Power lines,roads, and avian nest survival: effects on predator identity and predation intensity

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Snake predation on North American bird nests: culprits,patterns and future directions

Predation is the leading cause of nest failure for most birds. Thus, for ornithologists interested in the causes and consequences of variation in nest success, knowing the identity and understanding the behavior of dominant nest predators is likely to be important. Video documentation of nests has shown that snakes are frequent predators. Here we reviewed 53 North American studies that used nest cameras and used these data to identify broad patterns in snake predation. Snakes accounted for 26% (range: 0–90%) of recorded predation events, with values exceeding 35% in a third of studies. Snakes were more frequent nest predators at lower latitudes and less frequent in forested habitat relative to other nest predators. Although 12 species of snakes have been identified as nest predators, ratsnakes Elaphe obsoleta, corn snakes E. guttata and fox snakes E. vulpina were the most frequent, accounting for > 70% of all recorded nest predation events by snakes and have been documented preying on nests in 30–65% of studies conducted within their geographic ranges. Endotherm‐specialist snakes (Elaphe and Pituophis genera) were more likely to depredate nests in forests and the canopy relative to other snakes, due to their affinity for edge habitat. Predation by only ratsnakes and corn snakes was predominantly nocturnal and only ratsnakes were more likely to prey on nests during the nestling stage. Snakes were not identified to species in over 30% of predation events, underlining the need for more complete reporting of results. A review of research to date suggests the best approach to investigating factors that bring snakes and nests into contact involves combining nesting studies with radio tracking of locally important snake nest predators.     

Effects of parents and Brown‐headed Cowbirds (Molothrus ater) on nest predation risk for a songbird

Nest predation limits avian fitness, so ornithologists study nest predation, but they often only document patterns of predation rates without substantively investigating underlying mechanisms. Parental behavior and predator ecology are two fundamental drivers of predation rates and patterns, but the role of parents is less certain, particularly for songbirds. Previous work reproduced microhabitat‐predation patterns experienced by Yellow Warblers (Setophaga petechia) in the Mono Lake basin at experimental nests without parents, suggesting that these patterns were driven by predator ecology rather than predator interactions with parents. In this study, we further explored effects of post‐initiation parental behavior (nest defense and attendance) on predation risk by comparing natural versus experimental patterns related to territory density, seasonal timing of nest initiation, and nest age. Rates of parasitism by Brown‐headed Cowbirds (Molothrus ater) were high in this system (49% nests parasitized), so we also examined parasitism‐predation relationships. Natural nest predation rates (NPR) correlated negatively with breeding territory density and nonlinearly (U‐shaped relationship) with nest‐initiation timing, but experimental nests recorded no such patterns. After adjusting natural‐nest data to control for these differences from experimental nests other than the presence of parents (e.g., defining nest failure similarly and excluding nestling‐period data), we obtained similar results. Thus, parents were necessary to produce observed patterns. Lower natural NPR compared with experimental NPR suggested that parents reduced predation rates via nest defense, so this parental behavior or its consequences were likely correlated with density or seasonal timing. In contrast, daily predation rates decreased with nest age for both nest types, indicating this pattern did not involve parents. Parasitized nests suffered higher rates of partial predation but lower rates of complete predation, suggesting direct predation by cowbirds. Explicit behavioral research on parents, predators (including cowbirds), and their interactions would further illuminate mechanisms underlying the density, seasonal, and nest age patterns we observed.     

Effects of Mesopredators on Nest Survival of Shrub-Nesting Songbirds

ABSTRACT Although nest predation is often the single largest source of mortality in avian populations, manipulative studies to determine predator impacts on nest survival are rare, particularly studies that examine impacts of mid-size mammalian predators (hereafter, mesopredators) on nest survival of shrub-nesting birds. We quantified nest survival and identified nest predators of shrub-nesting songbirds within 4 large (approx. 40-ha) exclosures and 4 control sites within a longleaf pine (Pinus palustris) ecosystem. During 2003–2006, we located and monitored 535 shrub nests (222 with videography) for 4,804 nest-days to quantify daily nest survival and document predation events. We found no support for a treatment effect, suggesting mesopredators had little impact on daily nest survival (0.9303 in controls and 0.9260 in exclosures) of shrub-nesting songbirds. For the 5 most commonly monitored species, daily nest survival within species was constant. Our analysis suggested that shrub nests were most vulnerable during the nestling stage and presence of cameras on nests increased survival with the increase in survival being more pronounced during the incubation stage. We filmed 107 nest predation events, identifying predators at 88 nests. Of these 88 nests, snakes caused 33%, red imported fire ants (hereafter fire ants, Solenopsis invicta) 28%, raptors 17%, corvids 8%, mesopredators 6%, and small mammals 8% of nest predations. Cause-specific nest predation in controls and exclosures did not differ from expectation, providing evidence that compensatory predation did not occur. Nest predators differed from expectation with regard to nest stage; fire ants and raptors only depredated nests during the nestling stage. Presence of cameras had no effect on nest abandonment. Fire ants were the most prevalent nest predator, and nest predation by fire ants was only observed on nestlings, potentially reducing likelihood of renesting. Magnitude and timing of fire ant predation suggests that fire ants may be the most influential nest predator of shrub-nesting birds within the longleaf pine ecosystem. Our data suggest that controlling mesopredators will have no effect on nest success of shrub-nesting birds within longleaf pine forests.     

Predators at bird nests in a northern hardwood forest in New Hampshire

ABSTRACT.   Nest predation is the primary cause of nest failure in most passerine birds, and increases in nest predation associated with anthropogenic habitat disturbance are invoked as explanations for population declines of some bird species. In most cases, however, the identity of the nest predators is not known with certainty. We monitored active bird nests with infrared time-lapse video cameras to determine which nest predators were responsible for depredating bird nests in northern New Hampshire. We monitored 64 nests of 11 bird species during three breeding seasons, and identified seven species of predators during 14 predation events. In addition, we recorded two instances of birds defending nests from predators and, in both cases, these nests were ultimately lost to predation. These results contrast with other studies in terms of the relatively high proportion of nests depredated by raptors and mice, as well as the absence of any predation by snakes. The diverse suite of predators in this and other studies is likely to confound our understanding of patterns of nest predation relative to fragmentation and habitat structure.     

Nest predators of Lance-tailed Manakins on Isla Boca Brava, Panamá

ABSTRACT.   Nest predation is often the primary cause of nest failure for passerines. Despite this, little is known about predation rates and the nest predators of birds in the tropics. I used video cameras to monitor seven Lance-tailed Manakin ( Chiroxiphia lanceolata ) nests on Isla Boca Brava, Panamá. One nest fledged young and six nests failed due to predation. I recorded five predation events involving four avian predators and one mammalian predator. Crested Oropendolas ( Psarocolius decumanus ) predated two nests and a Roadside Hawk ( Buteo magnirostris ) and a Black-chested Jay ( Cyanocorax affinis ) each predated one. The mammalian predator was a common opossum ( Didelphis marsupialis ). All avian predation was diurnal; the mammalian predation was nocturnal. My results suggest that tropical birds are subject to a diverse suite of nest predators, and that avian predators may be an important cause of nest failure at my study site.     

Do seasonal patterns of rat snake (Pantherophis obsoletus) and black racer (Coluber constrictor) activity predict avian nest predation?

Avian nest success often varies seasonally and because predation is the primary cause of nest failure, seasonal variation in predator activity has been hypothesized to explain seasonal variation in nest success. Despite the fact that nest predator communities are often diverse, recent evidence from studies of snakes that are nest predators has lent some support to the link between snake activity and nest predation. However, the strength of the relationship has varied among studies. Explaining this variation is difficult, because none of these studies directly identified nest predators, the link between predator activity and nest survival was inferred. To address this knowledge gap, we examined seasonal variation in daily survival rates of 463 bird nests (of 17 bird species) and used cameras to document predator identity at 137 nests. We simultaneously quantified seasonal activity patterns of two local snake species (N = 30 individuals) using manual (2136 snake locations) and automated (89,165 movements detected) radiotelemetry. Rat snakes (Pantherophis obsoletus), the dominant snake predator at the site (~28% of observed nest predations), were most active in late May and early June, a pattern reported elsewhere for this species. When analyzing all monitored nests, we found no link between nest predation and seasonal activity of rat snakes. When analyzing only nests with known predator identities (filmed nests), however, we found that rat snakes were more likely to prey on nests during periods when they were moving the greatest distances. Similarly, analyses of all monitored nests indicated that nest survival was not linked to racer activity patterns, but racer‐specific predation (N = 17 nests) of filmed nests was higher when racers were moving the greatest distances. Our results suggest that the activity of predators may be associated with higher predation rates by those predators, but that those effects can be difficult to detect when nest predator communities are diverse and predator identities are not known. Additionally, our results suggest that hand‐tracking of snakes provides a reliable indicator of predator activity that may be more indicative of foraging behavior than movement frequency provided by automated telemetry systems.     

Meta‐analyses of nest predation in temperate Australian forests and woodlands

Nest predation is the leading cause of nesting failure. Thus it is a crucial area of research needed to inform conservation management and to understand the life history of birds. I surveyed the literature to review the identity of nest predators and the factors affecting nest predation, in Australia using 177 studies. Overall, 94 nest predators were identified when incorporating artificial nests, 69 without. Using only natural nests, the Pied Currawong Strepera graculina was the most frequently reported nest predator. Five nest predators, including Pied Currawong, depredated 40% of the prey measured by the number of prey species taken. Yet, 60% of predation was carried out by the other 64 species, which included by the order of importance birds, mammals, reptiles, frogs and ants. Predation at cup and dome nests was more frequently reported than at burrow, ground and hollow nests. Only 28% of predators were observed at both artificial and natural nests suggesting artificial nests have limited, but not negligible, ability as tools for identifying predators. There was a highly significant and positive correlation between predator and prey masses. The predator prey mass ratio was calculated with a mean 0.25 and a median 0.22, a result closely matching with the proportional size of prey taken by raptors. The finding that predator size is proportional to prey opens a pathway for more life history and conservation research.     

Does cowbird parasitism increase predation risk to American redstart nests?

Brood parasitism and nest predation are major causes of reproductive failure for many bird species nesting in fragmented landscapes. While brood parasites and predators may act independently, they could also interact if brood parasites increase the likelihood that predators detect nests. In this study, we examined the interaction between cowbird parasitism and nest predation in a 10 year study on 466 American redstart Setophaga ruticilla nests in central Alberta, Canada. We used advanced nest survival models to examine the support for three mechanisms that might lead to a positive correlation between brood parasitism and nest predation: 1) the presence of a cowbird nestling might increase the detection of the nest by predators, 2) nests with lower cover are more likely to be detected by both cowbirds and predators, and 3) cowbirds and predators may co-occur in landscapes of similar structure. Twelve percent of nests were parasitized and those nests had a 16–19% higher rate of failure due to predators compared to unparasitized nests. Daily nest predation rates increased during the nestling stage for both groups, but more strongly for parasitized nests. Loud begging by the cowbird nestling and/or higher parental feeding rates for the cowbird may have increased nest detectability to predators. Brood parasitism and nest predation were also positively related to forest cover, indicating landscape level effects were influential. Most nest predators were forest species and we suspect cowbirds responded positively to forest cover because of the increased abundance of songbird hosts. Nest-site features had less of an impact on nest predation or brood parasitism, although nests with higher overhead cover were less susceptible to predators. Our study shows how multiple mechanisms, particularly the behavioral effects of the brood parasite nestling and landscape structure, can lead to a positive relationship between nest predation and brood parasitism.     

Impact of brood parasitism and predation on nest survival of the fan-tailed gerygone in New Caledonia

Predation and brood parasitism are common reasons for nesting failure in passerine species and the additive impact by invasive species is a major conservation concern, particularly on tropical islands. Recognising the relative contribution of the different components of nesting failure rates is important to understand co-evolutionary interactions within brood parasite–host systems. In the remote archipelago of New Caledonia, the fan-tailed gerygone Gerygone flavolateralis is the exclusive host of the brood-parasitic shining bronze-cuckoo Chalcites lucidus. Additionally, invasive rodents also possibly have an impact on breeding success. To estimate the impact of potential nest predators, we 1) video monitored nests to identify predators, 2) estimated the probability of predation based on nest visibility and predator abundance and 3) tested the possibility that the location of experimental nests and lack of odour cues decrease the predation by rodents. In addition, we estimated nest survival rates using data collected in different habitats over the course of eight breeding seasons. Nesting success of fan-tailed gerygones was relatively low and predation was the main cause of nesting failure. We recorded mainly predation by native birds, including the shining bronze-cuckoo, whereas predation by rats was rare. In open habitats predation by cuckoos was much lower than predation by other avian predators. Neither predator activity around nests nor nest visibility influenced the probability of predation. Experimental nests in more accessible locations and containing an odorous bait were more exposed to rodent predation. Apparently, the fan-tailed gerygone has either never been specifically vulnerable to predation by rats or has developed anti-predator adaptations.     

Increased nest defence of upland‐nesting ducks in response to experimentally reduced risk of nest predation

Parent birds should take greater risks defending nests that have a higher probability of success. Given high rates of mammalian nest predation, therefore, parents should risk more for nests in areas with a lower risk of mammalian predation. We tested this hypothesis using nest defence data from over 1300 nests of six species of dabbling ducks studied in an area where predation risk had been reduced through removal of mammalian predators. When predator removal reduced nest predation, the ducks increased risk taking as predicted. Also as predicted, risk taking varied inversely with body size, an index of annual survival, among species. For ducks to vary nest defence in response to variation in predation risk they must be able to assess the risk of nest predation. Because ducks modified nest defence in the breeding season immediately following predator removal, ducks may be able to assess predator abundance indirectly (e.g. by UV reflection from urine) rather than by seeing or interacting directly with the predators.     

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     

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.     

Nest height, nest concealment, and predator type predict nest predation in superb fairy-wrens (Malurus?cyaneus)

Three factors and their interaction effects are increasingly recognized as important determinants of nest predation: nest concealment, nest height, and predator type. The risk of nest predation is predicted to vary across these variables because of nest detectability and accessibility. In general, however, few studies examine how these three variables interact in relation to nest predation, focusing instead on either nest concealment or nest height (whereby predator identity is usually not known). In this study, we examine the role of nest concealment and nest height for nest survival using both artificial and natural nests in the superb fairy-wren (Malurus cyaneus). We indirectly identified potential predators through marks left on artificial eggs and footprints left on tracking tunnels. Predation level at artificial nests was lower than at natural nests, and this could be due to a failure of some nest predators to locate cryptic nests in the absence of cues provided by parental activity. Our results supported the prediction that exposed and concealed nests have different levels of nest predation, which can be explained by variation in predator type. Visual predators were only detected at exposed nests, and survival from visual predators was lower for high nests that were also exposed. However, olfactory predators were detected irrespective of nest height or nest concealment. Because rodents use olfaction to locate nests, this could explain the lack of association between nest concealment and predation outcome at low nests. In addition, rodent footmarks near nests were significantly associated with rodent tooth marks on eggs.     

Repeatability of nest predation in passerines depends on predator species and time scale

It has been proposed that some specific locations of bird's nests have higher intrinsic chances of being depredated than other locations. This predicts that fates of consecutive nesting attempts at the same site should be repeatable. We used 20 pairs of old thrush nests to simulate repeated nesting attempts at the same sites, both within and between breeding seasons (n=40  sites×2  trials×2  years=160). Each nest was monitored by a camera to record multiple predation events and to identify predators. Predation by all predator species was repeatable during a 15-day trial. Predation by principal predators (jay Garrulus glandarius , marten Martes martes / foina ) and total predation (all species combined) was not correlated within pairs of simultaneously exposed nests or within samples of nests from particular study plot, and not repeatable for individual nests between-trials or between-years. These findings suggest short-term effect of predator memory causing revisitation of previously depredated nests during a current nesting trial (all predators); do not support an effect of nest site features on multiple nest discoveries and/or an effect of nest location on repeated random encounters with the same nest (principal predators). Long-term repeatability and correlation within pairs of simultaneously exposed nests was detectable only in occasional predators (great spotted woodpecker Dendrocopos major , possibly also squirrel Sciurus vulgaris ), which suggests effect of nest location combined with site fidelity and individual foraging specialization of these predators. We conclude that repeatability of nest predation depends on the time scale considered and the local predator community. We caution against spurious findings of repeatable nest predation resulting simply from statistical properties of correlation in binary data (nest fates).     

Black-capped vireo nest predator assemblage and predictors for nest predation

Nest predation is a major limiting factor for songbird productivity, including the federally endangered black-capped vireo (Vireo atricapilla). However, nest predator information is limited across the range of the black-capped vireo in central and southwest Texas. We monitored nests in 3 counties within the breeding range of black-capped vireos in Texas in 2008 and 2009 and used continuous recording digital video cameras to record predation events. We video-monitored 115 nests and documented 39 predation events by at least 9 predator species. Overall, we observed avian species (51%, n = 39), specifically brown-headed cowbirds (Molothrus ater; n = 12), and snakes (26%, n = 39) as the most frequent nest predators. The estimated daily nest survival rate during the laying and incubation stage was 0.985 (95% CI = 0.967–0.993) and 0.944 (95% CI = 0.921–0.961) during the nestling stage. In addition, we analyzed models of predator-specific nest predation using multinomial logistic regression. Effect of nest height on predation rate was significant for snakes; nest stage was significant for nests depredated by avian predators. By identifying and increasing our knowledge of nest predators and vegetation characteristics associated with greater risk of predation in multiple locations within the black-capped vireo's range, we can effectively manage habitat to benefit recovery efforts of the species. © 2012 The Wildlife Society.     

Breeding Phenology of Birds: Mechanisms Underlying Seasonal Declines in the Risk of Nest Predation

Seasonal declines in avian clutch size are well documented, but seasonal variation in other reproductive parameters has received less attention. For example, the probability of complete brood mortality typically explains much of the variation in reproductive success and often varies seasonally, but we know little about the underlying cause of that variation. This oversight is surprising given that nest predation influences many other life-history traits and varies throughout the breeding season in many songbirds. To determine the underlying causes of observed seasonal decreases in risk of nest predation, we modeled nest predation of Dusky Flycatchers (Empidonax oberholseri) in northern California as a function of foliage phenology, energetic demand, developmental stage, conspecific nest density, food availability for nest predators, and nest predator abundance. Seasonal variation in the risk of nest predation was not associated with seasonal changes in energetic demand, conspecific nest density, or predator abundance. Instead, seasonal variation in the risk of nest predation was associated with foliage density (early, but not late, in the breeding season) and seasonal changes in food available to nest predators. Supplemental food provided to nest predators resulted in a numerical response by nest predators, increasing the risk of nest predation at nests that were near supplemental feeders. Our results suggest that seasonal changes in foliage density and factors associated with changes in food availability for nest predators are important drivers of temporal patterns in risk of avian nest predation.     

Nocturnal nest predation: a potential obstacle to recovery of a Florida Scrub-Jay population

ABSTRACT.   Population declines among birds are often linked to habitat change and associated increases in nest predation rates. In species of conservation concern identifying nest predators is an important first step in developing management strategies to mitigate low nesting success caused by depredation. Because predator composition varies geographically and with landscape factors habitat restoration may need to be tailored to reduce locally important predators. We used miniature video cameras to identify nest predators in a population of Florida Scrub-Jays ( Aphelocoma coerulescens ) significant to conservation. At 22 nests we observed 25 predation events; 22 (88%) of these events were nocturnal. Yellow rat snakes ( Elaphe obsoleta ) had the highest daily predation rate and accounted for 76% of egg and nestling losses. Florida Scrub-Jays are vulnerable to nocturnal nest predation because their vigilance behavior is ineffective against nocturnal predators, breeders cannot defend against nocturnal predators, and brooding females are at risk of being killed by nocturnal predators. If current habitat restoration efforts do not reduce numbers of yellow rat snakes and improve scrub-jay nesting success, management actions to reduce populations of nocturnal snakes may need to be considered.     

Effects of nest-site characteristics and parental activity on cowbird parasitism and nest predation in Brown-and-yellow Marshbirds

ABSTRACT Nest‐site selection and nest defense are strategies for reducing the costs of brood parasitism and nest predation, two selective forces that can influence avian nesting success and fitness. During 2001–2002, we analyzed the effect of nest‐site characteristics, nesting pattern, and parental activity on nest predation and brood parasitism by cowbirds (Molothrus spp.) in a population of Brown‐and‐yellow Marshbirds (Pseudoleistes virescens) in the Buenos Aires province, Argentina. We examined the possible effects of nest detectability, nest accessibility, and nest defense on rates of parasitism and nest predation. We also compared rates of parasitism and nest predation and nest survival time of marshbird nests during the egg stage (active nests) with those of the same nests artificially baited with passerine eggs after young fledged or nests failed (experimental nests). Most nests (45 of 48, or 94%) found during the building or laying stages were parasitized, and 79% suffered at least one egg‐predation event. Cowbirds were responsible for most egg predation, with 82 of 107 (77%) egg‐predation events corresponding to eggs punctured by cowbirds. Nests built in thistles had higher rates of parasitism and egg predation than nests in other plant, probably because cowbirds were most active in the area where thistles were almost the only available nesting substrate. Parasitism rates also tended to increase as the distance to conspecific nests increased, possibly due to cooperative mobbing and parental defense by marshbirds. The proportion of nests discovered by cowbirds was higher for active (95%) than for experimental (29%) nests, suggesting that cowbirds used host parental activity to locate nests. Despite active nest defense, parental activity did not affect either predation rates or nest‐survival time. Thus, although nest defense by Brown‐and‐yellow Marshbirds appears to be based on cooperative group defense, such behavior did not reduce the impact of brood parasites and predators.     

Contrast in edge vegetation structure modifies the predation risk of natural ground nests in an agricultural landscape

Nest predation risk generally increases nearer forest-field edges in agricultural landscapes. However, few studies test whether differences in edge contrast (i.e. hard versus soft edges based on vegetation structure and height) affect edge-related predation patterns and if such patterns are related to changes in nest conspicuousness between incubation and nestling feeding. Using data on 923 nesting attempts we analyse factors influencing nest predation risk at different edge types in an agricultural landscape of a ground-cavity breeding bird species, the Northern Wheatear (Oenanthe oenanthe). As for many other bird species, nest predation is a major determinant of reproductive success in this migratory passerine. Nest predation risk was higher closer to woodland and crop field edges, but only when these were hard edges in terms of ground vegetation structure (clear contrast between tall vs short ground vegetation). No such edge effect was observed at soft edges where adjacent habitats had tall ground vegetation (crop, ungrazed grassland). This edge effect on nest predation risk was evident during the incubation stage but not the nestling feeding stage. Since wheatear nests are depredated by ground-living animals our results demonstrate: (i) that edge effects depend on edge contrast, (ii) that edge-related nest predation patterns vary across the breeding period probably resulting from changes in parental activity at the nest between the incubation and nestling feeding stage. Edge effects should be put in the context of the nest predator community as illustrated by the elevated nest predation risk at hard but not soft habitat edges when an edge is defined in terms of ground vegetation. These results thus can potentially explain previously observed variations in edge-related nest predation risk.