Testing ecological and behavioral correlates of nest predation

Variation in nest predation rates among bird species are assumed to reflect differences in risk that are specific to particular nest sites. Theoretical and empirical studies suggest that parental care behaviors can evolve in response to nest predation risk and thereby differ among ecological conditions that vary in inherent risk. However, parental care also can influence predation risk. Separating the effects of nest predation risk inherent to a nest site from the risk imposed by parental strategies is needed to understand the evolution of parental care. Here we identify correlations between risks inherent to nest sites, and risk associated with parental care behaviors, and use an artificial nest experiment to assess site-specific differences in nest predation risk across nesting guilds and between habitats that differed in nest predator abundance. We found a strong correlation between parental care behaviors and inherent differences in nest predation risk, but despite the absence of parental care at artificial nests, patterns of nest predation risk were similar for real and artificial nests both across nesting guilds and between predator treatments. Thus, we show for the first time that inherent risk of nest predation varies with nesting guild and predator abundance independent of parental care.     

Do alien predators pose a particular risk to duck nests in Northern Europe? Results from an artificial nest experiment

Several alien predator species have spread widely in Europe during the last five decades and pose a potential enhanced risk to native nesting ducks and their eggs. Because predation is an important factor limiting Northern Hemisphere duck nest survival, we ask the question, do alien species increase the nest loss risk to ground nesting ducks? We created 418 artificial duck nests in low densities around inland waters in Finland and Denmark during 2017–2019 and monitored them for seven days after construction using wildlife cameras to record whether alien species visit and prey on the nests more often than native species. We sampled various duck breeding habitats from eutrophic agricultural lakes and wetlands to oligotrophic lakes and urban environments. The results differed between habitats and the two countries, which likely reflect the local population densities of the predator species. The raccoon dog (Nyctereutes procyonoides), an alien species, was the most common mammalian nest visitor in all habitats and its occurrence reduced nest survival. Only in wetland habitats was the native red fox (Vulpes vulpes) an equally common nest visitor, where another alien species, the American mink (Neovison vison), also occurred among nest visitors. Although cautious about concluding too much from visitations to artificial nests, these results imply that duck breeding habitats in Northern Europe already support abundant and effective alien nest predators, whose relative frequency of visitation to artificial nests suggest that they potentially add to the nest predation risk to ducks over native predators.

     

Landscape and habitat affect frequency of artificial duck nest predation by native species,but not by an alien predator

Annual Finnish breeding duck surveys over the last 30 years show declining abundance among several species and greater declines on eutrophic waters than oligotrophic lakes. It has been suggested that habitat-related differences in the rate of increase in predation pressure is a potential explanation for contrasting duck population trajectories between habitats. We assessed potential duck nest predation risk and predator presence in various duck breeding habitats in Finland and Denmark by monitoring 333 artificial duck nests with wildlife cameras during 2017–2019. Predation rates differed between landscapes and habitats: nest predation rate and predator diversity were lowest in forested and highest in agricultural landscapes. Forest nests further from water bodies survived better than nests around shorelines of permanent lakes. Of the 16 different predator species detected, the most common were Eurasian magpie (Picapica), hooded crow (Corvus corone) and raccoon dog (Nyctereutes procyonoides). While predation by specific native predator species was typically associated with particular habitats and landscapes, the alien raccoon dog appeared to be a true habitat generalist, ubiquitous and common across all habitats and landscapes. Based on these results, the higher duck nest predation pressure along shorelines, especially in agricultural landscape lakes, due to increased diversity and abundance within the predator community, may contribute to the declining population trends of ducks.     

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     

Parent birds assess nest predation risk: influence of cavity condition and avian nest predator activity

Skutch hypothesized that nest predators visually assess parental activities to locate a prey nest, whereas parents modify fitness‐related traits to reduce the probability of nest predation. We examined how cavity condition and parental activity interact with avian nest predators to shape the nest success of two coexisting parid species, marsh tits Poecile palustris and oriental tits Parus minor, breeding in nest‐boxes during the incubation period. Nest‐boxes were manipulated to create a prolonged risk of nest predation (entrance diameter 2.6 cm control vs 5.5 cm treatment) soon after clutch completion. To measure changes in parental behavior, we also simultaneously simulated a pulsed risk of nest predation, using sound playbacks of a coexisting control bird and an avian nest predator. We found that the parent tits merely responded the pulsed risk, presumably due to an environment with high avian nest predator encounters, compared to the prolonged risk. Instead, both species spent more time on vigilance at the nest, only under prolonged risk conditions. The activity of corvids near the nest‐box was higher in the marsh tit than that in oriental tits. This activity was also higher in the treatment nest box than that in the control nest‐box. Nest predation during the incubation period was higher in marsh tits than in oriental tits, presumably due to higher and more plastic vigilance in oriental tits, compared to marsh tits. Our results highlight that the differences in cavity condition and parental activities at the nests of two coexisting non‐excavators may contribute to differential nest predation by attracting avian nest predators.     

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.     

Mothers vigilantly guard nests after partial brood loss: a cue of nest predation risk in a paper wasp

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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).     

Adaptive nest clustering and density‐dependent nest survival in dabbling ducks

Density‐dependent population regulation is observed in many taxa, and understanding the mechanisms that generate density dependence is especially important for the conservation of heavily‐managed species. In one such system, North American waterfowl, density dependence is often observed at continental scales, and nest predation has long been implicated as a key factor driving this pattern. However, despite extensive research on this topic, it remains unclear if and how nest density influences predation rates. Part of this confusion may have arisen because previous studies have studied density‐dependent predation at relatively large spatial and temporal scales. Because the spatial distribution of nests changes throughout the season, which potentially influences predator behavior, nest survival may vary through time at relatively small spatial scales. As such, density‐dependent nest predation might be more detectable at a spatially‐ and temporally‐refined scale and this may provide new insights into nest site selection and predator foraging behavior. Here, we used three years of data on nest survival of two species of waterfowl, mallards and gadwall, to more fully explore the relationship between local nest clustering and nest survival. Throughout the season, we found that the distribution of nests was consistently clustered at small spatial scales (?50–400 m), especially for mallard nests, and that this pattern was robust to yearly variation in nest density and the intensity of predation. We demonstrated further that local nest clustering had positive fitness consequences – nests with closer nearest neighbors were more likely to be successful, a result that is counter to the general assumption that nest predation rates increase with nest density.     

A predator's perspective of nest predation: predation by red squirrels is learned,not incidental

Nest predation has been used to explain aspects of avian ecology ranging from nest site selection to population declines. Many arguments rely on specific assumptions regarding how predators find nests, yet these predatory mechanisms remain largely untested. Here we combine artificial nest experiments with behavioural observations of individual red squirrels Tamiasciurus hudsonicus to differentiate between two common hypotheses: predation is incidental versus learned. Specifically, we tested: 1) whether nest survival could be explained solely by a squirrel's activity patterns or habitat use, as predicted if predation was incidental; or 2) if predation increased as a squirrel gained experience preying on a nest, as predicted if predation was learned. We also monitored squirrel activity after predation to test for evidence of two search mechanisms: area‐restricted searching and use of microhabitat search images. Contrary to incidental predation and in support of learning, squirrels did not find nests faster in areas with high use (e.g. forest edges). Instead, survival of artificial nests was strongly related to a squirrel's prior experience preying on artificial nests. Experience reduced nest survival times by over half and increased predation rates by 150–200%. Squirrels returned to and doubled their activity at the site of a previously preyed on nest. However, neither area‐restricted searching nor microhabitat search images can explain how squirrels located artificial nests more readily with experience. Instead, squirrels likely used cues associated with the nests or eggs themselves. Learning implies that squirrels could be increasingly effective predators as the density or profitability of nests increases. Our results add support to the view that nest predation is complex and broadly influenced (e.g. by predator experience, motivation), and is unlikely to be predicted consistently by simple relationships with predator activity, abundance or habitat.     

Effects of northern pike on patterns of nest use and resproductive behavior of male fathead minnows in a boreal lake

We conducted a two-part study to assets predator avoidance byreproductive male fathead minnows (Pimephales promelos) subjectedto predation threat from northern pike (Esox lucius). First,we determined if patterns of nest use by egg-guarding male minnowsin a boreal lake were related to pike densities. We samplednorthern pike and identified four areas of "high pike-density"and three areas of "low pike-density." We censused natural nestsand placed nest boards in these areas. We found eggs on naturalnests more frequently in areas with low densities of pike thanin areas with high densities of pike. However, we could notfully explain the distribution of nests by predation risk. Second,we evaluated the behavioral response of egg-guarding males toa control stimulus (a piece of wood) or a live pike in a wirecage. We used time to return to the nest after a stimulus asa measure of risk taking. Males took different amounts of riskbased on predation threat; males in the predator treatment tooklonger to return to their nests than control males. Risk takingwas not related to the number or age of the eggs but to distanceto nearest egg-guarding neighbor; males with close neighborsreturned sooner than more isolated males. Males in the predatortreatment had lower total activity and egg rubbing than controlmales after they returned to their nests. We conclude that malefathead minnows altered their reproductive behavior in waysthat reduced predation risk, but the cost of predator avoidancemight include egg predation, lost mating opportunities, or usurpationof nests     

The influence of parental behavior on vulnerability to nest predation in tropical thrushes of an Andean cloud forest

The Skutch hypothesis predicts that parental activity around the nest may attract the attention of predators and thus, in the tropics where predation pressure may be high, selection favors reduced parental activity. This hypothesis has been questioned by studies demonstrating that parents can decrease the risk of nest predation through nest defense. The link between parental activity and predation risk may be further confounded by nest site characteristics. We examined the effects of parental behavior and nest site on clutch survival in two sympatric tropical thrushes (Myadestes ralloides and Turdus leucops). We compared survival rates of clutches in three treatments: 1) natural nests at the incubation stage, 2) unattended nests (un‐manipulated nests of the same species, with clutches unattended by parents), and 3) exposed clutches (eggs exposed in unconcealed positions, unprotected by the nest). Parental activity had a positive effect on clutch survival, which was revealed by significantly higher survival rate of clutches in attended nests compared to unattended nests. The effect of nest site was less clear: clutches in unattended natural nests survived better than clutches in exposed sites selected by humans, but results were insignificant. We propose that parent birds can exclude a group of opportunistic predators, that are able to destroy unattended clutches. Nest site characteristics may be less important in determining clutch survival in the tropics, where predator guilds are more diverse, making completely safe sites difficult to find. Our results challenge Skutch's hypothesis and point to the need for more data from tropical latitudes.     

A retrospective analysis of sea turtle nest depredation patterns

Nest predation can significantly reduce hatchling recruitment in sea turtle populations. We examined 20 yr of data from Canaveral National Seashore, Florida, which has pristine and altered beaches. We used chi-squared test to determine if secondary predation events were related to the nests' primary predation events, and proportional hazard regression analysis to determine the relative risk of individual nest predation. To determine if human beach use and nest predation risk were spatially or temporally linked, we ranked human beach use and examined predation frequency across all screened and marked nests. We found that once a nest has experienced predation it has an increased likelihood of experiencing a subsequent predation event when compared to other nests on the beach. Primary and multiple predation events occurred with greater frequency in limited beach use areas and with lowest frequency in moderate use areas. Predation risk decreased by an average of 29.5% from 2000 to 2008, relative to the initial year of study. Nests deposited mid-season were 9.8% more likely to be predated than early or late season nests. We conclude that where anti-nest predator management efforts are required but restricted (e.g., by funds, available personnel, or multiple management goals) resources should be concentrated to protect mid-season nests and those in limited access areas, if enhancing hatchling numbers is a management goal. © 2012 The Wildlife Society.     

Appearance and vulnerability of artificial duck nests to avian predators

Use of artificial nests in studies of nest success in birds has become increasingly popular, but this practice has been criticised because the appearance and success of artificial nests may differ from natural nests. Typically, artificial nests are presented with few visible eggs throughout the entire exposure period while natural nests during their life have different appearances, possibly producing stage-dependent predation rates. In ducks, nests may appear with openly visible eggs, eggs covered with nest material and covered by an incubating female. To test how nest appearance may affect nest survival, I simulated such duck nest appearances, and used direct observations to record responses of marsh harriers Circus aeruginosus , the dominant duck nest predator at the Lake Engure, Latvia, during the breeding seasons 2000–2002. I recorded 274 harrier flights over all types of artificial nests, 74 nest discoveries and 54 predation events. Logistic regression analyses showed that nest appearance was a significant predictor of the fate of the nest: nests with openly visible eggs were discovered more often than nests covered with nest material. Nests with dummy females had discovery rates similar to the average of all artificial nest types suggesting that they best reflect the predation risk of artificial nests and could be used in future studies. After discovering nests covered with dummy females, harriers never attacked but tried to scare them off to get access to the eggs. In 52% of cases of nest discovery, harriers gave up before touching the dummy. This shows that duck passive nest defence may prevent clutch predation.     

Incidental nest predation in freshwater turtles: inter- and intraspecific differences in vulnerability are explained by relative crypsis

There has long been interest in the influence of predators on prey populations, although most predator–prey studies have focused on prey species that are targets of directed predator searching. Conversely, few have addressed depredation that occurs after incidental encounters with predators. We tested two predictions stemming from the hypothesis that nest predation on two sympatric freshwater turtle species whose nests are differentially prone to opportunistic detection—painted turtles (Chrysemys picta) and snapping turtles (Chelydra serpentina)—is incidental: (1) predation rates should be density independent, and (2) individual predators should not alter their foraging behavior after encountering nests. After monitoring nest survival and predator behavior following nest depredation over 2 years, we confirmed that predation by raccoons (Procyon lotor), the primary nest predators in our study area, matched both predictions. Furthermore, cryptic C. picta nests were victimized with lower frequency than more detectable C. serpentina nests, and nests of both species were more vulnerable in human-modified areas where opportunistic nest discovery is facilitated. Despite apparently being incidental, predation on nests of both species was intensive (57% for painted turtles, 84% for snapping turtles), and most depredations occurred within 1 day of nest establishment. By implication, predation need not be directed to affect prey demography, and factors influencing prey crypsis are drivers of the impact of incidental predation on prey. Our results also imply that efforts to conserve imperiled turtle populations in human-modified landscapes should include restoration of undisturbed conditions that are less likely to expose nests to incidental predators.     

Predation of Lapwing Vanellus vanellus nests on lowland wet grassland in England and Wales: effects of nest density, habitat and predator abundance

There is concern that predation of Lapwing Vanellus vanellus nests may create additional pressure on declining populations of this species in Europe. At seven sites in England and Wales, daily nest predation rates on 1,390 nests were related to variables using Generalised Linear Mixed Models. The strongest predictor was Lapwing nest density (number of nests within 100 m): predation rates declined as nest density increased. Since nocturnal species, probably mammals, have been identified as the major predators of Lapwing nests at these sites, these results suggest that Lapwings are able to deter mammalian predators or may settle to nest at high densities in areas with low predation pressure. At the site level, there was no relationship between Lapwing nesting density and fox density, and a positive relationship with Carrion Crow Corvus corone nesting density. There was a weaker effect of distance to field boundary: nests closer to boundaries were more likely to be predated. Weak interactive effects between crow density and both nest visibility and distance to vantage point were identified in models using a reduced subset of nests. These were counter-intuitive, did not persist in the larger data set, and do not have obvious explanations. If Lapwings nesting at high density are able to deter predators, there are implications for land management. Smaller areas could be managed within potential breeding habitat to encourage Lapwings to nest in dense colonies. Selection of larger fields for such management, where nests could be located far from the field boundary should improve the value of such measures.     

Intraseasonal patterns in shorebird nest survival are related to nest age and defence behaviour

Nest survival may vary throughout the breeding season for many bird species, and the nature of this temporal variation can reveal the links between birds, their predators, and other components of the ecosystem. We used program Mark to model patterns in nest survival within the breeding season for shorebirds nesting on arctic tundra. From 2000 to 2007, we monitored 521 nests of five shorebird species and found strong evidence for variation in nest survival within a nesting season. Daily nest survival was lowest in the mid-season in 5 of 8 years, but the timing and magnitude of the lows varied. We found no evidence that this quadratic time effect was driven by seasonal changes in weather or the abundance of predators. Contrary to our prediction, the risk of predation was not greatest when the number of active shorebird nests was highest. Although nest abundance reached a maximum near the middle of the breeding season, a daily index of shorebird nest activity was not supported as a predictor of nest survival in the models. Predators’ access to other diet items, in addition to shorebird nests, may instead determine the temporal patterns of nest predation. Nest survival also displayed a positive, linear relationship with nest age; however, this effect was most pronounced among species with biparental incubation. Among biparental species, parents defended older nests with greater intensity. We did not detect a similar relationship among uniparental species, and conclude that the stronger relationship between nest age and both nest defence and nest survival for biparental species reflects that their nest defence is more effective.     

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.     

Predator recognition and differential behavioural responses of adult wood warblers <Emphasis Type="Italic">Phylloscopus sibilatrix</Emphasis>

Birds often engage in nest defence against predators to improve breeding success, but defence efficiency requires the capability to assess the threat level posed by potential predators. For species with low breeding-site tenacity, which may encounter varying occurrence and density of predators in different areas, threat recognition could be compromised due to naivety, and so predator recognition may focus on broad key features to diminish the risk of misidentification. We experimentally tested this hypothesis by recording behavioural reactions of the nomadic wood warbler Phylloscopus sibilatrix to objects reflecting various levels of threat: least weasel and Eurasian jay taxidermy mounts, an inanimate object and an empty display mount. To assess actual nest predators, we used remote cameras to record predation events at wood warbler nests. As in other studies in Western Europe, Eurasian jay was found to be the main nest predator, with occasional predation by least weasel. The reaction of adult warblers to the models was generally to remain silent and on nests during the incubation stage presumably due to the need to maintain efficient nest camouflage and concealment. During the nestling stage, behavioural responses of adult warblers, calling and suspended feeding of young, showed the strongest effects from the jay taxidermy mount, moderate to the weasel and weakest to the inanimate object and empty mount. As the reaction of wood warblers reflected the degree of genuine threat posed by the predators depicted by the models, we conclude that predator recognition may be present in this species.     

Supplemental food alters nest defence and incubation behaviour of an open‐nesting wetland songbird

Climate‐driven increases in spring temperatures are expected to result in higher prey availability earlier in the breeding season for insectivorous birds breeding in wetland habitats. Predation during the incubation phase is a major cause of nesting failure in open‐nesting altricial birds such as the Eurasian reed warbler. The nest predation rate in this species has recently been shown to be substantially reduced under conditions of experimentally elevated invertebrate prey availability. Food availability near the nest may be an important determinant of adult incubation and nest defence behaviours during the incubation period. We used two experimental studies to compare incubation behaviour and nest defence in food‐supplemented and unsupplemented adult Eurasian reed warblers during the incubation phase. In the first study we measured nest defence behavioural responses to a taxidermic mount of a native predator (stoat Mustela erminea). In the second study we used temperature loggers installed in nests to measure breaks in incubation as a measure of nest vulnerability. Food‐supplemented birds responded aggressively to the presence of a predator more quickly than those in the unsupplemented group, suggesting they are closer to their nest and can more quickly detect a predator in the vicinity. Food‐supplemented birds also had shorter breaks in incubation (both in terms of maximum and mean off‐bout durations), presumably because they were foraging for shorter periods or over shorter distances from the nest. This study therefore identifies the behavioural mechanisms by which changes in food availability may lead to changes in nest survival and thus breeding productivity, in open‐nesting insectivorous birds.