Differences in Predators of Artificial and Real Songbird Nests: Evidence of Bias in Artificial Nest Studies

Abstract:  In the past two decades, many researchers have used artificial nests to measure relative rates of nest predation. Recent comparisons show that real and artificial nests may not be depredated at the same rates, but no one has examined the mechanisms underlying these patterns. We determined differences in predator-specific predation rates of real and artificial nests. We used video cameras to monitor artificial nests baited with quail and plasticine eggs and Field Sparrow ( Spizella pusilla ) and Indigo Bunting ( Passerina cyanea ) nests in field habitats in central Missouri (U.S.A.). Although daily predation estimates (all predators pooled) were similar between artificial and real nests, predators differed substantially in their depredation of artificial versus real nests. Snakes were the major predator at real nests, and raccoons ( Procyon lotor) were the major predator at artificial nests. We found strong support for models that distinguished predation between two or among three predator groups and between artificial and real nests. There was no snake predation of artificial nests, and the odds of predation of artificial nests was 115–551% (95% confidence interval) and 2–154% of the odds of predation of real nests by mammals and birds, respectively. Artificial nests with plasticine eggs could not be used reliably to identify predators. In several cases plasticine eggs were marked by mice, and raccoons were recorded on video removing the quail egg. Because biases for artificial nests were positive for some predators and negative for other predators (and could be compensating), and potentially existed for all predator groups, conclusions based on artificial nest studies should be suspect even when there is evidence that overall predation rates are similar among real and artificial nests.     

Evidence of an Edge Effect on Avian Nest Success

Abstract:  Habitat fragmentation may modify ecological patterns by increasing the importance of edge effects, including elevating rates of predation on avian nests. Conventional wisdom suggests an increased rate of predation along habitat edges, and previous reviews support this view. These reviews did not apply recent statistical approaches, however, and some were based on a small number of studies. In our meta-analysis of 64 nest-predation experiments, our results supported prior reviews of the general pattern of increased nest predation along habitat edges (  p < 0.01). We separated studies into ecologically relevant categories and found the following patterns: (1) Edge effects were more pronounced in North America and northwestern Europe than in central Europe or Central America. This result may be biased, however, by the different habitats studied in the regions. (2) Marshes and deciduous forests had significant edge effects, whereas edge effects were not apparent in coniferous forests, tropical forests, or fields. (3) Ground and natural nest studies were more likely to exhibit edge effects. (4) Edge effects were detected in studies that used quail eggs and real eggs. (5) Edge effects were not significant when artificial nests were exposed for typical incubation periods, but were significant for shorter exposures. Three alternative hypotheses may explain increased nest predation along edges. The edge-effects hypothesis states that increased nest losses along edges are the result of the habitat discontinuity. The landscape-structure hypothesis states that more fragmented landscapes are more heavily depredated by nest predators. The human-disturbance hypothesis states that near anthropogenic edges increased nest predation is related to human activities. Nest-predation experiments should be placed in a landscape context to reveal differences between the hypotheses.     

Does egg colour affect predation rate on open passerine nests?

The breeding success of many passerines is strongly reduced by egg predation. The adaptive significance of egg crypsis in open nesters is often taken for granted, but visually searching predators may first detect the nest or adult bird and not the eggs. Götmark predicted that selection should favour egg crypsis in the absence of conspicuous nests, whereas birds with conspicuous nests should have non-cryptic eggs. I compared the effect of egg colour treatment (white, blue, brown-spotted) on nest survival (1) among species characterized by different egg coloration, nest size and nest placement, and (2) between relatively well and poorly concealed nests within species. I used artificial nests (n=1,296) and eggs mimicking (except in egg colour) those of the yellowhammer (Emberiza citrinella), blackcap (Sylvia atricapilla) and song thrush (Turdus philomelos). Concurrently, I monitored survival of real nests (n=1,106). Nest survival differed among species, increased with nest concealment and throughout the breeding season, but was not significantly related to egg colour in any species. Nevertheless, the data for the yellowhammer suggest a trend in survival rates across the colour treatments. Brown eggs survived better than white eggs by 11% and 4% in 2 years, but this study had insufficient power to detect effects of this size. The results thus suggest that egg coloration in the song thrush and blackcap (shrub nesters) may be a neutral trait with regard to nest predation, whereas egg crypsis may be an anti-predation feature for the yellowhammer (ground/near-ground nester). The role of predation in the evolution of eggshell colour may vary not only between cavity and open nesters, but also across nest sites within the latter group.     

Antipredation role of clumped nesting by marsh-nesting red-winged blackbirds

Summary Red-winged blackbirds, Agelaius phoeniceus, breed in marshes in high densities and their nests are frequently clumped. Because predation is consistently the most important cause of redwing nesting mortality, high densities of breeding individuals could be an anti-predation adaptation. In our study site predation by marsh wrens, Cistothorus palustris, was the main cause of redwing nesting losses. In situations when marsh wrens were near, predation rates on redwing nests decreased with increasing female density. Group life could reduce predation because of improved nest defense, selfish herd effects, or predator dilution effects. We differentiated between these possibilities by introducing experimental colonies consisting of 3, 6, and 9 artificial nests near and away from active redwing nests. The experimental colonies near active nests suffered less predation, but predation rates were not correlated with colony size or a nest's location within the colony. Therefore, the advantage of group life in this population is probably mutual nest protection.     

Blue eggs do not reduce nest predation in the song thrush,Turdus philomelos

Summary Many passerine birds with open cup-shaped nests lay blue or blue-green eggs. In thrushes, blue eggs may be cryptic and provide camouflage by imitating spots of light on green leaves. Alternatively, egg coloration may be selectively neutral because nest predators detect nests and not eggs, or it may be maladaptive because organisms are not always well adapted to their present environment. I evaluated these hypotheses by studying predation on artificial song thrush (Turdus philomelos) nests with quail eggs, painted either white, blue, or spotted (cryptic to a human eye). Corvids were the major nest predators. For concealed as well as exposed nests, I found no differences in the predation rates of nests with white, blue, or spotted eggs. Predators apparently detected the nests, and not the eggs, first. In a second experiment, I placed egg groups without nests in trees to study the effect of color per se. The predation rate of the spotted egg groups was significantly lower than that of the white and blue egg groups, for concealed as well as exposed egg groups. These results suggest that blue eggs in the song thrush are not cryptic but may be selectively neutral or even maladaptive with regard to nest predation.     

Effects of climate and exurban development on nest predation and predator presence in the southern Appalachian Mountains (USA)

In the eastern United States, land-use and climate change have likely contributed to declines in the abundance of Neotropical migrant birds that occupy forest interiors, but the mechanisms are not well understood. We conducted a nest-predation experiment in southern Appalachian Mountain forests (North Carolina, U.S.A.) during the 2009 and 2010 breeding seasons to determine the effects of exurban development and temperature on predator presence and the average number of days until eggs in an artificial nest were disturbed by predators. We baited artificial nests with quail (Excalfactoria chinensi) eggs and monitored them for 18 days. We used clay eggs, track plates, and motion-triggered cameras to detect and identify nest predators. The average number of days a nest was undisturbed decreased as mean temperature increased and, to a lesser extent, as the density of buildings increased. Nests on the ground were more often depredated than those in trees, likely due to increased predation by opossum (Didelphis virginiana) and other carnivores. Raccoons (Procyon lotor), opossums, corvids (Corvus brachyrhynchos and Cyanocitta cristata), chipmunks (Tamias striatus), black bears (Ursus americanus), and domestic cats (Felis catus) were the most commonly detected predators. Presence of these predators did not vary as a function of mean temperature. Domestic cats and corvids were detected more frequently in plots with high rather than low densities of buildings. Forest-interior specialists and Neotropical migrants often nest in cool, high-elevation areas with low housing density. These bird species, especially those that nest on the ground, may be most vulnerable to increased nest predation if temperature and exurban development increase at higher elevations as anticipated.     

Potential Impact of Mammalian Nest Predators on Endemic Forest Birds of Western Mauna Kea, Hawaii

I used dummy nest experiments to investigate the role of nest predation by introduced mammals asa significant limiting factor for the endangered Palila ( Loxioides bailleui ) and other endemic birds on the western slope of Mauna Kea. Overall predation rates on dummy nests were extremely low. Rates were comparable to those on actual Palila nests, indicating that dummy nests give a valid representation of the dynamics of nest predation. The black rat ( Rattus rattus ) was the only important predator. Feral cats ( Felis catus ) played only a minor role, and there was no evidence to implicate house mice ( Mus musculus ) as nest predators. Four factors appear to be responsible for the low nest predation rates: (1) only a single species of predator (black rat) is involved; (2) rat densities are extremely low on Mauna Kea; (3) low prey (nest)densities preclude density-dependent predation, and (4) rats have alternative foods that are more abundant and accessible than arboreal bird nests. Since mammalian predation appears unlikely to have a significant impact on the Palila, other factors must be limiting the abundance of this endangered species.     

Nest-size variation reflecting anti-predator strategies in social spider mites of<Emphasis Type="Italic"> Stigmaeopsis</Emphasis> (Acari: Tetranychidae)

The social spider mites (Acari: Tetranychidae) of Stigmaeopsis weave dense nests on the underside of host leaves. Four species occur on the leaves of bamboo in Japan: Stigmaeopsis longus, S. celarius, S. takahashii and S. saharai. We initially reconfirmed the occurrence of distinct variation in nest size among the species. Based on the hypothesis that this variation plays a role in protecting the spider mites from predators, we looked at the behavior of the natural enemies that occur on the host plants along with members of Stigmaeopsis. We found considerable variation in the ability of nests to protect the spider-mite eggs. The smallest nests protected the eggs against three predators, whereas the largest nests protected the eggs against only one predator species. So, decreases in nest size increased egg defense. Thus we concluded that nest-size variation reflects a strategy for reducing predation.Communicated by D. Gwynne     

Evolution of Nesting Height in an Endangered Hawaiian Forest Bird in Response to a Non‐Native Predator

Abstract: The majority of bird extinctions since 1800 have occurred on islands, and non‐native predators have been the greatest threat to the persistence of island birds. Island endemic species often lack life‐history traits and behaviors that reduce the probability of predation and they can become evolutionarily trapped if they are unable to adapt, but few studies have examined the ability of island species to respond to novel predators. The greatest threat to the persistence of the Oahu Elepaio (Chasiempis ibidis), an endangered Hawaiian forest bird, is nest predation by non‐native black rats (Rattus rattus). I examined whether Oahu Elepaio nest placement has changed at the individual and population levels in response to rat predation by measuring nest height and determining whether each nest produced offspring from 1996 to 2011. Average height of Oahu Elepaio nests increased 50% over this 16‐year period, from 7.9 m (SE 1.7) to 12.0 m (SE 1.1). There was no net change in height of sequential nests made by individual birds, which means individual elepaios have not learned to place nests higher. Nests ≤3 m off the ground produced offspring less often, and the proportion of such nests declined over time, which suggests that nest‐building behavior has evolved through natural selection by predation. Nest success increased over time, which may increase the probability of long‐term persistence of the species. Rat control may facilitate the evolution of nesting height by slowing the rate of population decline and providing time for this adaptive response to spread through the population.     

Direct Effects of Cattle on Grassland Birds in Canada

Effects of grazing on grassland birds are generally thought to be indirect, through alteration of vegetation structure; however, livestock can also affect nest survival directly through trampling and other disturbances (e.g., livestock‐induced abandonment). We extracted data on nest fates from 18 grazing studies conducted in Canada. We used these data to assess rates of nest destruction by cattle among 9 ecoregions and between seasonal and rotational grazing systems. Overall, few nests were destroyed by cattle (average 1.5% of 9132 nests). Nest destruction was positively correlated with grazing pressure (i.e., stocking rate or grazing intensity), but nest survival was higher in more heavily grazed areas for some species. Because rates of destruction of grassland bird nests by cattle are low in Canada, management efforts to reduce such destruction may not be of ecological or economic value in Canada. Efectos Directos del Ganado sobre las Aves de Pastizales en Canadá     

Survival and productivity benefits to social nesting in the sweat bee <Emphasis Type="Italic">Megalopta genalis</Emphasis> (Hymenoptera: Halictidae)

Facultatively solitary and eusocial species allow for direct tests of the benefits of group living. We used the facultatively social sweat bee Megalopta genalis to test several benefits of group living. We surveyed natural nests modified for observation in the field weekly for 5 weeks in 2003. First, we demonstrate that social and solitary nesting are alternative behaviors, rather than different points on one developmental trajectory. Next, we show that solitary nests suffered significantly higher rates of nest failure than did social nests. Nest failure apparently resulted from solitary foundress mortality and subsequent brood orphanage. Social nests had significantly higher productivity, measured as new brood cells provisioned during the study, than did solitary nests. After accounting for nest failures, per capita productivity did not change with group size. Our results support key predictions of Assured Fitness Return models, suggesting such indirect fitness benefits favor eusocial nesting in M. genalis. We compared field collections of natural nests to our observation nest data to show that without accounting for nest failures, M. genalis appear to suffer a per capita productivity decrease with increasing group size. Calculating per capita productivity from collected nests without accounting for the differential probabilities of survival across group sizes leads to an overestimate of solitary nest productivity.     

Mapping risk for nest predation on a barrier island

Barrier islands and coastal beach systems provide nesting habitat for marine and estuarine turtles. Densely settled coastal areas may subsidize nest predators. Our purpose was to inform conservation by providing a greater understanding of habitat-based risk factors for nest predation, for an estuarine turtle. We expected that habitat conditions at predated nests would differ from random locations at two spatial extents. We developed and validated an island-wide model for the distribution of predated Diamondback terrapin nests using locations of 198 predated nests collected during exhaustive searches at Fisherman Island National Wildlife Refuge, USA. We used aerial photographs to identify all areas of possible nesting habitat and searched each and surrounding environments for nests, collecting location and random-point microhabitat data. We built models for the probability of finding a predated nest using an equal number of random points and validated them with a reserve set (N?=?67). Five variables in 9 a priori models were used and the best selected model (AIC weight 0.98) reflected positive associations with sand patches near marshes and roadways. Model validation had an average capture rate of predated nests of 84.14 % (26.17–97.38 %, Q1 77.53 %, median 88.07 %, Q3 95.08 %). Microhabitat selection results suggest that nests placed at the edges of sand patches adjacent to upland shrub/forest and marsh systems are vulnerable to predation. Forests and marshes provide cover and alternative resources for predators and roadways provide access; a suggestion is to focus nest protection efforts on the edges of dunes, near dense vegetation and roads.     

Nesting Success of a Neotropical Migrant in a Multiple-Use, Forested Landscape

Abstract: We studied the nesting success of an individually marked population of Kentucky Warblers ( Oporornis formosus ), a species that nests in disturbed and undisturbed forests, in a heterogeneous, managed forest site in the Shawnee National Forest in southern Illinois from 1992 to 1995. We examined the effects of forest stand type (clearcuts of various ages, tree plantations, and older forest) and distance from habitat edges on rates of nest predation and brood parasitism by Brown-headed Cowbirds (   Molothrus ater ). Brood parasitism levels gradually decreased from 60% to 3% ( nests) over a distance of 2 km from an agricultural edge proximal to a known cowbird foraging site (a pig feedlot), but they did not vary with distance from any other kinds of edges or with forest stand type. Rates of nest predation ( nests) did not vary with distance from any edges, but they were significantly lower in older forest than within even-aged clearcuts, a tree plantation, and in successional vegetation adjacent to a residential facility. These results suggest that, even in fragmented landscapes with high overall levels of parasitism and nest predation, management practices within and immediately adjacent to forest tracts can affect the nesting success of some species, but not necessarily as a simple function of distance from edge. For the Kentucky Warbler, our results suggest that a management strategy that avoids even-age silviculture and leaves core stands of older forest far from cowbird feeding areas can increase nesting success to levels similar to those measured in more forested landscapes.     

Nest concealment and parental behaviour interact in affecting nest survival in the blackcap (Sylvia atricapilla): an experimental evaluation of the parental compensation hypothesis

Nest concealment varies strongly within populations of many species. Although some studies have revealed the beneficial effects of concealment in mitigating predation pressure on nests, other studies were unable to find similar effects. One potential reason for the mixed results is that parental behaviour may compensate for the effects of nest cover, and specifically designed experimental studies are needed to reveal this compensation. I studied the effects of concealment on the probability of nest survival in the blackcap (Sylvia atricapilla), by experimentally manipulating the degree of nest-foliage cover. There was a significant effect of the treatment depending on nest type and the phase of nesting. Whereas there was no effect of concealment on nest survival in natural nests, there was a positive effect in real nests baited with plasticine clutches (i.e. without parental activity). Parents probably behaviourally compensated for poor concealment in natural nests (nest guarding, defence). In line with this, there was no effect of concealment on nest survival during incubation, whereas there was probably a positive effect in the nestling phase. Parents spent more time on the nest during incubation (80%) than during the care of nestlings (40%) and, consequently, had more opportunities to compensate for poor cover. In general, we cannot use single measures of behaviours or states (nest concealment) as an indication of predation risk because of the capacity for compensation in other behaviours.Communicated by C. Brown     

Individual differences in nest defense in the colonial breeding Black-tailed Gulls

Often in colonial seabirds, all colony members are believed to defend against nest predators and experience equal nest predation risk. However, the variation of defense behavior among members and its reproductive consequences are largely unknown. We investigated (1) individual variation in the nest defense of breeding Black-tailed Gulls Larus crassirostris against a natural egg predator, the Jungle Crow Corvus macrorhynchos and (2) how this behavioral variation affects an individual’s own nest predation risk and that of their neighbors. Results were compared between 2 years where crow attack levels were manipulated to average 5 and 22 times normal rates (“low” and “high” predation risk years, respectively) by the placement of varying numbers of artificial nests containing unguarded eggs at the perimeter of the gull colony. In both years, 23–38% of parents, mostly males, showed “aggressive” defense behavior (strikes or chases) against crows and decoys. Other “non-aggressive” gulls showed no defense. In the year of low predation risk, intrusion rates by crows (landing within 0.5 m of an individual gull’s nest) were similar for aggressive and non-aggressive gulls. In the year of high predation risk, however, the rates of intrusion for aggressive gulls (4%) and for non-aggressive gulls with an aggressive neighbor (37%) were significantly lower than for non-aggressive gulls without an aggressive neighbor (76%). These results indicate that aggressive individuals reduce nest predation risk for themselves and conspecific neighbors in a colonially breeding species.     

The Effect of Edge on Avian Nest Success: How Strong Is the Evidence?

Wildlife biologists historically considered the edge between adjacent habitat types highly productive and beneficial to wildlife. A current dogma is that edges adversely affect a wide range of avian species by increasing depredation and parasitism rates of nests. I critically evaluated existing empirical evidence to test whether there was a gradation in nest success as a function of distance from an edge. Researchers investigating this question have been inconsistent in their experimental designs, making generalizations about edge-effect patterns difficult. The majority of studies I examined found nest success varied near edges, with both depredation rates (10 of 14 artificial nest studies, and 4 of 7 natural nest studies) and parasitism rates (3 of 5 studies) increasing near edges. In addition, there was apositive relationship between nest success and patch size (8 of 8 studies). The most conclusive studies suggest that edge effects usually occur within 50 m of an edge, whereas studies proposing that increased depredation rates extend farther than 50 m from an edge are less convincing. Prior research has probably focused on distances too far from an edge to detect threshold values, and future research should emphasize smaller scales. 100–200 m from an edge at 20–25 m increments. Researchers often use relatively arbitrary habitat characteristics to define an edge. Therefore, I propose that only openings in the forest canopy with a diameter three times or more the height of the adjacent trees should be included in edge analyses. This review suggests that fragmentation of eastern North American temperate forests could lead to increased nest predation and parasitism, and there is need to determine if similar processes occur in other forested regions of North America.     

Assured fitness returns favor sociality in a mass-provisioning sweat bee,Megalopta genalis (Hymenoptera: Halictidae)

Assured fitness returns models for the evolution of sociality emphasize the selective value of ensuring that offspring receive adequate parental care to reach maturity. If a member of a social group dies, it can accrue returns on investment in offspring through the efforts of surviving social partners. We provide evidence that in the mass-provisioning, facultatively social sweat bee Megalopta genalis, adult presence in the nest throughout brood development provides protection from ant predation. Nests with adults present were well protected, and brood in nests with adults removed suffered higher predation. Females in observation nests showed effective defensive behavior against experimentally introduced ants, and bees in natural nests repulsed naturally occurring ant raids. Megalopta nest architecture and behavior are such that the brood of several cooperating females can be defended with little additional cost relative to solitary nesting. The benefits of cooperative defense may favor group living in mass provisioning bees. Our observations and experiments suggest that parental care throughout brood development can be adaptive in mass provisioning species, supporting the predictions of assured fitness returns models.     

Egg adoption can explain joint egg-laying in common eiders

Hypotheses regarding the evolution and maintenance of intraspecific nest parasitism were tested with data collected during a 3-year study of common eiders (Somateria mollissima) breeding near Churchill, Manitoba. The nest parasitism rate was highest (42.4% of nests) during the year with the highest nest density and the best environmental conditions, and lowest (20.2% of nests) in the year with the lowest nest density and the poorest environmental conditions. Over the nesting season, parasitic eggs were laid at the same time as normally laid eggs. Most parasitic eggs (>75%) were laid before the host female laid her third egg. The majority of the parasitic eggs were the first or second egg produced by the parasitic female. When a parasitic egg was laid before or on the same day as the host female initiated her clutch, the probability of her first egg being depredated before incubation was significantly lowered. First- and second-laid eggs suffered a high rate of predation probably because nesting females do not attend their clutch until their second or third egg is laid. Hypotheses that some females use intraspecific nest parasitism to parasitize the parental care of other females were inconsistent with these data. Egg adoption is a likely explanation for the prevalence of females incubating parasitic eggs in this population. Received: 30 September 1997 / Accepted after revision: 6 May 1998     

Costs, benefits, and plasticity of construction of nest defensive structures in paper wasps

Various animals build nests with defensive structures to deter predation on offspring. Construction of nest defensive structures can reduce the probability of predation but will involve various costs. Here, we examined both the costs and benefits of the construction of a nest defensive structure in a paper wasp, Polistes chinensis antennalis, and clarify whether the paper wasp changes the level of defensive structures of nests depending on predation risk. A foundress (queen) of the paper wasp starts a colony in spring and maintains her nest alone until the emergence of workers. At this stage, pupae in the nests are sometimes preyed on by conspecifics of other nests. The intruder needs to break the cocoon, which seals the entrance of the cell, to extract the pupa from the cell. Foundresses often apply nest material (pulp) to the surface of cocoons in their nests. We found that pulp on a cocoon increased the time an intruder required to break the cocoon. This result shows that the pulp structure on cocoons helps to prevent predation on pupae. On the other hand, pulp on cocoons involved costs, including time required to collect pulp and being a potential obstacle to the emergence of workers from the cocoon. Additionally, we found that the amount of pulp on cocoons was greater in nests under higher predation risk than nests under lower predation risk. These results suggest that pulp on cocoons is a nest defensive structure, and foundresses adjusted the construction of the defensive structure depending on predation risk.     

Offspring reproductive value and nest defense in the magpie (Pica pica)

Summary Magpie (Pica pica) brood defense against a human at the nest was studied in a Mediterranean population with low renesting potential. Variations in two defense measures recorded during 106 trials at 41 different nests were positively correlated with brood age. Ineremental effects due to the number of successive visits to nests by us, brood size, and the time in the breeding season were not significant. Partial correlation analyses showed that visit rate was not an important determinant of nest defense, which thus favors an adaptive explanation of nest defense patterns. Two functional hypotheses to account for the increase in defense intensity with brood age were tested: whether (1) increased parental defense serves to compensate the higher predation risk of older nests or (2) increased parental defense reflects the increasing reproductive value of nestlings as they grow older. Daily mortality and incidende of predation (estimated from contribution of whole-brood losses to total mortality) was higher early in the nestling period, hence providing weak evidence for the assumption on which hypothesis (1) is based. The timing of parental defense intensity did not mirror variations in predation risk for the nest but variations in reproductive value of the brood, as can be estimated from daily mortality, thus supporting hypothesis (2). Magpie parents increased defense intensity in response to premature escaping by almost fully-developed nestlings. Since such a response lowers predation risk for the offspring and increases their probability of survival, this finding supports hypothesis (2), but runs contrary to hypothesis (1). Parents also increased defense in response to play-backs of alarm calls uttered by nestlings during escaping episodes. It is argued that parents should continuously monitor the degree of offspring development in order to assess their reproductive value and that, by alarm calling, chicks honestly make their parents aware of the gain in reproductive value that results from enhancement in locomotory abilities that occur at the end of the nestling period.