Winter territoriality in migrant Black Redstarts Phoenicurus ochrurus in the Mediterranean area

The nest predation hypothesis predicts that clutch size should be inversely related to the risk of nest predation. In this paper we analyse data on nest predation, clutch size and fledgling production in 128 Jackdaw nests in order to test if clutches are larger in the safer sites. The predation rate was positively correlated with the minimum nest-entrance dimensions (predation is greater in nests with large entrances). Clutch size was not related to the risk of nest predation, and our data do not therefore support the nest predation hypothesis.     

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.     

Food use is affected by the experience of nest predation: implications for indirect predator effects on clutch size

Indirect predator effects on prey demography include any effect not attributable to direct killing and can be mediated by perceived predation risk. Though perceived predation risk clearly affects foraging, few studies have yet demonstrated that it can chronically alter food intake to an extent that affects demography. Recent studies have used stable isotopes to gauge such chronic effects. We previously reported an indirect predator effect on the size of subsequent clutches laid by song sparrows (Melospiza melodia). Females that experienced frequent experimental nest predation laid smaller clutches and were in poorer physiological condition compared to females not subject to nest predation. Every female was provided with unlimited supplemental food that had a distinctive ¹³C signature. Here, we report that frequent nest predation females had lower blood δ¹³C values, suggesting that the experience of nest predation caused them to eat less supplemental food. Females that ate less food gained less fat and were in poorer physiological condition, consistent with the effect on food use contributing to the indirect predator effect on clutch size. Tissue δ¹⁵N values corroborated that clutch size was not likely constrained by endogenous resources. Finally, we report that the process of egg production evidently affects egg δ¹³C values, and this may mask the source of nutrients to eggs. Our results indicate that perceived predation risk may impose food limitation on prey even where food is unlimited and such predator-induced food limitation ought to be added to direct killing when considering the total effect of predators on prey numbers.     

Anti-predator function of not covering eggs in the initial phase of nesting in Grey Partridge Perdix perdix: a field experiment

ABSTRACT

Capsule: An experiment in the field supports the hypothesis that Grey Partridges Perdix perdix purposely expose their first laid eggs in order to test the predation risk at their nest site.

Aims: To test the hypothesis that female Grey Partridges leave first laid eggs uncovered to assess the predation risk at their chosen nesting site.

Methods: Four area-independent experiments with artificial nests were used. Predation risk was estimated by daily nest failure rate. Generalized linear mixed-effects models were used in statistical analysis.

Results: We found that Grey Partridge females could predict nest site safety. At nest sites where the first uncovered egg was depredated, there was a higher predation risk for the whole clutch.

Conclusion: Our data statistically support the hypothesis that leaving the first egg uncovered serves to provide a more conspicuous bait for potential predators and could be a female tactic for better recognizing predation risk at a nesting site. Thus, if the first uncovered egg is depredated, the female may start a new clutch elsewhere without wasting investment in the clutch at a site under high predation risk.     

Reproductive tactics under severe egg predation: an eider’s dilemma

Parental defence against predators may increase offspring survival but entail other costs. Egg predation is frequent early in the laying sequence of the common eider, which differs in this and in several other ways from most other waterfowl. We test the hypothesis that permanent presence at the nest from the second or third egg is an adaptation for reducing egg predation in eiders. Two other alternative hypotheses for lower predation at later nest stages are early predation loss of the most vulnerable nests and seasonal decrease in predation risk. Analyses of predation rates at the one-egg and later stages refute these two alternatives. Early nest attendance by eider females is estimated to increase clutch survival by about 20% in four-egg and 35% in five-egg clutches, albeit probably at a cost of smaller clutch size.     

Parent birds assess nest predation risk and adjust their reproductive strategies

Avian life history theory has long assumed that nest predation plays a minor role in shaping reproductive strategies. Yet, this assumption remains conspicuously untested by broad experiments that alter environmental risk of nest predation, despite the fact that nest predation is a major source of reproductive failure. Here, we examined whether parents can assess experimentally reduced nest predation risk and alter their reproductive strategies. We experimentally reduced nest predation risk and show that in safer environments parents increased investment in young through increased egg size, clutch mass, and the rate they fed nestlings. Parents also increased investment in female condition by increasing the rates that males fed incubating females at the nest, and decreasing the time that females spent incubating. These results demonstrate that birds can assess nest predation risk at large and that nest predation plays a key role in the expression of avian reproductive strategies.     

人工巢箱条件下的大山雀巢捕食

于2004—2008年在次生阔叶林中,采用悬挂巢箱的方法对大山雀的巢捕食作了研究。结果表明:不同年龄巢箱的被捕食率显著不同,新巢箱被捕食率最低,第2年被捕食率最高,第3年下降很大,第4年又略有上升。被捕食巢的窝卵数极显著的低于未被捕食巢的窝卵数。影响巢捕食的主要生境因子为巢箱高度和巢上盖度,其次为灌木的密度和高度。     

Competition, predation and nest niche shifts among tropical cavity nesters: phylogeny and natural history evolution of parrots (Psittaciformes) and trogons (Trogoniformes)

Nest site selection by birds is a critically important life history trait as competition for suitable sites can be intense, and because birds are at their most vulnerable to predators during nesting. Previous studies show that the clutch size and nestling period evolve in response to competition for nest sites and nest predation, respectively. This provides the opportunity to study the relative contribution of competition and predation to the evolution of nesting niche. Using previously published phylogenies for parrots and trogons, I found evidence for at least 13 independent evolutionary transitions from tree cavities to alternative nesting niches (including termitaria, cliffs, and burrows). I analyzed variations in clutch size, incubation period and nestling period for 16 phylogenetically controlled pairs of species to test the relative roles of competition for tree cavities and nest predation, in favoring evolutionary switches to alternative nest sites. Tree cavity nesting species did not have larger clutch sizes as predicted if competition for tree cavities leads birds to invest heavily in nesting once they obtain a nest site (the limited breeding opportunities hypothesis). Instead I found that shifts to alternative nesting niches were accompanied by an increase in nestling period. As nestling period is a surrogate measure for long-term nest predation rates, this finding suggests that nest predation has been more important than competition in niche diversification among cavity nesting parrots and trogons. The timing of events in South America suggests that the explosive radiation of mammalian nest predators during the Upper-Oligocene, Lower-Miocene (20–30 million years ago) corresponded with the radiation of parrot and trogon taxa that exploit novel nesting niches.     

Life-history and ecological correlates of geographic variation in egg and clutch mass among passerine species

Broad geographic patterns in egg and clutch mass are poorly described, and potential causes of variation remain largely unexamined. We describe interspecific variation in avian egg and clutch mass within and among diverse geographic regions and explore hypotheses related to allometry, clutch size, nest predation, adult mortality, and parental care as correlates and possible explanations of variation. We studied 74 species of Passeriformes at four latitudes on three continents: the north temperate United States, tropical Venezuela, subtropical Argentina, and south temperate South Africa. Egg and clutch mass increased with adult body mass in all locations, but differed among locations for the same body mass, demonstrating that egg and clutch mass have evolved to some extent independent of body mass among regions. A major portion of egg mass variation was explained by an inverse relationship with clutch size within and among regions, as predicted by life-history theory. However, clutch size did not explain all geographic differences in egg mass; eggs were smallest in South Africa despite small clutch sizes. These small eggs might be explained by high nest predation rates in South Africa; life-history theory predicts reduced reproductive effort under high risk of offspring mortality. This prediction was supported for clutch mass, which was inversely related to nest predation but not for egg mass. Nevertheless, clutch mass variation was not fully explained by nest predation, possibly reflecting interacting effects of adult mortality. Tests of the possible effects of nest predation on egg mass were compromised by limited power and by counterposing direct and indirect effects. Finally, components of parental investment, defined as effort per offspring, might be expected to positively coevolve. Indeed, egg mass, but not clutch mass, was greater in species that shared incubation by males and females compared with species in which only females incubate eggs. However, egg and clutch mass were not related to effort of parental care as measured by incubation attentiveness. Ecological and life-history correlates of egg and clutch mass variation found here follow from theory, but possible evolutionary causes deserve further study.     

Causes of reduced clutch size in a tidal marsh endemic

We tested three hypotheses of clutch size variation in two subspecies of the swamp sparrow (Melospiza georgiana georgiana and M. g. nigrescens). Swamp sparrows follow the pattern of other estuarine endemics, where clutch size is smaller among tidal salt marsh populations (M. g. nigrescens) than their closest inland relatives (M. g. georgiana). Our results support predation risk and temperature, but not adult survival, as explanations of this pattern in swamp sparrows. Coastal nests were twice as likely to fail as inland nests, and parental activity around the nest site was positively related to clutch size at both sites. When brood size was controlled for, coastal adults visited nests less often and females vocalized less frequently during visits than inland birds, which may decrease nest detectability to predators. Coastal parents waited longer than inland birds to feed offspring in the presence of a model nest predator, but there was no difference in their response to models of predators of adults, as would be expected if coastal birds possessed increased longevity. Additionally, coastal females laid more eggs than inland females over a single season, following a within-season bet-hedging strategy rather than reducing within-season investment. Coastal territories experienced ambient air temperatures above the physiological zero of egg development more often, and higher temperatures during laying correlated with smaller clutches and increased egg inviability among coastal birds. Similar effects were not seen among inland nests, where laying temperatures were generally below physiological zero. Both subspecies showed an increase in hatching asynchrony and a decrease in apparent incubation length under high temperatures. Coastal individuals, however, showed less hatching asynchrony overall despite higher temperatures. Both air temperatures during laying and predation risk could potentially explain reduced clutch size in not only coastal plain swamp sparrows, but also other tidal marsh endemics.     

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.     

Effects of clutch sizes and incubation stage on nest desertion in the female Common Eider Somateria mollissima nesting in the high Arctic

In bird species, one of the trade-offs between reproduction and survival appears in the parental decision to desert the nest. Nest desertion is modulated by several factors including clutch size. However, the incubation stage at which predation occurs is also an important factor. In this study, we examined whether nest desertion was linked to initial clutch size, partial clutch predation (final clutch size) and the incubation stage at which it happened in a capital breeder: the female common eider (Somateria mollissima) nesting in the high Arctic. The study was performed in Kongsfjorden in 2002 on the western coast of the Svalbard Archipelago (78°55′N, 20°07′E). We observed that nest desertion was higher when the initial clutch size was small. Also, females deserted their nests more during the first third of incubation than later. Thus, as incubation proceeded, nest desertion was less likely to occur even after egg reduction. Our results pointed out that this parental decision in female eiders seemed to depend on initial clutch size and on the date into incubation of clutch reduction. Sophie Bourgeon and Francois Criscuolo contributed equally to the work     

两种山鹪莺的繁殖参数比较

生活史是鸟类生态学研究的重要内容之一,分析生活史的影响因子对于研究鸟类的生态适应具有重要意义。2007年3~9月,在广东省肇庆市江溪村对黄腹山鹪莺(Prinia flaviventris)和纯色山鹪莺(P.inornata)的繁殖参数进行了比较研究。结果表明:1)除筑巢集中期、窝卵数、巢捕食率和割草毁巢率外,两种山鹪莺各繁殖参数均存在显著性差异;2)黄腹山鹪莺的窝卵数相对较小,但卵重较大,而纯色山鹪莺则相反;3)与体重相似的9种雀形目鸟类相比,两种山鹪莺具有相对较高的年生产力;4)两种山鹪莺在部分繁殖参数上出现了分化,这可能是它们对不同巢捕食风险的响应,黄腹山鹪莺的巢捕食率相对较高,采取低窝卵数和高的卵重,而纯色山鹪莺则为高的窝卵数和低的卵重。     

Clutch size variation in passerine birds: The nest predation hypothesis

Summary The hypothesis that a negative relationship exists between clutch size and the probability that the nest will be robbed is tested, using data for passerine birds given in the literature. The data for four separate groups of species, viz. hole-nesters, semi hole-nesters and open-nesters nesting above and on the ground, respectively, were examined in relation to geographical gradients and seasonal and annual variation. In general, the data analysis results support the hypothesis, but cannot yet be considered as proven. More data on the riks of nest predation are needed. The most serious discrepancy is that for the Fieldfare, the protection from predation provided by nesting in colonies does not seem to be accompanied by a corresponding increase in clutch size. However, the clutch size of the Brambling, a species with seeks out such colonies for its breeding sites, does tend to increase in these hatitats.The pattern of clutch size variation was similar for the two groups of hole-nesting species, but differed significantly from that found for the two groups of open-nesters. The difference in the clutch size variation of the two groups of open-nesting species predicted from the hypothesis, viz. that, in northern regions, both latitudinal and altitudinal increases in clutch size should more commonly be found for those species which nest on the ground, compared to those nesting above ground level, was confirmed.A seasonal decrease in clutch size, in temperate regions, was found to be typical for species whose nests are subject to relatively little predation, particularly for those species which have a short breeding season. Advantages of laying small clutches, resulting in fewer nestlings than the number which would be possible for the parents to rear successfully, are discussed, and a simple model is presented which does not assume that nest predation is dependent on clutch size.     

Clutch desertion and re-nesting in pied flycatchers: an experiment with progressive clutch removal

Clutch desertion and re-nesting are important components of fitness when predation is frequent. In nestbox populations however, nest predation and desertion are rare but can be studied by experimental manipulations. We experimentally reduced clutches of pied flycatcher,Ficedula hypoleucaby removing one egg per day until desertion occurred. The size of the clutch at desertion and whether females re-nested or not were used as measures of the female response. Of the deserting females, 74% re-nested in our study area. Re-nesting frequency was correlated with date but not with the size of the clutch laid. The majority of the non-re-nesting females deserted empty nests, while the majority of re-nesting females deserted one egg. Clutch size at desertion was not correlated with the size of the clutch laid nor with laying date; it was smaller than the size predicted by an optimality analysis of the value of both the current (deserted) and the replacement clutch. For the re-nesting females, there was a negative correlation between fledging rate of the replacement clutch and the size of the clutch at desertion. Our predictions, made under the hypothesis that desertion and re-nesting are adaptive behaviours, were partly supported by the data; we explain the discrepancy by the constraint of searching for a new nest site or mate for re-nesting.     

Predictors of clutch predation of a globally significant avifauna in New Zealand's braided river ecosystems

Predation by introduced mammals is decimating New Zealand's indigenous fauna. Understanding factors that influence this process allows resources for predator control to be applied with maximum effect. This study examines how predation of a secondary prey species (a relatively common but declining native plover, the banded dotterel Charadrius bicinctus ) varied with reductions in abundance of a major prey source (rabbits), kill-trapping of predators, nest density and habitat complexity. Banded dotterels mostly nest in open braided riverbeds alongside a number of endemic threatened species. We measured the fate of 753 dotterel clutches exposed to predation by cats, ferrets and hedgehogs. We found key times and places of high predation risk. Immediately after widespread reduction in rabbit populations by rabbit haemorrhagic disease (RHD), clutch predation rates were almost as high (mean, 50%) as those recorded during past rabbit poisoning programmes (mean, 57%). Both rates were significantly higher than the mean predation rate of 22% without rabbit control, suggesting a shift in predator diet immediately after rabbit population declines. Unlike after rabbit poisoning, clutch predation rate remained high in the years after RHD. Other patterns observed included higher clutch predation rate where nest density was lower, suggesting that predation can potentially cause local extinction. Clutch predation was also higher along riverbed margins where vegetation was dense. There was equivocal evidence for an effect of predator kill-trapping on clutch predation rate. Management strategies that could potentially reduce clutch predation risks include focusing predator mitigation measures during periods of rabbit decline, maintaining them for more than one breeding season if the rabbit declines are widespread (e.g. RHD epidemics), and applying greater effort at sites with relatively low nest density and along riverbed margins where predator use is more frequent.     

Breeding Phenology of Birds: Mechanisms Underlying Seasonal Declines in the Risk of Nest Predation

Seasonal declines in avian clutch size are well documented, but seasonal variation in other reproductive parameters has received less attention. For example, the probability of complete brood mortality typically explains much of the variation in reproductive success and often varies seasonally, but we know little about the underlying cause of that variation. This oversight is surprising given that nest predation influences many other life-history traits and varies throughout the breeding season in many songbirds. To determine the underlying causes of observed seasonal decreases in risk of nest predation, we modeled nest predation of Dusky Flycatchers (Empidonax oberholseri) in northern California as a function of foliage phenology, energetic demand, developmental stage, conspecific nest density, food availability for nest predators, and nest predator abundance. Seasonal variation in the risk of nest predation was not associated with seasonal changes in energetic demand, conspecific nest density, or predator abundance. Instead, seasonal variation in the risk of nest predation was associated with foliage density (early, but not late, in the breeding season) and seasonal changes in food available to nest predators. Supplemental food provided to nest predators resulted in a numerical response by nest predators, increasing the risk of nest predation at nests that were near supplemental feeders. Our results suggest that seasonal changes in foliage density and factors associated with changes in food availability for nest predators are important drivers of temporal patterns in risk of avian nest predation.     

Predators and the breeding bird: behavioral and reproductive flexibility under the risk of predation

A growing body of work suggests that breeding birds have a significant capacity to assess and respond, over ecological time, to changes in the risk of predation to both themselves and their eggs or nestlings. This review investigates the nature of this flexibility in the face of predation from both behavioural and reproductive perspectives, and also explores several directions for future research. Most available work addresses different aspects of nest predation. A substantial change in breeding location is perhaps the best documented response to nest predation, but such changes are not always observed and not necessarily the best strategy. Changes in nesting microhabitat (to more concealed locations) following predation are known to occur. Surprisingly little work addresses the proactive avoidance of areas with many nest predators, but such avoidance is probably widespread. Individual birds could conceivably adopt anti‐predator strategies based on the nest predators actually present in an area, but such effects have yet to be demonstrated. In fact, the ways in which birds assess the risk of nest predation is unclear. Nest defence in birds has historically received much attention, but little is known about how it interacts with other aspects of decision‐making by parents. Other studies concentrate on predation risk to adults. Some findings suggest that risk to adults themselves influences territory location, especially relative to raptor nests. An almost completely unexplored area concerns the sorts of social protection from predators that might exist during the breeding season. Flocking typical of the non‐breeding season appears unusual while breeding, but a mated pair may sometimes act as a “flock of two”. Opportunistic heterospecific sociality may exist, with heterospecific protector species associations more prevalent than currently appreciated. The dynamics of singing during the breeding season may also respond to variation in predation risk, but empirical research on this subject is limited. Furthermore, a few theoretical and empirical studies suggest that changes in predation risk also influence the behaviour of lekking males. The major influence of predators on avian life histories is undoubtedly expressed at a broad phylogenetic scale, but several studies hint at much flexibility on an ecological time scale. Some species may forgo breeding completely if the risk of nest predation is too high, and a few studies document smaller clutch sizes in response to an increase in nest predation. Recent evidence suggests that a female may produce smaller eggs rather than smaller clutches following an increase in nest predation risk. Such an increase may also influence decisions about intraspecific brood parasitism. There are no clear examples of changes in clutch/egg size with changes in risk experienced by adults, but parental responses to predators have clear consequences for offspring fitness. Changes in risk to adults may also influence body mass changes across the breeding season, although research here is sparse. The topics highlighted herein are all in need more empirical attention, and more experimental field work whenever feasible.     

Age-specific nest-site preference and success in eiders

Variation in nest concealment is puzzling given the expected strong selection for safe nest sites. Selecting a concealed nest may decrease the risk of clutch predation but hinder parents from escaping predators, providing a possible solution to this paradox. Because the relative value of current versus future reproduction may vary with breeder age or state, nest concealment may also vary as a function of these attributes. We tested four predictions of the female and clutch safety trade-off hypothesis in eiders (Somateria mollissima): (1) nest concealment is negatively related to escape possibilities, (2) our capture rate of females is higher in covered nests, (3) egg predation is higher in open nests, and (4) overall nest success is unrelated to nest habitat. We also analysed nest microhabitat preferences and nest success relative to breeder age and body condition, controlling for nest spatial centrality. As expected, nest concealment and potential escape angle were negatively related, and capture by us, indicating female predation vulnerability, increased with nest cover. Clutch size was smaller in open nests, suggesting higher partial clutch predation, while it was larger among experienced and good-condition breeders. The probability of successful hatching was unrelated to nest habitat, positively associated with breeder experience, and negatively associated with hatching date. Experienced females selected more concealed and centrally located nests without sacrificing potential escape angles. The age-specific spatial distribution of nests on islands was unrelated to nest initiation dates, indicating no apparent competition. The age-specific preference of eiders for concealed nests may reflect declining reproductive value with age or confidence in surviving despite selecting a concealed nest. The apparently positive relationship between female age and survival and fecundity in eiders refutes the former alternative. Individual improvement in choosing safe nest sites, coupled with differential survival of individuals performing well, most likely explains age-specific nest-site preference and success.