Dynamic, multi-scale analyses indicate site- and landscape-level forest cover drive Yellow-billed and Black-billed Cuckoo interannual turnover.

Studies of habitat use in breeding birds often assume species have relatively stable breeding distributions. Some species, however, display considerable year-to-year variability, complicating efforts to determine suitable or preferred habitats. After returning to their breeding range, Black-billed Cuckoos (Coccyzus erythropthalmus) and Yellow-billed Cuckoos (C. americanus) are thought to range widely before nesting, resulting in high rates of interannual breeding-site turnover, potentially contributing to conflicting habitat associations found in past studies. However, difficulty detecting these rare and declining species could lead to overinflated estimates of interannual turnover. Using broadcast surveys to increase detection probability, we collected detection/non-detection data in 2019 and 2020 at 41 publicly owned sites in Illinois and performed a dynamic, multi-scale occupancy analysis for each species to separate detection probability from potential interannual turnover and determine landscape and small-scale variables driving habitat use and occupancy dynamics. We found strong support for interannual turnover for both species based on poor performance of non-dynamic models and variation in estimated annual occupancy (20% and 21% increase between years for Black-billed and Yellow-billed Cuckoos, respectively). Black-billed Cuckoos persisted at sites with less forest in the surrounding landscape and used areas with denser understory vegetation. Yellow-billed Cuckoos colonized sites with greater canopy cover, avoided developed landscapes, and used areas with a shorter subcanopy layer. The dynamic nature of habitat use in these two cuckoo species suggests the importance of coordinating management and conservation across a broader spatial scale. Managing for larger patches of dense shrubs in less forested landscapes would benefit Black-billed Cuckoos while Yellow-billed cuckoos would benefit from management creating forested areas with open understories in less-developed landscapes.

Water depth influences survival and predator-specific patterns of nest loss in three secretive marsh bird species.

Wetlands have become increasingly rare in the United States, negatively influencing wetland-dependent birds, and many remaining wetlands are intensively managed through seasonal dewatering mimicking historic flood pulses during spring and summer. However, water around nests may provide protection from terrestrial predators, and lowering water levels during the breeding season of wetland birds may increase predation risk and exacerbate marsh bird population declines. Understanding interactions between water depth, nesting marsh birds, and nest predators is critical to aid managers in developing a multi-species management approach in emergent wetlands. During the 2020 and 2021 breeding seasons, we examined nest survival of 148 marsh bird nests (American Coot, Fulica americana, n = 1; Common Gallinule, Gallinula galeata, n = 64; and Least Bittern; Ixobrychus exilis, n = 83) and installed cameras at 78 nests to identify predators at a large, restored floodplain wetland in Illinois where the primary management technique is seasonal water removal to stimulate germination of moist soil plants. We found nest predation of, and abandonment by, Least Bittern and Common Gallinule were related to shallower water, and early season, high volume dewatering. Least Bitterns nested more commonly along wetland edges and nests farther from the shore were more likely to survive. Similarly, we found mammalian depredation of nests and nest abandonment decreased when deeper water was present around nests. Alternatively, snake predation was observed earlier in the year prior to water removal from inundated emergent vegetation. Our results demonstrate water depth may be an important deterrent of nest predators, especially mammals, during the breeding season. Further, we recommend managers delay dewatering until after the nesting season at sites where management for conservation-priority marsh birds is a focus.

Extensive regional variation in the phenology of insects and their response to temperature across North America.

Climate change models often assume similar responses to temperatures across the range of a species, but local adaptation or phenotypic plasticity can lead plants and animals to respond differently to temperature in different parts of their range. To date, there have been few tests of this assumption at the scale of continents, so it is unclear if this is a large-scale problem. Here, we examined the assumption that insect taxa show similar responses to temperature at 96 sites in grassy habitats across North America. We sampled insects with Malaise traps during 2019-2021 (N = 1041 samples) and examined the biomass of insects in relation to temperature and time of season. Our samples mostly contained Diptera (33%), Lepidoptera (19%), Hymenoptera (18%), and Coleoptera (10%). We found strong regional differences in the phenology of insects and their response to temperature, even within the same taxonomic group, habitat type, and time of season. For example, the biomass of nematoceran flies increased across the season in the central part of the continent, but it only showed a small increase in the Northeast and a seasonal decline in the Southeast and West. At a smaller scale, insect biomass at different traps operating on the same days was correlated up to ~75 km apart. Large-scale geographic and phenological variation in insect biomass and abundance has not been studied well, and it is a major source of controversy in previous analyses of insect declines that have aggregated studies from different locations and time periods. Our study illustrates that large-scale predictions about changes in insect populations, and their causes, will need to incorporate regional and taxonomic differences in the response to temperature.

Long-term monitoring reveals widespread and severe declines of understory birds in a protected Neotropical forest.

Long-term studies on the population dynamics of tropical resident birds are few, and it remains poorly understood how their populations have fared in recent decades. Here, we analyzed a 44-y population study of a Neotropical understory bird assemblage from a protected forest reserve in central Panama to determine if and how populations have changed from 1977 to 2020. Using the number of birds captured in mist nets as an index of local abundance, we estimated trends over time for a diverse suite of 57 resident species that comprised a broad range of ecological and behavioral traits. Estimated abundances of 40 (∼70%) species declined over the sampling period, whereas only 2 increased. Furthermore, declines were severe: 35 of the 40 declining species exhibited large proportional losses in estimated abundance, amounting to ≥50% of their initial estimated abundances. Declines were largely independent of ecology (i.e., body mass, foraging guild, or initial abundance) or phylogenetic affiliation. These widespread, severe declines are particularly alarming, given that they occurred in a relatively large (∼22,000-ha) forested area in the absence of local fragmentation or recent land-use change. Our findings provide robust evidence of tropical bird declines in intact forests and bolster a large body of literature from temperate regions suggesting that bird populations may be declining at a global scale. Identifying the ecological mechanisms underlying these declines should be an urgent conservation priority.

Host community-wide patterns of post-fledging behavior and survival of obligate brood parasitic brown-headed cowbirds.

The antagonistic arms races between obligate brood parasites and their hosts provide critical insights into coevolutionary processes and constraints on the evolution of life history strategies. In avian brood parasites-a model system for examining host-parasite dynamics-research has primarily focused on the egg and nestling stage, while far less is known about the behavior and ecology of fledgling and juvenile brood parasites. To provide greater insights into the post-fledging period of generalist brood parasites, we used handheld and automated telemetry systems to examine the behavior and survival of fledgling brown-headed cowbirds (Molothrus ater). Our host community-wide analysis (data on cowbirds fledged from different host species were pooled) shows that fledgling cowbirds' follow patterns of movement and survival found across the post-fledging literature on parental passerine species. Cowbird fledgling survival was lowest during the first 3 days post-fledging, whereas daily rates of survival neared 100% after about 16 days post-fledging. Cowbird daytime post-fledging activity rates, perch heights, and distance from the natal area all increased with fledging age and young generally gained independence from host parents at 3-4 weeks post-fledging, with approximately the same latency as has been observed in studies on fledglings of cowbird host species. Our research demonstrates how automated telemetry systems can overcome past methodological limitations in post-fledging research and provides an important foundation for future studies examining adaptations that cowbirds and other brood parasites use to exploit hosts during the post-fledging period.

Ground nesting by arboreal American robins (Turdus migratorius).

Animals with dependent and vulnerable young need to decide where to raise their offspring to minimize ill effects of weather, competition, parasitism, and predation. These decisions have critical fitness consequences through impacting the survival of both adults and progeny. Birds routinely place their nest in specific sites, allowing species to be broadly classified based on nest location (e.g., ground- or tree-nesting). However, from 2018 to 2020, we observed 24 American robin (Turdus migratorius) nests placed not on their species-typical arboreal substrates or human-made structures but on the ground at a predator-rich commercial tree-farm in Illinois, USA. This behavior does not appear to be in response to competition and did not affect nest daily survival rate but was restricted to the early half of the breeding season. We hypothesize that ground nesting may be an adaptive response to avoid exposure and colder temperatures at sites above the ground early in the breeding season or a nonadaptive consequence of latent robin nest-placement flexibility.

Referential alarm calling elicits future vigilance in a host of an avian brood parasite.

Yellow warblers (Setophaga petechia) use referential 'seet' calls to warn mates of brood parasitic brown-headed cowbirds (Molothrus ater). In response to seet calls during the day, female warblers swiftly move to sit tightly on their nests, which may prevent parasitism by physically blocking female cowbirds from inspecting and laying in the nest. However, cowbirds lay their eggs just prior to sunrise, not during daytime. We experimentally tested whether female warblers, warned by seet calls on one day, extend their anti-parasitic responses into the future by engaging in vigilance at sunrise on the next day, when parasitism may occur. As predicted, daytime seet call playbacks caused female warblers to leave their nests less often on the following morning, relative to playbacks of both their generic anti-predator calls and silent controls. Thus, referential calls do not only convey the identity or the type of threat at present but also elicit vigilance in the future to provide protection from threats during periods of heightened vulnerability.

Egg Incubation Temperature Affects Development of Innate Immune Function in Nestling American Robins (Turdus migratorius).

The innate immune system provides important first-line defenses against invading pathogens and is considered especially important for developing organisms. However, we know little about how early-life conditions influence these defenses in wild animals. For oviparous species such as birds, embryonic development occurs in the egg, which can be subject to variation in thermal conditions. There is evidence from cavity-nesting species and species with precocial young that reduced incubation temperatures can result in reduced measures of innate immunity. Whether and how this thermal variation impacts innate immunity for open-cup-nesting species with altricial offspring has not been examined. In this study, we experimentally manipulated egg incubation temperature for American robins (Turdus migratorius) and compared the bacteria-killing ability (BKA) of the nestlings' blood plasma. We collected baseline and poststressor samples on day 7 and day 10 after hatch to gain additional insights into the ontogeny of this immune measure, as well as into whether any changes were linked to levels of the glucocorticoid hormone corticosterone (CORT). We found that nestlings incubated at the low treatment (36.1°C) had significantly reduced BKA compared with nestlings incubated at the high treatment (37.8°C) when controlling for the posthatch nest environment. We also documented a significant reduction in poststressor levels of BKA, as well as an increase in BKA from day 7 to day 10. We found a weak inverse association between CORT and BKA but no other indications that BKA was mediated via treatment-induced variation in CORT. Our results suggest that incubation temperature can affect development of innate immunity in open-cup-nesting passerines.

Avian fitness consequences match habitat selection at the nest-site and landscape scale in agriculturally fragmented landscapes.

Habitat selection theory suggests that when choosing breeding sites, animals should choose the best available habitat; however, studies show that individuals fail to choose habitats that maximize their fitness especially in drastically altered landscapes. Many studies have focused on selection at single scales, often using a single measure of fitness. However, links between habitat selection and fitness may vary depending on the spatial scale and measure of fitness, especially in situations where agricultural land use has altered the surrounding landscape.We examined multiscale habitat selection and fitness measures of the Bell's Vireo (Vireo bellii) and Willow Flycatcher (Empidonax traillii) using data collected in agriculturally fragmented landscapes.We found evidence for selection of nest sites with dense understory, larger patches, and increasing restored habitat cover and decreasing forest cover in the surrounding landscape.For both focal species, selection for dense understory significantly increased nest survival; however, there appeared to be no concurrent benefit for fledgling production. Selection for broader scale features positively influenced nest survival for the Willow Flycatcher with no concurrent benefit for fledgling production. The observed mismatches may be due to anthropogenic habitat fragmentation at broader scales or may represent reproductive trade-offs for a fitness benefit not measured in this study.Fine-scale habitat selection decisions by our focal species appear to match fitness outcomes, whereas habitat selection at broader scales only provided fitness benefits for the Willow Flycatcher. Though providing no fledgling production benefit, when combined with suitably dense nesting habitat, larger patches in landscapes with greater amounts of restored habitat cover for Bell's Vireo and lower amounts of forest cover for Willow Flycatcher will produce more fledglings per unit area than smaller patches in landscapes with less restored habitat and more forest cover, respectively, which could help inform conservation decisions for these at-risk species.

Mimicry-dependent lateralization in the visual inspection of foreign eggs by American robins.

Brain lateralization, or the specialization of function in the left versus right brain hemispheres, has been found in a variety of lineages in contexts ranging from foraging to social and sexual behaviours, including the recognition of conspecific social partners. Here we studied whether the recognition and rejection of avian brood parasitic eggs, another context for species recognition, may also involve lateralized visual processing. We focused on American robins (Turdus migratorius), an egg-rejecter host to occasional brood parasitism by brown-headed cowbirds (Molothrus ater) and tested if robins preferentially used one visual hemifield over the other to inspect mimetic versus non-mimetic model eggs. At the population level, robins showed a significantly lateralized absolute eyedness index (EI) when viewing mimetic model eggs, but individuals varied in left versus right visual hemifield preference. By contrast, absolute EI was significantly lower when viewing non-mimetic eggs. We also found that robins with more lateralized eye usage rejected model eggs at higher rates. We suggest that the inspection and recognition of foreign eggs represent a specialized and lateralized context of species recognition in this and perhaps in other egg-rejecter hosts of brood parasites.

Estimating apparent survival of songbirds crossing the Gulf of Mexico during autumn migration.

Many migratory bird species are declining, and the migratory period may limit populations because of the risk in traversing large geographical features during passage. Using automated radio-telemetry, we tracked 139 Swainson's thrushes (Catharus ustulatus) departing coastal Alabama, USA and crossing the Gulf of Mexico to arrive in the Yucatan Peninsula, Mexico during autumn. We estimated apparent survival and examined how extrinsic (weather variables and day of year) and intrinsic (fat load, sex and age) factors influenced survival using a mark-recapture approach. We also examined how favourability of winds for crossing the Gulf varied over the past 25 years. Fat load, day of year and wind profit were important factors in predicting which individuals survived crossing the Gulf. Survival estimates varied with wind profit and fat, but generally, fat birds departing on days with favourable wind profits had an apparent survival probability of greater than 0.90, while lean individuals with no or negative wind profits had less than 0.33. The proportion of favourable nights varied within and among years, but has increased over the last 25 years. While conservation strategies cannot improve extrinsic factors, they can provide opportunities for birds to refuel before crossing large geographical features through protecting and creating high-quality stopover sites.

Incubation temperature impacts nestling growth and survival in an open-cup nesting passerine.

For oviparous species such as birds, conditions experienced while in the egg can have long-lasting effects on the individual. The impact of subtle changes in incubation temperature on nestling development, however, remains poorly understood, especially for open-cup nesting species with altricial young. To investigate how incubation temperature affects nestling development and survival in such species, we artificially incubated American robin (Turdus migratorius) eggs at 36.1°C ("Low" treatment) and 37.8°C ("High" treatment). Chicks were fostered to same-age nests upon hatching, and we measured mass, tarsus, and wing length of experimental nestlings and one randomly selected, naturally incubated ("Natural"), foster nest-mate on days 7 and 10 posthatch. We found significant effects of incubation temperature on incubation duration, growth, and survival, in which experimentally incubated nestlings had shorter incubation periods (10.22, 11.50, and 11.95 days for High, Low, and Natural eggs, respectively), and nestlings from the Low treatment were smaller and had reduced survival compared to High and Natural nestlings. These results highlight the importance of incubation conditions during embryonic development for incubation duration, somatic development, and survival. Moreover, these findings indicate that differences in incubation temperature within the natural range of variation can have important carryover effects on growth and survival in species with altricial young.

Correction: Does flooding effect the apparent survival and body condition of a ground foraging migrant passerine?

[This corrects the article DOI: 10.1371/journal.pone.0175179.].

Rates of parasitism, but not allocation of egg resources, vary among and within hosts of a generalist avian brood parasite.

Brown-headed cowbirds (Molothrus ater) deposit their eggs into the nests of other birds, which then raise the cowbird chick. Female cowbirds thus have limited options for impacting their offspring's development via maternal effects compared to most other passerines. Cowbirds can impact their offspring's phenotype by choosing among potential host nests, and by adjusting egg resources based on host characteristics. To examine whether cowbirds exhibit either or both of these strategies, we investigated rates of cowbird parasitism and egg investment (egg size, yolk-to-albumen ratio, and yolk testosterone and androstenedione) among and within host species in a shrubland bird community. We found that the probability of being parasitized by cowbirds, controlling for host status as a cowbird egg accepter or rejecter and ordinal date, varied significantly among host species, indicating an apparent preference for some hosts. Parasitism rates did not differ with host size, however, and despite variation in cowbird egg size among host species, this variation was not related to host size or cowbird preference. Among host species with eggs that are larger than those of the cowbird, cowbirds were significantly more likely to parasitize nests with relatively smaller eggs, whereas parasitism rates did not vary with relative egg size in host species with smaller eggs. There was no evidence for variation in cowbird egg components among or within host species. Our data indicate that cowbirds discriminate among host nests, but do not appear to adjust the composition of their eggs based on inter- or intraspecific host variation.

Does flooding effect the apparent survival and body condition of a ground foraging migrant passerine?

Natural disturbances play a fundamental role in maintaining habitat and landscape heterogeneity; however, these events can also have negative effects on some species. While we know that disturbances can reduce habitat quality for many species, leading to diminished populations and altered community structure, the effect of these events on individuals that continue to occupy affected areas remains unknown. We focused on understanding the impact of flood-mediated reduction of habitat quality on Swainson's Warblers (Limnothlypis swainsonii). In 2008, a catastrophic flood event occurred on the Mississippi River and its tributaries, severely affecting one of two locations where we had studied territorial males since 2004. To determine the impact of flooding on this species, we evaluated how body condition and apparent survival of males differed between locations and in pre-flood (2004-2007) and post-flood (2008-2010) periods. Body condition did not differ between locations after the flood, suggesting that flooding did not cause food limitation for this obligate ground forager. Apparent survival in the post-flood period was lower at both locations and led to near population extirpation at the heavily flood-impacted site. Overall, this study demonstrates the vulnerability of species to extreme hydrological events, an increasing threat due to climate change. Future research should focus on identifying species that are vulnerable to these events and determining appropriate conservation strategies. Conservation for the Swainson's Warbler should focus on identifying and conserving the highest elevation, least flood prone areas within bottomland hardwood forests and managing those areas for thick understory vegetation.

Fat, weather, and date affect migratory songbirds' departure decisions, routes, and time it takes to cross the Gulf of Mexico.

Approximately two thirds of migratory songbirds in eastern North America negotiate the Gulf of Mexico (GOM), where inclement weather coupled with no refueling or resting opportunities can be lethal. However, decisions made when navigating such features and their consequences remain largely unknown due to technological limitations of tracking small animals over large areas. We used automated radio telemetry to track three songbird species (Red-eyed Vireo, Swainson's Thrush, Wood Thrush) from coastal Alabama to the northern Yucatan Peninsula (YP) during fall migration. Detecting songbirds after crossing ∼1,000 km of open water allowed us to examine intrinsic (age, wing length, fat) and extrinsic (weather, date) variables shaping departure decisions, arrival at the YP, and crossing times. Large fat reserves and low humidity, indicative of beneficial synoptic weather patterns, favored southward departure across the Gulf. Individuals detected in the YP departed with large fat reserves and later in the fall with profitable winds, and flight durations (mean = 22.4 h) were positively related to wind profit. Age was not related to departure behavior, arrival, or travel time. However, vireos negotiated the GOM differently than thrushes, including different departure decisions, lower probability of detection in the YP, and longer crossing times. Defense of winter territories by thrushes but not vireos and species-specific foraging habits may explain the divergent migratory behaviors. Fat reserves appear extremely important to departure decisions and arrival in the YP. As habitat along the GOM is degraded, birds may be limited in their ability to acquire fat to cross the Gulf.

Patch size and edge proximity are useful predictors of brood parasitism but not nest survival of grassland birds.

Declines of migratory birds have led to increased focus on causative factors for these declines, including the potential adverse effects of habitat fragmentation on reproductive success. Although numerous studies have addressed how proximity to a habitat edge, patch size, or landscape context influence nest survival or brood parasitism, many have failed to find the purported effects. Furthermore, many have sought to generalize patterns across large geographic areas and habitats. Here, we examined evidence for effects of edge proximity, patch size, and landscape context on nest survival and brood parasitism of grassland birds, a group of conservation concern. The only consistent effect was a positive association between edge proximity and brood parasitism. We examined effects of patch size on nest survival (37 studies) and brood parasitism (30 studies) representing 170 and 97 different estimates, respectively, with a total sample size of > 14000 nests spanning eastern North America. Nest survival weakly increased with patch size in the Great Plains, but not in the Midwestern or Eastern United States, and brood parasitism was inversely related to patch size and consistently greater in the Great Plains. The consistency in brood parasitism relative to nest survival patterns is likely due to parasitism being caused by one species, while nest survival is driven by a diverse and variable suite of nest predators. Often, studies assume that predators responsible for nest predation, the main driver of nest success, either are the same or exhibit the same behaviors across large geographic areas. These results suggest that a better mechanistic understanding of nest predation is needed to provide meaningful conservation recommendations for improving grassland bird productivity, and that the use of general recommendations across large geographic areas should only be undertaken when sufficient data are available from all regions.

Juvenile survival in a neotropical migratory songbird is lower than expected.

Attempts to estimate and identify factors influencing first-year survival in passerines, survival between fledging and the first reproductive attempt (i.e. juvenile survival), have largely been confounded by natal dispersal, particularly in long-distance migratory passerines. We studied Prothonotary Warblers (Protonotaria citrea) breeding in nest boxes to estimate first-year survival while accounting for biases related to dispersal that are common in mark-recapture studies. The natal dispersal distribution (median = 1420 m; n = 429) and a distance-dependent recruitment rate, which controls for effects of study site configuration, both indicated a pattern of short-distance natal dispersal. This pattern was consistent with results of a systematic survey for birds returning outside the nest box study sites (up to 30 km in all directions) within a majority (81%) of total available bottomland forest habitat, further suggesting that permanent emigration outside of the study system was rare. We used multistate mark-recapture modeling to estimate first-year survival and incorporated factors thought to influence survival while accounting for the potential confounding effects of dispersal on recapture probabilities for warblers that fledged during 2004-2009 (n = 6093). Overall, the average first-year survival for warblers reared without cowbird nestmates was 0.11 (95% CI = 0.09-0.13), decreased with fledging date (0.22 early to 0.03 late) and averaged 40% lower for warblers reared with a brood parasite nestmate. First-year survival was less than half of the rate thought to represent population replacement in migratory passerines (∼0.30). This very low rate suggests that surviving the first year of life for many Neotropical migratory species is even more difficult than previously thought, forcing us to rethink estimates used in population models.

Implications of spatial patterns of roosting and movements of American robins for West Nile virus transmission.

The arrival of West Nile virus (WNV) in North America has led to interest in the interaction between birds, the amplification hosts of WNV, and Culex mosquitoes, the primary WNV vectors. American robins (Turdus migratorius) are particularly important amplification hosts of WNV, and because the vector Culex mosquitoes are primarily nocturnal and feed on roosting birds, robin communal roosting behavior may play an important role in the transmission ecology of WNV. Using data from 43 radio-tracked individuals, we determined spatial and temporal patterns of robin roosting behavior, and how these patterns related to the distribution of WNV-infected mosquitoes. Use of the communal roost and fidelity to foraging areas was highly variable both within and among individual robins, and differed markedly from patterns documented in a previous study of robin roosting. Although there were clear seasonal patterns to both robin roosting and WNV occurrence, there was no significant relationship between communal roosting by robins and temporal or spatial patterns of WNV-positive mosquitoes. Our results suggest that, although robins may be important as WNV hosts, communal roosts are not necessarily important for WNV amplification. Other factors, including the availability and distribution of high-quality mosquito habitat and favorable weather for mosquito reproduction, may influence the importance of robin roosts for local WNV amplification and transmission.

Identifying predators clarifies predictors of nest success in a temperate passerine.

1. Nest predation negatively affects most avian populations. Studies of nest predation usually group all nest failures when attempting to determine temporal and parental activities, habitat or landscape predictors of success. Often these studies find few significant predictors and interpret patterns as essentially random. 2. Relatively little is known about the importance of individual predator species or groups on observed patterns of nest success, and how the ecology of these predators may influence patterns of success and failure. 3. In 2006 and 2007, time-lapse, infrared video systems were deployed at nests of Swainson's warblers (Limnothlypis swainsonii Audubon) in east-central Arkansas to identify dominant nest predators and determine whether factors predicting predation differed among these predators. 4. Analysis of pooled data yielded few predictors of predation risk, whereas separate analyses for the three major predator groups revealed clear, but often conflicting, patterns. 5. Predation by ratsnakes (Elaphe obsoleta) and raptors was more common during the nestling period, whereas predation by brown-headed cowbirds (Molothrus ater) occurred more during incubation. Additionally, the risk of predation by raptors and cowbirds decreased throughout the breeding season, whereas ratsnake predation risk increased. 6. Contrary to expectations, predation by ratsnakes and cowbirds was more common far from edges, whereas raptor predation was more common close to agricultural edges. 7. Collectively, our results suggest that associating specific predators with the nests they prey on is necessary to understand underlying mechanisms.