Using artificial nests to explore predation by introduced predators inhabiting alpine areas in New Zealand

Abstract

Many bird species endemic to alpine New Zealand are now at critically low densities and restricted in range, making predator‐prey research difficult. We used artificial nests in the Borland Valley, Fiordland National Park, to investigate (1) which introduced species is the most frequent nest predator in the two habitats, (2) whether nest survival differs between habitats, and (3) the utility of artificial nests for guiding conservation management. We used different types of artificial nest in 2 different years and undertook a calibration study of the two types. In 2003, survival of artificial nests containing wax eggs and chicken eggs was high in both habitats. In 2004, survival of artificial nests containing plastilina eggs and chicken eggs was low in both habitats, but was higher in alpine grassland compared with beech forest. Stoats and possums were the most frequent predators (36 and 22% respectively of artificial plastilina nests in alpine grassland and high‐altitude beech forest combined); these percentages did not vary significantly between habitats. Given the low density and sparse distribution of vulnerable species in much of New Zealand, data from artificial nests can be a useful tool for studying predation in these remote and difficult habitats, or at least, preferable to ignorance. However, the type of artificial nest used can strongly affect the rate at which they are destroyed.     

An ecological paradox: More woodland predators and less artificial nest predation in landscapes colonized by noisy miners

Many passerine bird populations, particularly those that have open‐cup nests, are in decline in agricultural landscapes. Current theory suggests that an increase in habitat generalist predators in response to landscape change is partially responsible for these declines. However, empirical tests have failed to reach a consensus on how and through what mechanisms landscape change affects nest predation. We tested one hypothesis, the Additive Predation Model, with an artificial nest experiment in fragmented landscapes in southern Queensland, Australia. We employed structural equation modelling of the influence of the relative density of woodland and habitat generalist predators and landscape features at the nest, site, patch and landscape scales on the probability of nest predation. We found little support for the Additive Predation Model, with no significant influence of the density of woodland predators on the probability of nest predation, although landscape features at different spatial scales were important. Within woodlands fragmented by agriculture in eastern Australia, the presence of noisy miner colonies appears to influence ecological processes important for nest predation such that the Additive Predation Model does not hold. In the absence of colonies of the aggressive native bird, the noisy miner, the influence of woodland predators on the risk of artificial nest predation was low compared with that of habitat generalist predators. Outside noisy miner colonies, we found significant edge effects with greater predation rates for artificial nests within woodland patches located closer to the agricultural matrix. Furthermore, the density of habitat generalist predators increased with the extent of irrigated land‐use, suggesting that in the absence of noisy miner colonies, nest predation increases with land‐use intensity at the landscape scale.     

Location of bumblebee nests is predicted by counts of nest‐searching queens

1. Bumblebee nests are difficult to find in sufficient numbers for well replicated studies. Counts of nest‐searching queens in spring and early summer have been used as an indication of preferred nesting habitat, but this relationship has not yet been validated; high densities of nest‐searching queens may indicate habitat with few nesting opportunities (meaning that queens have to spend longer looking for them). 2. From mid April 2010, queen bumblebees were counted along 20 transects in grassland and woodland habitats in central Scotland, U.K. The number of inflorescences of suitable forage plants were also estimated at each transect visit. The area surrounding each transect was searched for nests in the summer. 3. In total, 173 queen bumblebees were recorded on transects, and, of these, 149 were engaged in nest‐searching. Searches subsequently revealed 33 bumblebee nests. 4. The number of nest‐searching queens on transects was significantly, positively related to the number of nests subsequently found. Estimated floral abundance along the transect did not correlate with numbers of nest‐searching queens or with the number of nests found, suggesting that queens do not target their searching to areas that are locally high in spring forage. 5. The data suggest that counts of nest‐searching queens provide a useful positive indication of good nesting habitat, and hence where bumblebee nests are likely to be found later in the year.     

Do trade‐offs between predation pressures on females versus nests drive nest‐site choice in painted turtles?

Predation strongly influences reproductive behaviours because reproducing individuals must balance mortality risks to themselves and to their offspring. In many freshwater turtles, the nest predation risk decreases with nest distance from water, whereas the predation risk to females increases farther from water. To determine whether predation pressure influences the distance from water at which female turtles nest, we measured predation pressure on nesting females and on nests, as well as the distances of nests to water, in two populations of painted turtles. Using models, we found that female survival in both populations was high and did not vary with distance from water. Nest survival was also uncorrelated with nest distance to water, although it was significantly lower than adult survival in both populations and was only 1.2% in one population. Our results suggest that nest sites are not predictably safe from predators. Instead, turtles may hedge their bets by nesting over a wide range of distances from water because any distance is risky for nests and no distance is particularly risky for the nesting female. We suggest that other factors, such as suitable incubation conditions and/or post‐emergence hatchling survival, probably play a larger role than predation in driving nest‐site choice in painted turtles.     

Different nest predator faunas and nest predation risk on ground and shrub nests at forest ecotones: an experiment and a review

This study examined predator faunas of artificial ground and shrub nests and whether nest predation risk was influenced by nest site, proximity to forest edge, and habitat structure in 38 grassland plots in south-central Sweden. There was a clear separation of predator faunas between shrub and ground nests as identified from marks in plasticine eggs. Corvids accounted for almost all predation on shrub nests whereas mammals mainly depredated ground nests. Nest predation risk was significantly greater for shrub than for ground nests at all distances (i.e. 0, 15 and 30 m) from the forest edge. However, nest predation risk was not significantly related to distance to forest edge, but significantly increased with decreasing distance to the nearest tree. Different corvid species robbed nests at different distances from the forest edge, with jays robbing nests closest to edges. We conclude that the relationship between the predation risk of grassland bird nests and distance to the forest edge mainly depends on the relative importance of different nest predator species and on the structure of the forest edge zone. A review of published articles on artificial shrub and ground nest predation in the temperate zone corroborated the results of our own study, namely that shrub nests experienced higher rates of depredation in open habitats close to the forest edge and that avian predators predominantly robbed shrub nests. Furthermore, the review results showed that predation rates on nests in general are highest <50 m inside the forest and lower in open as well as forest interior habitats (≥50 m from the edge). Received: 16 March 1998 / Accepted: 30 July 1998     

Predators of Greater Sage‐Grouse nests identified by video monitoring

ABSTRACT Nest predation is the primary cause of nest failure for Greater Sage‐Grouse (Centrocercus urophasianus), but the identity of their nest predators is often uncertain. Confirming the identity of these predators may be useful in enhancing management strategies designed to increase nest success. From 2002 to 2005, we monitored 87 Greater Sage‐Grouse nests (camera, N= 55; no camera, N= 32) in northeastern Nevada and south‐central Idaho and identified predators at 17 nests, with Common Ravens (Corvus corax) preying on eggs at 10 nests and American badgers (Taxidea taxis) at seven. Rodents were frequently observed at grouse nests, but did not prey on grouse eggs. Because sign left by ravens and badgers was often indistinguishable following nest predation, identifying nest predators based on egg removal, the presence of egg shells, or other sign was not possible. Most predation occurred when females were on nests. Active nest defense by grouse was rare and always unsuccessful. Continuous video monitoring of Sage‐Grouse nests permitted unambiguous identification of nest predators. Additional monitoring studies could help improve our understanding of the causes of Sage‐Grouse nest failure in the face of land‐use changes in the Intermountain West.     

Exploitation of underground bee nests by three sympatric consumers in Loango National Park,Gabon

Honey represents a highly nutritious resource for animals, but is difficult to obtain given bees' defensive strategies. We investigated exploitation of the underground nests of stingless bees (Meliplebeia lendliana) by three sympatric consumers in Loango National Park, Gabon: the central African chimpanzee (Pan troglodytes troglodytes), forest elephant (Loxodonta cyclotis) and honey badger (Mellivora capensis). Given the differences in their respective morphological traits and sensory abilities, we hypothesized that chimpanzees would be more limited in digging out the bee nests, compared to the other two competitors, and would show behavioral strategies to overcome such constraints. Our dataset comprised camera trap footage recorded over 60 mo at 100 different bee nests. Chimpanzees visited the nests more often than the other consumers, showing a frequency of extraction success comparable to that observed in honey badgers, the most efficient digger. Both chimpanzees and honey badgers increased their extractive attempts across the dry season, whereas elephants did not. The soil hardness was greater during the dry season than the wet season and, possibly in order to compensate for this, chimpanzees showed a tendency toward digging at nests found in relatively softer soil. They also seemed to be inhibited by indirect cues left by other consumers, possibly as a risk‐avoidance strategy. Overall, chimpanzees and honey badgers extracted the underground nests of stingless bees with similar frequencies, whilst forest elephants did so only occasionally. Moreover, chimpanzees can use tools and other behavioral strategies to overcome the physical limitations that may constrain their exploitation of this resource.     

Absence from the nest due to human disturbance induces higher nest predation risk than natural recesses in Common Eiders Somateria mollissima

Human disturbance of nesting birds may cause reduced breeding success. It is therefore necessary to assess the impact of disturbance to identify steps that minimize negative impacts. We carried out a study of nesting success at two adjacent colonies of Common Eider Somateria mollissima on the islands of Grindøya and Håkøya in northern Norway between 2006 and 2011. Over the study period, nesting success was consistently higher on Håkøya (69–82%) than on Grindøya (35–60%). Between 2009 and 2011 we used camera monitoring of individual nests to identify determinants of nest survival and predation, focusing in particular on the effect of departures from the nest due to human disturbance, which differed between the colonies due to a long‐term research project on Grindøya. Overall, absence of Common Eiders from nests due to disturbance increased the predation risk by a factor of 6.42 for an increase of one additional daily disturbance. In contrast, absence due to natural recesses did not increase nest losses. Under high levels of human disturbance, camera monitoring indicated that the main cause of breeding failure was predation, primarily by Hooded Crows Corvus cornix, but also to some extent Great Black‐backed Gulls Larus marinus. The presence of cameras did not increase the predation risk. Both the presence of researchers and the sight of Common Eider females conspicuously departing from nests are likely to have provided cues to these predators. We suggest management trials to reduce nesting disturbance through the guarding of unoccupied nests to mitigate the effects of human disturbance on reproductive success.     

Predation of experimental nests is linked to local population dynamics in a fragmented bird population

Artificial nest experiments (ANEs) are widely used to obtain proxies of natural nest predation for testing a variety of hypotheses, from those dealing with variation in life-history strategies to those assessing the effects of habitat fragmentation on the persistence of bird populations. However, their applicability to real-world scenarios has been criticized owing to the many potential biases in comparing predation rates of artificial and natural nests. Here, we aimed to test the validity of estimates of ANEs using a novel approach. We related predation rates on artificial nests to population viability analyses in a songbird metapopulation as a way of predicting the real impact of predation events on the local populations studied. Predation intensity on artificial nests was negatively related to the species' annual population growth rate in small local populations, whereas the viability of large local populations did not seem to be influenced, even by high nest predation rates. The potential of extrapolation from ANEs to real-world scenarios is discussed, as these results suggest that artificial nest predation estimates may predict demographic processes in small structured populations.     

Does land use change influence predation of bird nests?

Worldwide, many areas of agricultural land which were once covered with native vegetation have been converted to tree plantations. Such landscape transformation can influence the dynamics of wildlife populations through, for example, altering rates of predation (e.g. predation of nests of birds). Nest predation can influence reproductive success and, in turn, may alter populations by affecting juvenile recruitment. We quantified predation of bird nests in woodland remnants surrounded by two types of land use, grazing farmland and exotic Radiata pine (Pinus radiata) plantation. We also examined differences in predation rates between artificial and natural nests. We found both artificial and natural nests were more susceptible to nest predation in woodland remnants surrounded by a pine plantation than in woodland remnants located within farmland. Our study suggests that higher levels of nest predation may reduce occupancy of woodland remnants by small‐bodied birds over time, including species of conservation concern. This may have been occurred as a result of the conversion of semi‐cleared grazing land to exotic pine plantation.     

Appearance and vulnerability of artificial duck nests to avian predators

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

Predation on artificial,solitary and aggregated wader nests on farmland

Predation rates on artificial wader nests, solitary curlew (Numenius arquata) and lapwing (Vanellus vanellus) nests and lapwing nests in colonies were studied on a farmland site in central Sweden. Predation rates were highest on artificial wader nests, intermediate on solitary curlew and lapwing nests and lowest on lapwing nests in colonies, probably because of active defence of adults at real nests and/or because of selection of nest sites with lower predation risk by breeding birds. A comparison of nests close to (50 m) and far away from (200 m) forest edges revealed no increased predation risk close to edges for any of the studied nest types. Predation risk changed during the season for artificial nests (highest in the middle of May), while predation rates on lapwing and curlew nests were more stable. Artificial nests seem to be inappropriate for measuring actual predation rates and temporal differences in predation rates on real nests, but they might be suitable for use as an index of spatial differences.     

Compounding effects of habitat fragmentation and predation on bird nests

Habitat fragmentation and invasive species are two of the greatest threats to species diversity worldwide. This is particularly relevant for oceanic islands with vulnerable endemics. Here, we examine how habitat fragmentation influences nest predation by Rattus spp. on cup‐nesting birds in Samoan forests. We determined models for predicting predation rates by Rattus on artificial nests at two scales: (i) the position of the bird's nest within the landscape (e.g. proximity to mixed crop plantations, distance to forest edge); and (ii) the microhabitat in the immediate vicinity of the nest (e.g. nest height, ground cover, slope). Nest cameras showed only one mammal predator, the black rat (Rattus rattus), predating artificial nests. The optimal model predicting nest predation rates by black rats included a landscape variable, proximity to plantations and a local nest site variable, the percentage of low (<15 cm) ground cover surrounding the nest tree. Predation rates were 22 ± 13% higher for nests in forest edges near mixed crop plantations than in edges without plantations. In contrast, predation rates did not vary significantly between edge habitat where the matrix did not contain plantations, and interior forest sites (>1 km from the edge). As ground cover reduced, nest predation rates increased. Waxtags containing either coconut or peanut butter were used as a second method for assessing nest predation. The rates at which these were chewed followed patterns similar to the predation of the artificial nests. Rural development in Samoa will increase the proportion of forest edge near plantations. Our results suggest that this will increase the proportion of forest birds that experience nest predation from black rats. Further research is required to determine if rat control is needed to maintain even interior forest sites populations of predator‐sensitive bird species on South Pacific islands.     

Coastal saltpans are a good alternative breeding habitat for Kentish plover Charadrius alexandrinus when umbrella species are present

The loss and degradation of natural habitats in coastal areas worldwide has adversely affected many waterbird species, changing their breeding distribution and reducing their productivity. Anthropogenic habitats such as saltpans can provide alternative or complementary habitats for waterbirds and mitigate the increasing human impact on natural coastal habitats. Unvegetated linear paths between salt ponds are used by ground‐nesting waterbird species to breed but their linear structure may facilitate the detection of nests by predators. This negative effect may, however, be counterbalanced by the advantages of breeding in mixed colonies. To evaluate the importance and the risks of breeding in saltpans we used the Kentish plover Charadrius alexandrinus breeding in coastal saltpans of southern Portugal as a model species. Specifically, we assessed the role of nest‐site characteristics, predation and nesting proximity to species with aggressive antipredator behaviour (black‐winged stilt Himantopus himantopus) on their breeding success. Kentish plovers selected nest‐sites on the edges of paths, with about 20% of water around the nest, and a mean visibility of more than 72%; however nest‐site characteristics were not correlated with nesting success. Predation was the main cause of nest loss in saltpans (42%); carrion crows Corvus corone were responsible for most daylight nest predation (58%) and red fox (Vulpes vulpes; 73%) for night predation. An 8‐yr monitoring plan of a Kentish plover population showed a linear increase in their breeding success as their breeding season increasingly overlapped with that of the black‐winged stilt. An experiment with artificial nests showed a significant increase in the number of exposure days (7 to 12) when these nests were within close distance of black‐winged stilt nests. Overall, our results showed that saltpans are an important alternative breeding habitat for the Kentish plover, especially for the maintenance of mixed species colonies.     

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

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

Predation rate on artificial nests increases with human housing density in suburban habitats

Predation causes most nest failure in birds. Predator communities are likely to vary across a gradient of increasing urbanization, so nest predation also is likely to vary across this gradient. Although predation is thought to decline with increasing urbanization, relatively little is known about variation in predation pressure within strata along an urban gradient and how factors known to affect nest success, such as nest location, interact with urban variables, such as human housing density. Native habitats are frequently fragmented and isolated by suburban residential development, thus we quantified predation rates on artificial nests located in natural oak scrub patches within a suburban matrix in south-central Florida. We examined patterns of predation based on nest location relative to habitat edges, artificial nest weathering treatment, nest shrub height, and human housing density. Over two 18-d trials, we placed a total of 240 nests, each containing a single quail egg and a clay sham, along three roadside transects. Nest predation was not influenced by proximity to edge, nest weathering, or trial date, but was highest at high housing density and lowest at low housing density. The proportion of quail eggs removed from nests increased with human housing density. Birds were the most frequent predators of artificial nests, but the relative frequency of predation by birds or mammals did not differ relative to any of our treatments. Higher rates of nest predation with increasing human housing density within suburban habitats may reflect changes in habitat structure and composition that increase the vulnerability of nests to predation or changes in the composition of the predator community. Our results modify the conclusions of previous studies by suggesting that at scales smaller than the entire urban gradient, nest predation may increase with human housing density, one common measure of urbanization.     

Phenology largely explains taller grass at successful nests in greater sage‐grouse

Much interest lies in the identification of manageable habitat variables that affect key vital rates for species of concern. For ground‐nesting birds, vegetation surrounding the nest may play an important role in mediating nest success by providing concealment from predators. Height of grasses surrounding the nest is thought to be a driver of nest survival in greater sage‐grouse (Centrocercus urophasianus; sage‐grouse), a species that has experienced widespread population declines throughout their range. However, a growing body of the literature has found that widely used field methods can produce misleading inference on the relationship between grass height and nest success. Specifically, it has been demonstrated that measuring concealment following nest fate (failure or hatch) introduces a temporal bias whereby successful nests are measured later in the season, on average, than failed nests. This sampling bias can produce inference suggesting a positive effect of grass height on nest survival, though the relationship arises due to the confounding effect of plant phenology, not an effect on predation risk. To test the generality of this finding for sage‐grouse, we reanalyzed existing datasets comprising >800 sage‐grouse nests from three independent studies across the range where there was a positive relationship found between grass height and nest survival, including two using methods now known to be biased. Correcting for phenology produced equivocal relationships between grass height and sage‐grouse nest survival. Viewed in total, evidence for a ubiquitous biological effect of grass height on sage‐grouse nest success across time and space is lacking. In light of these findings, a reevaluation of land management guidelines emphasizing specific grass height targets to promote nest success may be merited.     

Substrate and structure of ground nests have fitness consequences for an alpine songbird

Songbird nests are an important life‐history component with multiple functions, including the creation of a suitable microclimate for offspring development. Thus, functional nest characteristics may influence fitness correlates, such as nestling size traits, and may co‐vary with prevailing environmental conditions. We investigated among‐ and within‐female variation in nest substrate, lining and decoration structures with associated fitness consequences (hatching success, nestling size traits, nest survival) across two breeding seasons for an alpine population of Horned Lark Eremophila alpestris. We combined these observations with explicit measures of nest temperature to address the influence of nest characteristics on microclimate. Nests in heather substrate had the coldest microclimates compared with grass and bare‐ground substrate, but also the greatest nest survival rates (68% versus 37–44% in other substrates), indicating the potential for substrate use decisions to reflect a trade‐off between microclimate and nest survival in response to prevailing weather and predation risk conditions. Furthermore, nest lining and nest decoration patterns indicated some support for a thermoregulatory function. Nests that were lined with willow (Salix sp.) seed‐down were associated with larger, heavier nestlings and the use of down lining decreased in frequency as the season warmed up. Nest decoration placed in front of the nest (e.g. stones or dirt clumps varying in mass from 5.3 to 186.6 g) was positively associated with warmer nest microclimates. Females demonstrated high phenotypic flexibility, as 61–94% of the observed variance in nest characteristics was explained by within‐female rather than among‐female differences. Such flexible nesting behaviour suggests the capacity to adjust to changing environmental conditions to maintain vital fitness correlates such as nest survival and nestling size development.     

Targeted agri‐environment schemes significantly improve the population size of common farmland bumblebee species

Changes in agricultural practice across Europe and North America have been associated with range contractions and local extinction of bumblebees (Bombus spp.). A number of agri‐environment schemes have been implemented to halt and reverse these declines, predominantly revolving around the provision of additional forage plants. Although it has been demonstrated that these schemes can attract substantial numbers of foraging bumblebees, it remains unclear to what extent they actually increase bumblebee populations. We used standardized transect walks and molecular techniques to compare the size of bumblebee populations between Higher Level Stewardship (HLS) farms implementing pollinator‐friendly schemes and Entry Level Stewardship (ELS) control farms. Bumblebee abundance on the transect walks was significantly higher on HLS farms than ELS farms. Molecular analysis suggested maximum foraging ranges of 566 m for Bombus hortorum, 714 m for B. lapidarius, 363 m for B. pascuorum and 799 m for B. terrestris. Substantial differences in maximum foraging range were found within bumblebee species between farm types. Accounting for foraging range differences, B. hortorum (47 vs 13 nests/km²) and B. lapidarius (45 vs 22 nests/km²) were found to nest at significantly greater densities on HLS farms than ELS farms. There were no significant differences between farm type for B. terrestris (88 vs 38 nests/km²) and B. pascuorum (32 vs 39 nests/km²). Across all bumblebee species, HLS management had a significantly positive effect on bumblebee nest density. These results show that targeted agri‐environment schemes that increase the availability of suitable forage can significantly increase the size of wild bumblebee populations.     

Predator–prey interactions between the South Polar skua Catharacta maccormicki and Antarctic tern Sterna vittata

Antarctic terns have to co‐exist in a limited space with their major nest predator, the skuas. We conducted artificial nest experiments to evaluate the roles of parental activity, nest location and nest and egg crypsis in this simple predator–prey system. Predation on artificial (inactive) nests was higher in traditional nesting sites than in sites previously not occupied by terns, which suggests that skuas memorized past tern breeding sites. Predation on artificial nests in inactive colonies was higher than in active (defended) colonies. Parental defense reduced predation in colonies to the level observed in artificial nests placed away from colonies. This suggests that communal defense can balance the costs of attracting predators to active colonies. Within colonies, predation was marginally higher on experimental eggs put in real nests than on bare ground. Although it seems that the presence of a nest is costly in terms of increased predation, reductions in nest size might be constrained by the need for protective nest structures and/or balanced by opposing selection on nest size. Predation did not differ markedly between artificial (quail) and real tern eggs. A simultaneous prey choice experiment showed that the observed predation rates reflected egg/nest detectability, rather than discrimination of egg types. In summary, nesting terns probably cannot avoid being detected, and they cannot defend their nest by attending them. Yet, by temporarily leaving the nest, they can defend it through communal predator mobbing, and at the same time, they can benefit from crypsis of unattended nest and eggs.