Predator-mediated interactions between and within guilds of nesting songbirds: experimental and observational evidence

ABSTRACT Apparent competition (i.e., a mutually negative indient rect interaction between prey species through shared predation) arises when predator abundance or foraging effort increases with spetotal prey availability. We review and formalize several patch-use models from which we derive predictions for how the degree of coupling (from the predators' perspective) between nesting guilds (defined as species nesting within a vegetation stratum) affects the outcome of shared predation. We then determine which model best applies to nest predation on woodland songbirds and artificial nests by a natural population of raccoons. Using artificial nests, we showed that increasing the density of nests placed either in shrubs or on the ground increased overall predation (i.e., proportion of nests) on both types. We also tested for apparent competition between American robin and wood thrush, two coexisting woodland songbirds that commonly nest within the shrub stratum. Nest predation increased for wood thrushes but not robins as the combined density of robin and thrush nests within two individual substrate types, Lonicera and Rhamnus, increased. Thus, we documented apparent competition both within and among nesting guilds. We discuss the possible relevance of this interaction in determining species diversity, particularly in the light of increasing generalist nest predators through anthropogenically driven changes in human-altered landscapes.     

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.     

Nest height, nest concealment, and predator type predict nest predation in superb fairy-wrens (Malurus?cyaneus)

Three factors and their interaction effects are increasingly recognized as important determinants of nest predation: nest concealment, nest height, and predator type. The risk of nest predation is predicted to vary across these variables because of nest detectability and accessibility. In general, however, few studies examine how these three variables interact in relation to nest predation, focusing instead on either nest concealment or nest height (whereby predator identity is usually not known). In this study, we examine the role of nest concealment and nest height for nest survival using both artificial and natural nests in the superb fairy-wren (Malurus cyaneus). We indirectly identified potential predators through marks left on artificial eggs and footprints left on tracking tunnels. Predation level at artificial nests was lower than at natural nests, and this could be due to a failure of some nest predators to locate cryptic nests in the absence of cues provided by parental activity. Our results supported the prediction that exposed and concealed nests have different levels of nest predation, which can be explained by variation in predator type. Visual predators were only detected at exposed nests, and survival from visual predators was lower for high nests that were also exposed. However, olfactory predators were detected irrespective of nest height or nest concealment. Because rodents use olfaction to locate nests, this could explain the lack of association between nest concealment and predation outcome at low nests. In addition, rodent footmarks near nests were significantly associated with rodent tooth marks on eggs.     

Predator‐mediated interactions between preferred,alternative and incidental prey in the arctic tundra

Apparent competition between prey is hypothesized to occur more frequently in environments with low densities of preferred prey, where predators are forced to forage for multiple prey items. In the arctic tundra, numerical and functional responses of predators to preferred prey (lemmings) affect the predation pressure on alternative prey (goose eggs) and predators aggregate in areas of high alternative prey density. Therefore, we hypothesized that predation risk on incidental prey (shorebird eggs) would increase in patches of high goose nest density when lemmings were scarce. To test this hypothesis, we measured predation risk on artificial shorebird nests in quadrats varying in goose nest density on Bylot Island (Nunavut, Canada) across three summers with variable lemming abundance. Predation risk on artificial shorebird nests was positively related to goose nest density, and this relationship was strongest at low lemming abundance when predation risk increased by 600% as goose nest density increased from 0 to 12 nests ha^?1. Camera monitoring showed that activity of arctic foxes, the most important predator, increased with goose nest density. Our data support our incidental prey hypothesis; when preferred prey decrease in abundance, predator mediated apparent competition via aggregative response occurs between the alternative and incidental prey items.     

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

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

Influence of Inter-Lake Variation in Natural Nest Predation Pressure on the Parental Care Behaviour of Smallmouth Bass (Micropterus dolomieu)

Predation risk has the ability to greatly influence the behaviour of reproducing individuals. In large long-lived species with low risk of predation for parents, reproductive behaviours often involve caring for offspring (i.e. defending broods from predators) and these behaviours are essential for offspring survival. Our objectives were to test for the presence of natural variation in nest predation pressure in an aquatic environment for a species that provides sole-paternal care, smallmouth bass ( Micropterus dolomieu ), and to determine if natural variation in predation pressure influences parental care behaviour. We used snorkeler observations and a series of metrics to assess predation pressure and parental care behaviour in six lakes within a narrow geographical range. Lakes differed in all predation pressure metrics: number of predators in proximity to nest when males were present, time to predator arrival and number of predators that consumed eggs when males were absent and total number of nests that was preyed upon. Similarly, parental behaviour varied between lakes. Parental smallmouth bass spent more time engaged in anti-predator defences in lakes with high predation pressure, while males from low predator pressure lakes remained close to their nest. Conversely, males from lakes with low and high predation pressure showed a similar willingness to defend their nests during simulated nest predation events. Our results show that natural variation in aquatic nest predation pressure across multiple lakes can be significant and has the ability to influence baseline parental care behaviour. Such variation provides opportunities to study the costs and consequences of parental care and to evaluate how this could influence demography and community interactions in aquatic systems.     

Relative effects of nest size and site on the risk of predation in open nesting passerines

Large nests may incur fitness cost in terms of conspicuousness to predators, but the effect of nest size on predation risk can be confounded by effects of nest site and parental characteristics. I examined relative effects of nest size and placement by experimentally exchanging subsets of inactive nests baited with artificial clutches, among three open-cup nesting passerine species characterized by different nest size, placement and predation rate. The prediction that increasing nest size (original nest replaced by nest of the larger species) would increase predation, while decreasing nest size (replacement by nest of the smaller species) would decrease predation, relative to control (replacement by conspecific nest) was not supported in any species. The prediction that predation should be higher for large nests compared to small ones, even after exchanging nests among species-specific sites, was not supported. Predation rate differed among species (combined site/nest effect) before manipulation, whereas only the effect of nest site was significant after manipulation. This means that predation differed between species-specific nest sites, irrespective of the nest placed at these sites, but not between large (thrush) and small (warbler) nests, irrespective of their placement. Results do not suggest that nest predation selects directionally for smaller nest size. This conclusion could be specific to the study system characterized by high nest densities and high predation rate.     

Higher nest predation risk in association with a top predator: mesopredator attraction?

Breeding close to top predators is a widespread reproductive strategy. Breeding animals may gain indirect benefits if proximity to top predators results in a reduction of predation due to suppression of mesopredators. We tested if passerine birds gain protection from mesopredators by nesting within territories of a top predator, the Ural owl (Strix uralensis). We placed nest boxes for pied flycatchers (Ficedula hypoleuca) in Ural owl nest sites and in control sites (currently unoccupied by owls). The nest boxes were designed so that nest predation risk could be altered (experimentally increased) after flycatcher settlement; we considered predation rate as a proxy of mesopredator abundance. Overall, we found higher nest predation rates in treatment than in control sites. Flycatcher laying date did not differ between sites, but smaller clutches were laid in treatment sites compared to controls, suggesting a response to perceived predation risk. Relative nest predation rate varied between years, being higher in owl nest sites in 2 years but similar in another; this variation might be indirectly influenced by vole abundance. Proximity to Ural owl nests might represent a risky habitat for passerines. High predation rates within owl territories could be because small mesopredators that do not directly threaten owl nests are attracted to owl nest sites. This could be explained if some mesopredators use owl territories to gain protection from their own predators, or if top predators and mesopredators independently seek similar habitats.     

Nest predation increases with parental activity: separating nest site and parental activity effects.

Alexander Skutch hypothesized that increased parental activity can increase the risk of nest predation. We tested this hypothesis using ten open-nesting bird species in Arizona, USA. Parental activity was greater during the nestling than incubation stage because parents visited the nest frequently to feed their young during the nestling stage. However, nest predation did not generally increase with parental activity between nesting stages across the ten study species. Previous investigators have found similar results. We tested whether nest site effects might yield higher predation during incubation because the most obvious sites are depredated most rapidly. We conducted experiments using nest sites from the previous year to remove parental activity. Our results showed that nest sites have highly repeatable effects on nest predation risk; poor nest sites incurred rapid predation and caused predation rates to be greater during the incubation than nestling stage. This pattern also was exhibited in a bird species with similar (i.e. controlled) parental activity between nesting stages. Once nest site effects are taken into account, nest predation shows a strong proximate increase with parental activity during the nestling stage within and across species. Parental activity and nest sites exert antagonistic influences on current estimates of nest predation between nesting stages and both must be considered in order to understand current patterns of nest predation, which is an important source of natural selection.     

It’s a dog-eat-croc world: dingo predation on the nests of freshwater crocodiles in tropical Australia

Predation on eggs is an important source of mortality for many long-lived organisms, but causes of egg mortality from specific predators remain poorly known in most cases. Understanding the identity of predators, and the rates and determinants of their effects on a cohort of recruits, can provide a valuable background for attempts to exploit, control or conserve populations. We used remotely triggered cameras to study predation on the nests of freshwater crocodiles (Crocodylus johnstoni) inhabiting Lake Argyle, in tropical Australia. We also supplemented our work on natural crocodile nests with artificial nests. Overall, 80 of 111 natural nests were opened by predators, and predation occurred throughout the study period (7 weeks). Unlike in other parts of the species’ range, most nest-robbers were dingoes (Canis lupus dingo, responsible for 98% of all predator visits in the northern sites, and 54% in the Ord River site), with minimal additional predation by reptiles and birds. Contrary to expectation, rates of nest predation were not influenced by spatial clumping of nests: the probability of predation per nest did not change with total numbers of nests laid in an area, and artificially aggregated versus dispersed nests experienced similar levels of predation. Nest vulnerability was linked to abiotic features including slope of surrounding banks, compactness of nesting substrate, and distance from the nearest forest. Abundant aquatic food resources support a large crocodile population, but a lack of suitable nest-sites forces the crocodiles to concentrate nesting in small areas readily accessible to wide-ranging nest predators. Collectively, our results suggest that distinctive attributes of the lakeside landscape alter predator guilds and fashion unique predator–prey interactions.     

Nest Predation Deviates from Nest Predator Abundance in an Ecologically Trapped Bird

In human-modified environments, ecological traps may result from a preference for low-quality habitat where survival or reproductive success is lower than in high-quality habitat. It has often been shown that low reproductive success for birds in preferred habitat types was due to higher nest predator abundance. However, between-habitat differences in nest predation may only weakly correlate with differences in nest predator abundance. An ecological trap is at work in a farmland bird (Lanius collurio) that recently expanded its breeding habitat into open areas in plantation forests. This passerine bird shows a strong preference for forest habitat, but it has a higher nest success in farmland. We tested whether higher abundance of nest predators in the preferred habitat or, alternatively, a decoupling of nest predator abundance and nest predation explained this observed pattern of maladaptive habitat selection. More than 90% of brood failures were attributed to nest predation. Nest predator abundance was more than 50% higher in farmland, but nest predation was 17% higher in forest. Differences between nest predation on actual shrike nests and on artificial nests suggested that parent shrikes may facilitate nest disclosure for predators in forest more than they do in farmland. The level of caution by parent shrikes when visiting their nest during a simulated nest predator intrusion was the same in the two habitats, but nest concealment was considerably lower in forest, which contributes to explaining the higher nest predation in this habitat. We conclude that a decoupling of nest predator abundance and nest predation may create ecological traps in human-modified environments.     

Landscape features associated to wind farms increase mammalian predator abundance and ground-nest predation

Wind farm implementation is a rapidly growing source of landscape transformation that may alter ecological processes such as predator–prey interactions. We tested the hypothesis that wind farms increase the activity of nest predators and, ultimately, increment ground-nest predation rates. We placed 18 plots in Iberian shrub-steppes (11 at control and seven at wind farm sites), each one comprised nine artificial ground-nests (three quail eggs/nest). Artificial nests were placed during two events: at the beginning (April) and at the end (June) of the breeding season in 2016 (n?=?324 artificial nests). We estimated the relative abundance of avian and large mammalian predators in the surroundings of each plot and recorded nest fate after 12 days exposure. We also measured variables at landscape and microhabitat scale that potentially affect predator abundance and nest predation. Wind farm sites contained higher cover of gravel roads and more large mammalian predators. Moreover, the abundance of large mammalian predators increased with surrounding cover of both trees and gravel-roads. Avian predator abundance and nest predation rates did not differ between control and wind farm sites, though nest predation did increase with the surrounding cover of crops and gravel roads. Lastly, nest predation was higher at the end of the breeding season and decreased with moss and lichen cover. Our results support previous evidence on the increase of mammalian predator abundance as the surface area of gravel-roads increases, pointing towards a potential mechanism for wind farms leading to rise ground-nest predation. Future wind energy projects should minimize the development of gravel-roads for wind turbine access or maintenance.

     

Arboreal nesting as anti-predator adaptation by savanna chimpanzees (Pan troglodytes verus) in southeastern Senegal

Chimpanzees (Pan troglodytes) make nests for resting and sleeping, which is unusual for anthropoid primates but common to all great apes. Arboreal nesting has been linked to predation pressure, but few studies have tested the adaptive nature of this behavior. We collected data at two chimpanzee study sites in southeastern Senegal that differed in predator presence to test the hypothesis that elevated sleeping platforms are adaptations for predator defense. At Assirik in the Parc National du Niokolo-Koba, chimpanzees face four species of large carnivore, whereas at Fongoli, outside national park boundaries, humans have exterminated almost all natural predators. We quantified the availability of vegetation at the two sites to test the alternative hypothesis that differences in nesting reflect differences in habitat structure. We also examined possible sex differences in nesting behavior, community demographic differences, seasonality and nest age differences as variables also potentially affecting nest characteristics and nesting behavior between the two sites. Chimpanzees at Fongoli nested at lower heights and farther apart than did chimpanzees at Assirik and sometimes made nests on the ground. The absence of predators outside of the national park may account for the differences in nest characteristics at the two sites, given the similarities in habitat structure between Fongoli and Assirik. However, Fongoli chimpanzees regularly build arboreal nests for sleeping, even under minimal predation pressure, and this requires explanation.     

Effects of nest features and surrounding landscape on predation rates of artificial nests

Artificial nest predation experiments were carried out in northern Italy in woods which varied in size, isolation and surrounding landscape structure. Multivariate analyses, including logistic regression, showed that: (1) size and isolation of woods did not significantly affect predation rates; (2) nests on the edge of woods did not suffer higher predation rates than nests inside the wood; (3) nest camouflage greatly influenced predation rates, suggesting that predators were mainly using visual clues to identify nests; (4) the type of habitat that surrounded the woods emerged as a crucial factor in nest survival and the amount of human settlements in the vicinity of the wood was inversely correlated with nest survival, probably due to predators associated with humans; (5) other habitat variables, which were apparently individually unimportant, were found to have an effect on nesting success, if combined in a single ‘suitability index’. It is impossible to generalize about the influence of landscape fragmentation on nest predation because local landscape history and predator guilds, together with the scale of fragmentation, probably interact to determine the suitability of nest sites and their vulnerability to predators.     

Do artificial nests reveal relative nest predation risk for real nests?

Present knowledge of the effects of nest predation on spatial distribution, habitat selection and community structure of birds is to a large extent based on results from experiments with artificial nests. Although nest predation risk is likely to differ between artificial and real nests, most previous studies of nest predation using artificial nests have been lacking a proper control. We investigated whether predation rates on artificial nests predicted those on real nests by simultaneously comparing the fate of real and artificial nests (containing quail Coturnix coturnix and plasticine eggs) in 92 territories of the northern wheatear Oenanthe oenanthe in 1996. We also investigated whether risk for artificial nests was related to relative average risk for real nests in these territories (based on data collected two years before and two years after the experiment). Nest predation on artificial nests did predict relative predation risk for real nests only when quail egg depredation was used as the criterion for artificial nest predation. Despite plasticine egg depredation being the most common type of predation it was not associated with predation risk for real nests. Small mice and vole species dominated among cases with only plasticine egg depredation, while predatory mammals and snakes destroyed most quail eggs in artificial nests and most eggs in real wheatear nests. The results suggest that artificial nests may only predict the risk for real nests when the nest predator species are similar among the two types of nest. Furthermore, our data suggest that small mice and vole species rarely depredate nests of mid-sized passerine birds . Our results cast doubt on many previous conclusions based on experiments with artificial nests, since predation risk for such nests is likely to be uncorrelated with risk for real nests due to nest-type-specific differences in nest preying species.     

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

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

     

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

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

Nest predation in forest birds: influence of predator type and predator's habitat quality

We used the introduction of a generalist nest predator, the red squirrel Tamiasciurus hudsonicus, and of a large herbivore, the Sitka black-tailed deer Odocoileus hemionus sitkensis, to the islands of Haida Gwaii (Queen Charlotte Islands, British Columbia, Canada) to study how predator assemblage and habitat quality and structure influenced nest predation in forest birds. We compared losses of natural nests to predators on islands with and without squirrels. We selected nine islands with or without squirrel or deer and used 506 artificial nests put on the ground or in shrubs to further analyse variation of nest predation with predator assemblage and habitat quality for the predators. For both natural and artificial nests predation risk was higher in presence of squirrels. But predation risk varied within island categories. In presence of squirrels it was highest in stands with mature conifers where it fluctuated from year to year, in response to fluctuations in squirrel abundance. Vegetation cover around the nest had little effect on nest predation by squirrels. Where squirrels were absent, nest predation concentrated near predictable food sources for corvids, the main native predators, and increased with decreasing vegetation cover, suggesting that removal of the vegetation by deer increased the risk of predation by native avian nest predators that use visual cues. Predation risk in these forests therefore varies in space and time with predator composition and with quality of the habitat from the predators' perspective. This temporal and spatial variation in predation risk should promote trade-offs in the response of birds to nest predation, rather than fine-tuned adaptations to a given predation pattern.     

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.     

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

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