Sources of variation in the nesting success of understory tropical birds

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

Nest position and type affect predation rates of artificial avian nests in the tropical lowland forest on Mount Cameroon

Nest predation is the leading cause of reproductive failure in birds and thus it shapes their life history strategies. Intensities of nest predation appear to differ among nest locations and types in both temperate and tropical regions. However, there is limited knowledge of factors influencing susceptibility of avian nests to predation in Africa. The aim of our study was to investigate artificial nest predation rates of different ground and shrub nests located at different heights in the rainforest undergrowth. We placed artificial avian nests within a homogeneous lowland forest interior with sparse forest undergrowth in the Mount Cameroon National Park, Cameroon. We exposed three sets of nests: 50 bare-ground, 50 cup-ground and 50 cup-shrub nests, for 10 d. Predation was higher for cup-ground nests compared to cup-shrub nests, and bare-ground nests were more depredated than cup-ground nests. We concluded that the presence of a cup as well as higher nest position significantly increased probability of artificial nest survival. The results of this study suggest a potential selection pressure on nest type and placement in lowland forest birds for a poorly known tropical region.     

Effects of Edge Habitat and Nest Characteristics on Depredation of Artificial Nests in Fragmented Australian Tropical Rainforest

Variation in nest predation levels associated with rainforest fragmentation (edge effects) was assessed in Australia's Wet Tropics bioregion. Artificial nests were placed in the forest understorey at seven edge sites where continuous forest adjoined pasture, seven interiors (about 1 km from the edge), and six linear riparian forest remnants (50–100 m wide) that were connected to continuous forest. Four nest types were also compared, representing different combinations of two factors; height (ground, shrub) and shape (open, domed). At each site, four nests of each type, containing one quail egg and two model plasticine eggs, were interspersed about 15 m apart within a 160 m transect during September–October 2001. Predators were identified from marks on the plasticine eggs. The overall depredation rate was 66.5% of 320 nests' contents damaged over a three-day period. Large rodents, especially the rat Uromys caudimaculatus, and birds, especially the spotted catbird Ailuroedus melanotis, were the main predators. Mammals comprised 56.5% and birds 31.0% of predators, with 12.5% of unknown identity. The depredation rate did not vary among site-types, or between open and domed nests, and there were no statistically significant interactions. Nest height strongly affected depredation rates by particular types of predator; depredation rates by mammals were highest at ground nests, whereas attacks by birds were most frequent at shrub nests. These effects counterbalanced so that overall there was little net effect of nest height. Mammals accounted for 78.4% of depredated ground nests and birds for at least 47.4% of shrub nests (and possibly up to 70.1%). The main predators were species characteristic of rainforest, rather than habitat generalists, open-country or edge specialists. For birds that nest in the tropical rainforest understorey of the study region, it is unlikely that edges and linear remnants presently function as ecological population sinks due to mortality associated with increased nest predation.     

Artificial nest predation in hedgerows and scrub forest in a human-dominated landscape of central Mexico

Hedgerows as well as other narrow corridors could be valuable habitats for birds in regions of intensive agriculture, however, it is still not clear how successful breeding birds are in different types of hedgerows as compared to birds nesting in their natural habitats. We used artificial nests to examine whether hedgerows were sinks (ecological traps) for birds by comparing rates of predation in two types of hedgerows with different vegetation structure (simple and complex), and in a tract of scrub forest in an agricultural landscape of central Mexico. We determined also the types of predators responsible for egg predation. Ground and elevated nests were baited with one Japanese quail Coturnix japonica egg and one plasticine egg and placed alternately along transects. Significantly, greater predation rates were found in scrub forest and complex hedgerows than in simple hedgerows. Higher predation rates in complex habitats seemed to reflect the higher number of predator types found there. The most important predator types were carnivores followed by rodents, birds, and humans. Carnivores and rodents mainly predated ground nests, whereas birds and humans predated elevated nests. Simple hedgerows in this landscape appeared to offer relatively safe nest sites in terms of predation pressure when compared to more complex habitats (complex hedgerows and scrub forest).     

Predation on artificial nests in a forest dominated landscape – the effects of nest type, patch size and edge structure

We studied the effects of forest patch size and forest edge structure on nest predation in a boreal coniferous forest landscape. The following predictions were tested. Nest predation should be higher in small than in large stands, in edges than in interior areas of forest stands, and in barren forest/clear–cut edges created by forestry than in natural forest/open marsh edges. Four types of artificial above ground nests (total of 261) were used; open cup nests with reindeer Rangifer t. tarandus hair, open cup nests with domestic hen Gallus domesticus feathers, and unlined open cup and nest–box nests. Nests were baited with one Japanese quail Coturnix coturnix japonica egg. Nest–boxes were depredated significantly less than open cup nests of all types. No edge- or stand size–related nest predation was found. The predation rate, regardless of the nest type, did not differ relative to the edge type and vegetation characteristics. However, better horizontal visibility of open cup nests due to more open vegetation structure increased predation risk in man–made edges compared to inherent edges. The results suggest that edge–related nest predation is absent or weak in forest dominated landscapes. This may be due to predator types present in the landscape and/or predators habitat use in forest dominated areas. Therefore, it might be that findings documented in other areas, such as in agricultural dominated landscapes, cannot be directly applied to managed forest landscapes.     

Artificial nest predation rates in tropical and temperate forests: a review of the effects of edge and nest site

Nest predation rates are believed to be higher in tropical than in temperate forests. This notion is central in explaining different life history traits of tropical and temperate birds, but it is not known whether this assertion is true for all nest sites, such as ground and shrub nests, and at different distances from forest edge. I reviewed 22 studies using artificial nest experiments which concurrently contrasted predation rates of ground and shrub nests in temperate and tropical forests and found, contrary to the current dogma, no overall difference in predation rates between regions. However, there was a significant interaction between region and nest site. Ground nest predation rates were significantly higher in the tropical region, while predation rates on shrub nests were similar between regions. Within the tropical region, ground nests had significantly higher predation rates than shrub nests. Elevated nest predation rates at forest edges were found in both temperate and tropical forests. The results may have great implications for expected patterns of avian life histories and for the effects of forest fragmentation in temperate and tropical regions. First, if nest predation affects avian life histories, my results predict ground-nesting species in tropical forests to have shorter nestling periods, more broods and smaller clutch sizes than shrub-nesting species. Second, vulnerability of ground- and shrub-nesting guilds is suggested to differ between regions due to differences in forest vegetation structure, and the composition of predator faunas and their specific responses to forest fragmentation. Data to test these hypotheses are limited, but agree with the results of this review.     

Why do grebes cover their nests? Laboratory and field tests of two alternative hypotheses

Egg predation is a common feature influencing the reproductive success of open nesting birds. Evolutionary pressure therefore favours building cryptic, inconspicuous nests. However, these antipredatory pressures may be in conflict with thermoregulatory constraints, which select for dry nest material maintaining optimum temperature inside a nest cup during the absence of incubating parents. Here we examined possible trade-offs between nest crypsis and thermoregulation in Little Grebes (Tachybaptus ruficollis), which lay their eggs in floating nests built from wet plant material. As this species regularly covers its eggs with nest material, we experimentally examined (1) the rates of egg predation on covered and uncovered artificial nests and (2) possible thermoregulatory costs from nest covering by comparing temperature and relative humidity changes inside the nest cup. Results revealed that covering clutches is beneficial in terms of deterring predators, because uncovered eggs were more vulnerable to predation. Moreover, covering clutches also had thermoregulatory benefits because the mean temperature and relative humidity inside nest cups covered by dry or wet materials were significantly higher for covered compared to uncovered treatments. Covering clutches in Little Grebes therefore does not pose thermoregulatory costs.     

Nest Predation Risk on Ground and Shrub Nests in Forest Margin Areas of Sulawesi, Indonesia

Forest loss and fragmentation in Indonesia may seriously affect the survivorship of forest birds and lead to local extinction of bird populations. We used 786 artificial nests baited with quail eggs to examine the effect of habitat alteration on nest predation in Lore Lindu National Park, Sulawesi. Natural forest and four habitats of forest margin areas: forest edge, forest gardens, coffee plantations, and secondary forest, were studied. Two types of artificial nests, ground and shrub nests were placed in these habitats at two different locations for a period of 8 days. In addition, we used automatic cameras and cage-traps to identify the predators. Nests in shrubs experienced significantly higher predation rates in forest margin areas than in natural forest. Predation on ground nests did not differ significantly between these habitat types, but was significantly higher than that on shrub nests in each habitat except forest edge. Rodents were the most common predators of both nests, but shrub nests were also susceptible to Dwarf cuscus (Strigocuscus celebensis), squirrels, and tree snakes. The nest predation rates we found were among the highest found in tropical rainforests, probably a consequence of the unique predator assemblages of Sulawesi. These results suggest that egg survival is negatively affected by human intervention and that human-induced habitats might have only limited importance for the conservation of Sulawesi's largely endemic understorey avifauna. These considerations might be important since forest margins comprise significant proportions of protected areas on Sulawesi and play an important role in future Park zoning concepts as well as in conservation-oriented land use management.     

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.     

Evaluation of disruptive camouflage of avian cup‐nests

Parent birds employ various strategies to protect their offspring against nest predators. Two well‐researched anti‐nest‐predation strategies involve visual concealment of the nest by way of parental camouflage and egg camouflage. By contrast, camouflage of nest structures is relatively under‐researched, particularly in the case of cup‐nests in trees and bushes. We explored how birds camouflage cup‐nests in nature. Specifically, we tested Hansell’s hypothesis that birds use externally applied pale and white objects such as spider cocoons and lichens to achieve cup‐nest camouflage. To test Hansell’s hypothesis, three complementary experiments were performed: (1) an in situ nest predation experiment; (2) a photo‐based visual search experiment; and (3) contrast analyses using PAT‐GEOM software in IMAGEJ. White paper and chalk spots were used to mimic white objects used by birds in nature. Whereas predation rates in Experiment 1 were not affected by white spots, location rates in Experiment 2 were lower for natural nests with white spots than without white spots. Experiment 3 demonstrated that white spots significantly increased the contrast between different visual elements of nests. It was concluded that white objects can potentially camouflage nests against some nest predators, and that any improved camouflage was probably achieved via disruptive camouflage.     

千岛湖栖息地片段化效应对鸟类巢捕食风险的影响

为了解栖息地片段化对鸟类巢捕食风险的影响,我们于2010年4-8月,在千岛湖地区选取16个岛屿,分别在岛屿边缘和内部区域用放置鸡蛋(大型卵)和鹌鹑(Coturnix japonica)蛋(小型卵)的方法进行人工模拟地面巢试验,研究不同体积大小鸟卵的巢捕食率及其差异,分析巢捕食率与岛屿面积、隔离度、形状指数和植物物种丰富...     

Impact of brood parasitism and predation on nest survival of the fan-tailed gerygone in New Caledonia

Predation and brood parasitism are common reasons for nesting failure in passerine species and the additive impact by invasive species is a major conservation concern, particularly on tropical islands. Recognising the relative contribution of the different components of nesting failure rates is important to understand co-evolutionary interactions within brood parasite–host systems. In the remote archipelago of New Caledonia, the fan-tailed gerygone Gerygone flavolateralis is the exclusive host of the brood-parasitic shining bronze-cuckoo Chalcites lucidus. Additionally, invasive rodents also possibly have an impact on breeding success. To estimate the impact of potential nest predators, we 1) video monitored nests to identify predators, 2) estimated the probability of predation based on nest visibility and predator abundance and 3) tested the possibility that the location of experimental nests and lack of odour cues decrease the predation by rodents. In addition, we estimated nest survival rates using data collected in different habitats over the course of eight breeding seasons. Nesting success of fan-tailed gerygones was relatively low and predation was the main cause of nesting failure. We recorded mainly predation by native birds, including the shining bronze-cuckoo, whereas predation by rats was rare. In open habitats predation by cuckoos was much lower than predation by other avian predators. Neither predator activity around nests nor nest visibility influenced the probability of predation. Experimental nests in more accessible locations and containing an odorous bait were more exposed to rodent predation. Apparently, the fan-tailed gerygone has either never been specifically vulnerable to predation by rats or has developed anti-predator adaptations.     

Habitat-specific clutch size and cost of incubation in eiders reconsidered

The energetic incubation constraint hypothesis (EICH) for clutch size states that birds breeding in poor habitat may free up resources for future reproduction by laying a smaller clutch. The eider (Somateria mollissima) is considered a candidate for supporting this hypothesis. Clutch size is smaller in exposed nests, presumably because of faster heat loss and higher incubation cost, and, hence, smaller optimal clutch size. However, an alternative explanation is partial predation: the first egg(s) are left unattended and vulnerable to predation, which may disproportionately affect exposed nests, so clutch size may be underestimated. We experimentally investigated whether predation on first-laid eggs in eiders depends on nest cover. We then re-evaluated how nesting habitat affects clutch size and incubation costs based on long-term data, accounting for confounding effects between habitat and individual quality. We also experimentally assessed adult survival costs of nesting in sheltered nests. The risk of egg predation in experimental nests decreased with cover. Confounding between individual and habitat quality is unlikely, as clutch size was also smaller in open nests within individuals, and early and late breeders had similar nest cover characteristics. A trade-off between clutch and female safety may explain nest cover variation, as the risk of female capture by us, mimicking predation on adults, increased with nest cover. Nest habitat had no effect on female hatching weight or weight loss, while lower temperature during incubation had an unanticipated positive relationship with hatching weight. There were no indications of elevated costs of incubating larger clutches, while clutch size and colony size were positively correlated, a pattern not predicted by the ‘energetic incubation constraint’ hypothesis. Differential partial clutch predation thus offers the more parsimonious explanation for clutch size variation among habitats in eiders, highlighting the need for caution when analysing fecundity and associated life-history parameters when habitat-specific rates of clutch predation occur.     

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     

An analysis of clutch-size in New World passerine birds

Variation in clutch-size among New World passerine birds was analysed with respect to four variables: body-mass, geographic latitude, the frequency of nest predation, and the structure of the nest. Data were analysed separately by averaging traits at three taxonomic levels: species, genus, and subfamily. Allometric scaling with body-mass did not account for significant variation in clutch-size regardless of the taxonomic level of analysis. Latitudinal effects on clutch-size were highly significant at all taxonomic levels. For species building small-pensile nests and open-cup nests, nest predation had a significant partial effect on clutch-size, with latitude held constant. When nest predation and latitude were held constant, clutch-size was significantly different among species building small-pensile nests, open-cup nests, and domed nests. These results suggest that New World passerine clutch-size is related to at least three variables: latitudinal effects, nest predation, and nest structure.     

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

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

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.     

Long‐term and large‐scale analyses of nest predation patterns in Australian songbirds and a global comparison of nest predation rates

Juvenile mortality is one of crucial drivers of life‐history evolution, and predation is the main cause of nest loss in birds. Thus, understanding how nest predation and failure vary in nature is important for understanding life history evolution and, moreover, for effective conservation. We used published data and unpublished records to study factors influencing nest predation and total failure in 138 populations of 90 species of Australian songbirds. Daily predation (average 2.0% d^?1) and failure rates (2.9%) increased from temperate regions to the tropics, over the last four decades, and were lowest in temperate south‐western Australia. Predation and failure were higher in smaller species, and failure rates were lower in species with closed nests than in species with open nests. There was no effect of nest height or nest site (ground, shrub, canopy) or social organization on nest predation or failure rates. Nest predation caused on average 72% of total nest failure, similar to other tropical, subtropical, and temperate areas. Our study spanning from the tropics to temperate regions and using > 10 000 nests confirmed that tropical birds faced higher nest failure rates. We identified an increase in nest depredation rates in the last four decades in Australia, suggesting that a large‐scale ecological phenomenon must be responsible. It may include increases in predator abundances and/or ranges, possibly connected with human‐caused habitat change. A global comparison of nest failure rates confirmed that predation is the main source of nest mortality in songbirds worldwide. We discuss implications of our results for the evolution of reproductive strategies and for the conservation of Australian birds.     

Support for the nest mimicry hypothesis in Yellow-rumped Thornbills Acanthiza chrysorrhoa

The nest of the Yellow-rumped Thornbill Acanthiza chrysorrhoa (endemic to Australia) consists of a lower domed nest where the young are cared for and an upper cup-shaped structure, referred to as the false nest. Numerous hypotheses have been proposed to explain the primary function of the false nest, but their predictions remain largely untested. Here, we test predictions of the nest mimicry hypothesis (with artificial nests) whereby the empty false nest acts as a decoy, deflecting visual nest predators and/or brood parasites away from the true nest below, thereby increasing nesting success. Clear detection of the false nest by adversaries is an intrinsic aspect of nest mimicry; thus we also test whether conspicuous nests (natural) are favoured by natural selection. Specifically, (1) we compare predation level at artificial domed nests with and without a false nest; and (2) we analyse associations between nest site concealment and nesting outcome in natural nests in a wild population. Supporting the nest mimicry hypothesis, predation level was significantly lower for artificial domed nests with false nests, and for natural nests with lower levels of concealment.     

Effects of nest-site characteristics and parental activity on cowbird parasitism and nest predation in Brown-and-yellow Marshbirds

ABSTRACT Nest‐site selection and nest defense are strategies for reducing the costs of brood parasitism and nest predation, two selective forces that can influence avian nesting success and fitness. During 2001–2002, we analyzed the effect of nest‐site characteristics, nesting pattern, and parental activity on nest predation and brood parasitism by cowbirds (Molothrus spp.) in a population of Brown‐and‐yellow Marshbirds (Pseudoleistes virescens) in the Buenos Aires province, Argentina. We examined the possible effects of nest detectability, nest accessibility, and nest defense on rates of parasitism and nest predation. We also compared rates of parasitism and nest predation and nest survival time of marshbird nests during the egg stage (active nests) with those of the same nests artificially baited with passerine eggs after young fledged or nests failed (experimental nests). Most nests (45 of 48, or 94%) found during the building or laying stages were parasitized, and 79% suffered at least one egg‐predation event. Cowbirds were responsible for most egg predation, with 82 of 107 (77%) egg‐predation events corresponding to eggs punctured by cowbirds. Nests built in thistles had higher rates of parasitism and egg predation than nests in other plant, probably because cowbirds were most active in the area where thistles were almost the only available nesting substrate. Parasitism rates also tended to increase as the distance to conspecific nests increased, possibly due to cooperative mobbing and parental defense by marshbirds. The proportion of nests discovered by cowbirds was higher for active (95%) than for experimental (29%) nests, suggesting that cowbirds used host parental activity to locate nests. Despite active nest defense, parental activity did not affect either predation rates or nest‐survival time. Thus, although nest defense by Brown‐and‐yellow Marshbirds appears to be based on cooperative group defense, such behavior did not reduce the impact of brood parasites and predators.