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.     

Alternative nesting behaviours following colonisation of a novel environment by a passerine bird

When birds settle in new environments, they may encounter different suites of nest-predators, resulting in strong selection to nest in novel locations. An important axis of variation is in height of nest placement, because off-ground nests are likely to be subject to different predation pressures than on-ground nests. In their native habitats, i.e. temperate montane forest, dark-eyed juncos Junco hyemalis predominantly nest on the ground. A population of juncos became established on the urban campus of the Univ. of California at San Diego in the early 1980s, and now has a relatively high frequency (∼20%) of off-ground nests. Off-ground nesting is associated with strong fitness benefits: based on returns from 579 nests, an egg laid off the ground has ∼80% higher chance of producing a recruit to the next generation than one laid on the ground. In addition, some nests were occasionally re-used by ∼10% of females and off-ground nests were more likely to be re-used. Many females that bred off-ground also built an on-ground nests and we did not detect an association in nest site positions of mothers and daughters, implying low heritability of this novel nest placement trait. By contrast, certain territories consistently had on-ground nests irrespective of the identity of the breeding female. We suggest that a limitation of suitable off-ground nest sites for a species predominantly adapted to on-ground nesting greatly slows the rate of evolution of this trait.     

Experimental enlargement of nest size does not increase risk of predation or brood parasitism in the Great Reed Warbler Acrocephalus arundinaceus

We assessed whether nest size affects the probability of nest loss using dyads of large and small (large being twice the size of small) inactive Great Reed Warbler Acrocephalus arundinaceus nests placed at similar sites in Great Reed Warbler territories. Large nests were not predated significantly more frequently than small nests. Experimentally enlarged active Great Reed Warbler nests suffered non‐significantly higher predation compared with non‐manipulated control nests. Our experiments did not support the nest‐size hypothesis and suggested that nest size does not appear to be a factor affecting the risk of nest predation in this species. The probability of brood parasitism by the Common Cuckoo Cuculus canorus was also unaffected by experimental nest enlargement, supporting the commonly accepted hypothesis that the Common Cuckoo searches for suitable host nests by host activity during nest building rather than nest size.     

Nest predation of grassland bird species increases with parental activity at the nest

Alexander Skutch predicted that nest predation will increase with activity at nests, and that predation should be greatest during the nestling stage when parents are feeding young. We tested this hypothesis using three ecologically similar grassland bird species nesting on the high altitude grasslands of Wakkerstroom, South Africa. Parental activity, measured as adult arrival and departure frequency from the nest, was greater during the nestling than incubation stage. Nest predation, however, did not increase with parental activity between these stages in all three study species. However, nest site effects could have confounded this result. We therefore conducted an experiment that controlled for parental activity (by reusing natural nests of the study species with artificial clutches) in order to test for nest site effects. Nests that had a high rate of predation when used by active parents had a correspondingly high rate of depredation when the same nests were reused with artificial clutches (i.e. after controlling for parental activity). This result supports the notion that variation in nest site quality is the primary factor affecting nest predation rate. We also tested whether high predation during incubation is related to nest site effects. Nest predation rates of experimental clutches placed in reused nests that had been originally (when active parents were present) depredated during incubation stage were significantly higher than those that were originally depredated during the nestling stage. Finally, once nest site effects were accounted for, nest predation showed a positive increase with parental activity during the nestling stage across species.     

Winter territoriality in migrant Black Redstarts Phoenicurus ochrurus in the Mediterranean area

The nest predation hypothesis predicts that clutch size should be inversely related to the risk of nest predation. In this paper we analyse data on nest predation, clutch size and fledgling production in 128 Jackdaw nests in order to test if clutches are larger in the safer sites. The predation rate was positively correlated with the minimum nest-entrance dimensions (predation is greater in nests with large entrances). Clutch size was not related to the risk of nest predation, and our data do not therefore support the nest predation hypothesis.     

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.     

Male Nest Choice in Sand Gobies, Pomatoschistus minutus

Although material resources can have a direct bearing on the fitness of both sexes, few studies have actually examined resource-based preferences from a male choice perspective. In sand gobies, Pomatoschistus minutus , the size of a male's nest influences his attractiveness to females and also dictates the number of eggs he can receive. Thus, one might expect males to prefer larger nests. However, an earlier study of marine sand gobies from a population with a surplus of nest sites and high nest predation found that males exhibited size-assortative nest preferences. Here, we investigated male nest preferences from a brackish population characterised by a chronic nest shortage but lower predation risk. A survey of naturally settled nests in the field (shells and rocks) showed a pattern of size-assortative nest occupancy consistent with the previously studied population, with larger males occupying larger (i.e. rock) nests. However, when offered a choice of potential nests in the absence of male competition, we found that all male gobies in our population, irrespective of their own body size, actually preferred larger nests. Moreover, a predilection towards large nests superseded any preferences based on nest colour. Our results not only indicate the existence of male preferences for material resources but, considered in the light of previous work, also suggest that such preferences may vary among populations and, importantly, may not necessarily be realised in a competitive setting.     

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).     

Exclosure fences around nests of imperiled Florida Grasshopper Sparrows reduce rates of predation by mammals

Increasing nest survival by excluding predators is a goal of many bird conservation programs. However, new exclosure projects should be carefully evaluated to assess the potential risks of disturbance. We tested the effectiveness of predator exclosure fences (hereafter, fences) for nests of critically endangered Florida Grasshopper Sparrows (Ammodramus savannarum floridanus) at a dry prairie site (Three Lakes; 2015–2018) and a pasture site (the Ranch; 2015–2016) in Osceola County, Florida, USA. We installed fences at nests an average of 8 days after the start of incubation, and nest abandonment after fence installation was rare (2 of 149 installations). Predation was the leading cause of failure for unfenced nests at both sites (48–73%). At Three Lakes, nest cameras revealed that mammals and snakes were responsible for 61.5% and 38.5% of predation events, respectively, at unfenced nests. Fences reduced the daily probability of predation (0.016 for fenced nests vs. 0.074 for unfenced nests). The probability that a fenced nest would survive from discovery to fledging was more than double that of unfenced nests (60.4% vs. 27.7%). However, we found no difference in daily nest survival at the Ranch between the year before nests were fenced (2015; 0.874) and the year when all but one nest were fenced (2016; 0.867) because red imported fire ants (Solenopsis invicta) were responsible for 86% of predation events at fenced nests at the Ranch. The use of cameras at fenced nests revealed that site‐specific differences in nest predators explained variation in fence efficiency between sites. Our fence design may be useful for other species of grassland birds, but site‐specific predator communities and species‐specific response of target bird species to fences should be assessed before installing fences at other sites.     

Experiments on clutch size and nest size in passerine birds

Summary Results of experiments on three passerine species suggest that brood size may be constrained by nest size, since the breeding success of pairs provided with large nestcups was greater than that of those provided with small artificial nestcups. These results may have important implications, e.g. to the design of experiments involving manipulation of clutch and brood size. A small nestcup is requisite for successful hatching during the incubation period, but a large one for successful rearing during the nestling period. In nature this difference may select for types of nesting materials that are elastic, such as mosses and lichens. However, experiments showed that such materials rapidly absorb rainwater but only slowly dry out. In addition, because large nests dry out more slowly than small nests, selection will favour small nests among those open-nesting species that have exposed nests. A further possible nest size constraint on open-nesters is nest predation. However, no difference in the predation rate was found in experiments with small and large artificial nests.     

Mixed species nesting associations in Darwin's tree finches: nesting pattern predicts predation outcome

Predation is the main cause of passerine nesting failure. Traditionally, large intraspecific group size is thought to accrue individuals with fitness benefits from increased predator vigilance and hence lower predation risk. To date, few studies have investigated interspecific group size in relation to predation risk. In the present study, we examined predation outcome in Darwin's small tree finch, Camarhynchus parvulus , in nests with many or few interspecific neighbours. We tested the predictions: (1) nests in mixed associations have lower predation than do more solitary nests; (2) mixed species nesting associations covary with nest site vegetation characteristics; (3) older (i.e. presumably experienced) males more commonly nest in mixed associations than younger males; (4) older males select more concealed nesting sites; and (5) controlling male age, females prefer to pair with males in mixed associations than at solitary nests. Almost half of all nests occurred in mixed associations (46%) compared to solitary nests (54%), and the overall distribution of nests was decidedly nonrandom, displaying a bimodal pattern. Nest site vegetation characteristics of the focal species were inconsistently associated with nesting pattern, but older males did select more concealed nesting sites. Controlling differences in surrounding vegetation characteristics, mixed nesting associations experienced markedly lower predation than solitary nests, and females showed a preference for males in mixed associations, as demonstrated by higher male pairing success.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 313–324.     

Conspecific nest parasitism is associated with inequality in nest predation risk in the common goldeneye (Bucephala clangula)

Previous studies of the role of nest predation in conspecificnest parasitism have not taken into account the possibilitythat predation risk may not be randomly distributed among nestsites and that breeding individuals may use different cues toassess the risk and adjust their reproductive tactic betweenyears accordingly. Especially in cavity-nesting species, therole of nest predation in conspecific nest parasitism has beendownplayed, while the role of nest site limitation has beenhighlighted. Using both observational and experimental data,I show that in the common goldeneye (Bucephala clangula), acavity-nesting species in which conspecific nest parasitism iscommon, predation risk varies considerably between nest sitesand does not follow a random expectation. The inequality inpredation risk between nest sites also showed up in the occurrenceof parasitized nests in an experimental setup. Nests parasitizedin year t were more frequent in those nest sites that were notdepredated during the previous nesting attempt in year t - nthan in nest sites that were depredated and in control nestsites that had not been used for nesting before. A nest siteaddition experiment revealed that conspecific nest parasitismwas not associated with nest site limitation. My findings givesupport for the hypothesis that nest predation is an importantecological factor explaining conspecific nest parasitism ingoldeneyes.     

Comparative nest survival of three sympatric loon species breeding in the Arctic

Identifying factors influencing nest survival among sympatric species is important for understanding and managing sources of variation in population dynamics of individual species. Three species of loons nest sympatrically in northern Alaska and differ in body size, life history characteristics, and population trends. We tested the effects of competition, nest site selection, and water level variations on nest survival of Pacific Gavia pacifica, yellow‐billed G. adamsii, and red‐throated loons G. stellata on the Arctic Coastal Plain in Alaska. Although overall nest survival rates did not differ between species, the factors influencing nest survival varied. Nest site selection influenced nest survival for Pacific and yellow‐billed loons, with both species having high nest survival when nesting on islands and peninsulas, likely due to a reduction in access by terrestrial predators. However, on mainland shorelines, Pacific loons had lower nest survival than yellow‐billed loons, and used a higher proportion of vegetation mats for nest sites suggesting that their smaller body size makes them less adept at nest defense. Nest site selection did not influence nest survival of red‐throated loons corresponding to our result of no nest site preferences by this species. Initiation date had a strong influence on nest survival for Pacific and yellow‐billed loons with nests laid earlier having higher survival. Pacific and yellow‐billed loon nests were susceptible to flooding due to precipitation, which contrasted with red‐throated loons that nest on smaller lakes with lower water level variations. Competition did not affect nest survival for any of the species likely due to most territorial encounters occurring prior to incubation. The only influence we found on red‐throated loon nest survival was differences among years. Our results indicate that loons chose nest sites based on predation risk and that factors influencing breeding success of closely related species may differ under similar breeding conditions.     

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.     

Plasticity of parental care under the risk of predation: how much should parents reduce care?

Predation can be an important agent of natural selection shaping parental care behaviours, and can also favour behavioural plasticity. Parent birds often decrease the rate that they visit the nest to provision offspring when perceived risk is high. Yet, the plasticity of such responses may differ among species as a function of either their relative risk of predation, or the mean rate of provisioning. Here, we report parental provisioning responses to experimental increases in the perceived risk of predation. We tested responses of 10 species of bird in north temperate Arizona and subtropical Argentina that differed in their ambient risk of predation. All species decreased provisioning rates in response to the nest predator but not to a control. However, provisioning rates decreased more in species that had greater ambient risk of predation on natural nests. These results support theoretical predictions that the extent of plasticity of a trait that is sensitive to nest predation risk should vary among species in accordance with predation risk.     

Nest predation is little affected by parental behaviour and nest site in two African Sylvia warblers

Parental behaviour and nest site are supposed to affect nest predation in birds. Few nest visits and high nest attentiveness are assumed to lead to low predation rates. Poorly concealed nests are thought to be more likely to be preyed upon than well concealed nests. Studies on the relationship between parental behaviour, nest site, and nest predation are rare and none have, so far, been conducted in the Afrotropics. We studied the effect of nest site, nest visitation rate, and nest attentiveness on the nest predation rate of the two tropical warblers Sylvia boehmi and Sylvia lugens in Kenya. Parental behaviour and predation on nests of 13 breeding pairs of both species were observed daily in two consecutive breeding seasons. In both species, parental activity at the nest was low [0.9 trips to the nest in 30 min during incubation, maximum 4.6 (S. boehmi) and 5.8 (S. lugens) trips to the nest in the nestling stage]. Predation rates in both species were high (Mayfield nest success 19.4 and 33.2%). Our analysis revealed only weak evidence for an effect of nest site, nest visitation rate, and nest attentiveness on the predation rate. It is suggested that smaller clutches of tropical in comparison to northern temperate birds result from lower feeding rates in tropical ecosystems with high predation rates (Skutchs hypothesis). The underlying mechanism could not be proven in this study.     

Microhabitat structure and avian nest predation risk in an open Argentinean woodland: an experimental study

We studied the effect of the general structure of the nest plant, especially the presence of thorns, and the structural homogeneity of the nest patch, on the vulnerability of nests to predation, using natural and artificial nests. Artificial nests placed in non-thorny plants had a significantly lower predation rate and higher daily survival rate than those in thorny plants. The addition of a ‘thorny microhabitat’ around the immediate proximity of nests placed in non-thorny plants did not have any effect on vulnerability of nests to predation. Conversely, natural nests were located in patches of habitat with a higher density of the species of plant that supported the nest compared to patches selected at random. However, daily survival rate was similar for natural nests placed in patches with a higher or lower density of the species of nest plant in the four bird species analysed. Similarly, survival of artificial nests did not increase with the presence of a higher number of plants similar to the nest plant in the nest patch. Thus, the observed patterns of survival for natural and artificial nests did not seem to support the potential prey-site hypotheses. Birds appeared to be the main nest predators in this ecosystem. Behavioural aspects of the identified predators and habitat structure could explain the lack of effect of thorns and nest patch characteristics on nest survival.     

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.     

Sources of variation in the nesting success of understory tropical birds

Survival of offspring is a key fitness component and, for birds, the threat of predation on nests is especially influential. Data on rates of nest success from tropical regions are comparatively few, conservation‐relevant, and essential for assessing the validity of models comparing the life histories and behavior or birds across latitudinal gradients. We monitored over 2 000 nests in the lowland forests of central Panama and, using the logistic exposure to model the fate of nests, explored the importance of variation in rate of nest success according to type of nest, height of nests, among years, in early versus late nests, and at different stages of the nest cycle. Analyses of over 1 400 nests for 18 species revealed considerable variation among species in the daily survival rate of nests (range among 18 species=0.91 to 0.98), but nest type and stage of the nesting cycle were generally influential on the probability of nest success. Cavity or enclosed nesters experienced greater nest success than open cup nesters and rates of nest loss were generally greatest in the nestling stage. We found limited evidence that height of nests affected probability of success, but no indication that timing of nesting effort was influential. Despite the occurrence of a severe ENSO event during our sampling, annual variation in nest success was not consistent among species. Interspecific variation in the rates and patterns of nest predation in our study, coupled with reports of high rates of nest loss at temperate latitudes, lead us to question long standing assumptions about latitudinal trends in rates of nest loss. We urge further work to understand the implications of nest predation on the evolutionary ecology of tropical birds.     

Nest placement and nesting success in two finch species colonizing a recently established plantation in an arid region

This study examines variations in nest placement and its effects on nesting success in the Desert Finch (Rhodospiza obsoleta) and the Linnet (Carduelis cannabina) which coexist in a recently established plantation (novel habitat) in a desert region in Jordan. A comparison with other habitats (old plantation in the desert and native, Mediterranean-type, habitat of the Linnet) showed that each species displayed similarity in nest placement patterns across habitats. Nest concealment predicted nesting outcome in the Linnet, whereas nest height predicted nesting outcome in the Desert Finch. Daily survival rates were significantly higher in Desert Finch nests compared to the Linnet's, perhaps because Desert Finches placed nests higher in the novel habitat, which lacked aerial nest predators. In accordance with predation theories of life history evolution, we provide evidence that variation in nest placement among the two species in a novel habitat resulted from previous exposure to different predation risks.