to vegetation change near forest edges. We censured birds along 300‐m line transects run into the open habitat perpendicularly to forest edges. We recorded habitat variables and landscape context along each transect and characterized edges and forest patches. We recorded 33 bird species in 153 transects for a total of 654 individuals. We analyzed species response to edges with generalized linear mixed models. Habitat preference was prevalent to explain species response to forest edges. The abundance of open‐habitat birds such as skylark Alauda arvensis decreased significantly in the vicinity of edges. This negative response extended within 150 m from the edge. The effect was disproportionately higher in open‐habitat species with high conservation concern. The abundance of species feeding or/and breeding in both forest and open habitat, such as woodlarks Lullula arborea, sharply increased near edges (positive edge response). Abundance of shrub and non‐shrub dependent species increased with distance to edge. The two species groups did no differ in abundance/distance to edge relationship. Intensity of species response to forest edges varied among transects in relation to transect vegetation characteristics. Edge length or aspect, diet and nest height had no direct effect. We discuss the possible role of variation in resources and nest predation risk to explain observed patterns.     

Artificial nest and seed predation experiments on tropical southeast Asian islands

Southeast Asia is rapidly losing native habitats and the consequences of this are poorly understood. Because habitat loss and disturbance can affect avian and seed survivorship, we conducted artificial nest and seed predation experiments on tropical southeast Asian islands. Data among islands and fragments or different forest types (e.g. primary versus exotic forest) within the islands are compared. On Singapore Island, predation among different forest types (primary, secondary and woodland) did not differ. Only at one of the sites, nest predation was higher at 75 m from the forest edge than at 25 m. In other sites, predation did not differ in relation to the distance from the forest edge. Predation among 10 small (0.8–1026 ha) Singaporean islands differed. However, none of the environmental variables (e.g. island area) could explain the predation differences. The lowest predation of both nests and seeds was recorded in the primary forest areas of a contiguous forest (25 500 ha) in central Java (Linggoasri). Small mammals were the main predators on Singapore and other surrounding islands. However, the index of potential predator abundance, overall, did not correlate with predation. While larger and more pristine forests may be better for avian and seed survivorship, pinpointing variables affecting both artificial nest and seed predation may be difficult.     

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.     

Edge effects reduce the nesting success of Acadian Flycatchers in a moderately fragmented forest

ABSTRACT.   Forest fragmentation can create negative edge effects that reduce the reproductive success of birds nesting near the forest/nonforest interface, and threaten bird populations deeper in remnant forest habitats. Negative edge effects may be more pronounced in landscapes that are moderately fragmented, particularly where agriculture is the primary land-use fragmenting forests. Information about the extent and strength of edge effects at a site can help guide conservation actions, and determine their effectiveness. We examined edge effects for birds breeding in a nearly contiguous forest fragmented by relatively narrow agricultural corridors in Illinois (USA). We measured rates of nest predation and brood parasitism for Acadian Flycatchers ( Empidonax virescens ) over a continuum of distances from the edge of an agricultural inholding. Nest predation and brood parasitism were highest near the edge and decreased with increasing distance from the edge. Given the cumulative effects of nest predation and brood parasitism on reproductive success, we determined that forest within 600 m of the inholding was sink habitat. We found, however, that deeper forest interior areas currently serve as source habitat, and that conversion of the entire 205 ha agricultural corridor to forest would add 1350 ha of source habitat for Acadian Flycatchers. Such results provide support for a local conservation strategy of forest consolidation and establish baseline measures necessary to determine the relative effectiveness of any subsequent reforestation efforts.     

Nest predation in forest birds: influence of predator type and predator's habitat quality

We used the introduction of a generalist nest predator, the red squirrel Tamiasciurus hudsonicus, and of a large herbivore, the Sitka black-tailed deer Odocoileus hemionus sitkensis, to the islands of Haida Gwaii (Queen Charlotte Islands, British Columbia, Canada) to study how predator assemblage and habitat quality and structure influenced nest predation in forest birds. We compared losses of natural nests to predators on islands with and without squirrels. We selected nine islands with or without squirrel or deer and used 506 artificial nests put on the ground or in shrubs to further analyse variation of nest predation with predator assemblage and habitat quality for the predators. For both natural and artificial nests predation risk was higher in presence of squirrels. But predation risk varied within island categories. In presence of squirrels it was highest in stands with mature conifers where it fluctuated from year to year, in response to fluctuations in squirrel abundance. Vegetation cover around the nest had little effect on nest predation by squirrels. Where squirrels were absent, nest predation concentrated near predictable food sources for corvids, the main native predators, and increased with decreasing vegetation cover, suggesting that removal of the vegetation by deer increased the risk of predation by native avian nest predators that use visual cues. Predation risk in these forests therefore varies in space and time with predator composition and with quality of the habitat from the predators' perspective. This temporal and spatial variation in predation risk should promote trade-offs in the response of birds to nest predation, rather than fine-tuned adaptations to a given predation pattern.