Exclosure fences around nests of imperiled Florida Grasshopper Sparrows reduce rates of predation by mammals

Increasing nest survival by excluding predators is a goal of many bird conservation programs. However, new exclosure projects should be carefully evaluated to assess the potential risks of disturbance. We tested the effectiveness of predator exclosure fences (hereafter, fences) for nests of critically endangered Florida Grasshopper Sparrows (Ammodramus savannarum floridanus) at a dry prairie site (Three Lakes; 2015–2018) and a pasture site (the Ranch; 2015–2016) in Osceola County, Florida, USA. We installed fences at nests an average of 8 days after the start of incubation, and nest abandonment after fence installation was rare (2 of 149 installations). Predation was the leading cause of failure for unfenced nests at both sites (48–73%). At Three Lakes, nest cameras revealed that mammals and snakes were responsible for 61.5% and 38.5% of predation events, respectively, at unfenced nests. Fences reduced the daily probability of predation (0.016 for fenced nests vs. 0.074 for unfenced nests). The probability that a fenced nest would survive from discovery to fledging was more than double that of unfenced nests (60.4% vs. 27.7%). However, we found no difference in daily nest survival at the Ranch between the year before nests were fenced (2015; 0.874) and the year when all but one nest were fenced (2016; 0.867) because red imported fire ants (Solenopsis invicta) were responsible for 86% of predation events at fenced nests at the Ranch. The use of cameras at fenced nests revealed that site‐specific differences in nest predators explained variation in fence efficiency between sites. Our fence design may be useful for other species of grassland birds, but site‐specific predator communities and species‐specific response of target bird species to fences should be assessed before installing fences at other sites.     

Building multiple nests is associated with reduced breeding performance in a south temperate population of Grass Wrens Cistothorus platensis platensis

Grass Wrens Cistothorus platensis build two types of non-breeding nest structures: platforms and dummy nests. Platforms are rudimentary accumulations of grasses concealed between vegetation. Dummy and breeding nests are dome-shaped with a similar structural layer. We used a nest-removal experiment and observational data to evaluate several hypotheses regarding the adaptive significance of building multiple nests in a south temperate population of Grass Wrens. Building non-breeding nests was not a strategy of males to attract additional females, as most of these nests were built after pair formation and both sexes collaborated during building. Building non-breeding nests was not a post-pairing display as the presence of multiple nests did not increase female investment in the breeding attempt: clutch size and female provisioning to nestlings did not differ between experimental and control territories where no non-breeding nests were removed. Similarly, in non-manipulated territories, clutch size and female provisioning were not correlated with the number of non-breeding nests or with males’ nest-building effort. Contrary to this hypothesis, the number of non-breeding nests was associated with delayed clutch initiation and reduced hatching success. The presence of non-breeding nests did not reduce nest predation and brood parasitism, which did not differ between experimental and control territories. We did not detect differences in concealment between non-breeding and breeding nests, suggesting that non-breeding nests were not the result of abandonment before egg-laying to reduce subsequent nest predation. Dummy nests did not provide shelter; they were not used frequently for roosting over the breeding season and were not maintained during the non-breeding season. We suggest that building non-breeding nests may be an attempt by males to manipulate the decision of females to breed with a mate they might otherwise reject or to start reproduction earlier than optimal for the females.     

Compounding effects of habitat fragmentation and predation on bird nests

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

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.     

Management of modern agricultural landscapes increases nest predation rates in Black‐tailed Godwits Limosa limosa

Effective conservation of endangered species requires a solid understanding of the demographic causes of population change. Bird populations breeding on agricultural grasslands have declined because their preferred habitat of herb‐rich meadows has been replaced by grassland monocultures. The timing of agricultural activities in these monocultural grasslands is critical, as they often coincide with the nesting phase of breeding birds. Here, we aim to identify the effect of habitat management and targeted nest protection on nest survival of Black‐tailed Godwits Limosa limosa in the Netherlands, a population that has shown a 70% reduction in breeding population size since the 1970s. To protect nests in monocultures from destruction, farmers are paid to either delay mowing or leave a patch of unmown grass around the nest, a patch which in practice varied in size. In herb‐rich meadows, which are typically managed for bird conservation purposes, mowing occurs after hatching. Nest survival declined as the season advanced, more steeply on monocultures than on meadows. Targeted nest protection was only partially successful, as nest predation was considerably higher on mown grassland monocultures with small unmown patches around the nest than in mown monocultures with large unmown patches and in unmown fields. Increased predator densities over the years have been suggested as an important cause of the trend towards lower nest survival, but here we show that nest survival was higher on herb‐rich meadows than on monocultures, and similar to the 1980s. It thus seems that increased predator densities are an increased threat during the egg stage only if habitat quality is low. High‐quality habitat in the form of herb‐rich meadows therefore provides a degree of protection against predators.     

The influence of landscape features on nest predation rates of grassland‐breeding waders

In Europe, lowland wet grasslands have become increasingly fragmented, and populations of waders in these fragments are subject to unsustainably high levels of nest predation. Patches of taller vegetation in these landscapes can support small mammals, which are the main source of prey for many predators. Providing such patches of habitat could potentially reduce levels of nest predation if predators preferentially target small mammals. However, predator attraction to patches of taller vegetation for foraging, shelter, perching and/or nesting could also result in local increases in predation rates, as a consequence of increased predator densities or spill‐over foraging into the surrounding area. Here we assess the influence of taller vegetation on wader nest predation rates, and the feasibility of managing vegetation structure to alter predator impacts. Between 2005 and 2011, the nest distribution and hatching success of Northern Lapwings Vanellus vanellus, which nest in the open, and Common Redshanks Tringa totanus, which conceal their nests in vegetation, were measured on a 487‐ha area of wet grassland in eastern England that is primarily managed for breeding waders. Predation rates of Lapwing nests increased significantly with distance from patches of taller vegetation, and decreased with increasing area of taller vegetation within 1 km of the nest, whereas neither variable influenced Redshank nest predation probability. These findings suggest that the distribution and activity of nest predators in lowland wet grassland landscapes may be influenced by the presence and distribution of areas of taller vegetation. For Lapwings at least, there may therefore be scope for landscape‐scale management of vegetation structure to influence levels of predation in these habitats.     

Diel patterns of predation and fledging at nests of four species of grassland songbirds

Although it is common for nestlings to exhibit a strong bias for fledging in the morning, the mechanisms underlying this behavior are not well understood. Avoiding predation risk has been proposed as a likely mechanism by a number of researchers. We used video surveillance records from studies of grassland birds nesting in North Dakota, Minnesota, and Wisconsin to determine the diel pattern of nest predation and fledging patterns of four ground‐nesting obligate grassland passerines (Grasshopper Sparrow (Ammodramus savannarum), Savannah Sparrow (Passerculus sandwichensis), Bobolink (Dolichonyx oryzivorus), and Eastern Meadowlark (Sturnella magna)). We used the nest predation pattern as a surrogate for predation activity to test whether nestlings minimized predation risk by avoiding fledging when predation activity was high and preferentially fledging when predation risk was low. Predation activity was significantly lower starting 3 hr before sunrise and ending 3 hr after sunrise, followed by a transition to a period of significantly higher activity lasting for 4 hr, before declining to an average activity level for the rest of the diel period. There was little evidence that the four grassland bird species avoided fledging during the high‐risk period and Savannah Sparrow fledged at higher rates during that period. All four species had hours during the low‐risk period where they fledged at higher rates, but only Grasshopper Sparrow fledged preferentially during that period. Bobolink and Eastern Meadowlark had multiple hours with high fledging rates throughout the daytime period, resulting in no relationship between probability of fledging and predation risk. Given the species variability in fledging pattern seen in our study, it is unlikely that there is a universal response to any driver that affects time of fledging. Further study is needed to understand the complex interplay between species ecology and drivers such as physiology, energetics, and predation in affecting grassland bird fledging behavior.     

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3.5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.     

Experimental evidence for edge-related predation in a fragmented agricultural landscape

Abstract This study tested the hypothesis that increased predation of experimental nests occurs close to a forest edge in a fragmented agricultural landscape. Artificial nests and eggs of willie wagtails Rhipidura leucophrys and superb fairy-wrens Malurus cyaneus were used in experiments to assess the extent and nature of predation occurring throughout the known breeding seasons of these species. Predators were identified by the imprints they left in plasticine eggs, and by remote photography. Surveys of avian predators were undertaken to investigate the relationship between predation intensity and predator distribution and abundance. Avian predators accounted for almost all predation for which a predator could be identified (96%). Five of seven predator species photographed attacking wagtail nests were corvids or artamids. Fairy-wren nests suffered relatively low rates of predation (29%) compared to wagtail nests (87%). Increased predation at the habitat edge was recorded for wagtail nests only; predation was correlated with the distribution and abundance of predatory avian species. The different extent and pattern of predation on fairy-wren nests could be explained by problems in detecting predation by mammals, and by possible failure of avian predators to locate the cryptic nests.     

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

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

Evidence of selection for egg crypsis in conspicuous nests

ABSTRACT The value of egg coloration as crypsis, once accepted as a general principle, has recently been questioned because most experiments have failed to show that egg coloration deters predation. The nest‐crypsis hypothesis postulates that, among species that build conspicuous nests, selection for egg crypsis is relaxed or absent because visually searching predators detect nests prior to eggs. I tested the nest‐crypsis hypothesis using the large, relatively conspicuous nests of American Robins (Turdus migratorius), and eggs that differed markedly in color that were collected from the nests of Red‐winged Blackbirds (Agelaius phoeniceus), Brewer's Blackbirds (Euphagus cyanocephalus), and Yellow‐headed Blackbirds (Xanthocephalus xanthocephalus). Each nest (N= 22) received a clutch of each species during three sequential predation trials that were 16 d in duration. The order of clutch presentation was randomized for each nest. Survival trends for Brewer's and Yellow‐headed Blackbirds were similar, and higher than those for clutches of Red‐winged Blackbirds. By the end of trials, overall survival of the three clutch types was roughly equivalent. However, clutches of Red‐winged Blackbird eggs, the most conspicuous egg type to the human eye, were discovered sooner by predators. Because the experimental design controlled for effects of nest crypsis, nest location, and nest size, this difference in egg survival can be attributed to differences in egg pigmentation. Thus, my results support a role for egg coloration as camouflage in conspicuous nests.     

Mothers vigilantly guard nests after partial brood loss: a cue of nest predation risk in a paper wasp

相似文献   

Predation on wader nests in Europe

The population declines of waders in Europe are widely considered to have resulted from habitat loss and degradation due to agricultural changes. However, recent empirical evidence suggests that levels of predation on wader nests are unsustainably high in many cases, even in some situations where breeding habitat is otherwise favourable. We review the published and 'grey' literature on nest predation on waders in Europe and quantify the relative importance of the major predators. Nest cameras offer the least biased method of identifying and quantifying nest predators. A small number of camera studies, in combination with others utilizing nest temperature loggers, indicate that nocturnal/mammalian predators make the largest contribution to wader nest predation. More than half of site-years or studies reviewed reported clutch failure rates of over 50% attributable to predation alone, a rate that is likely to be associated with declining populations, although parameters such as chick and adult survival will also affect population trends. Correlates of wader nest predation are documented, with time of season, field type and management, distance to habitat/field edge, wader nest density, and abundance of mammalian predators being most consistently identified. Future directions of research into wader productivity are discussed, and we suggest that studies quantify additional life-history parameters such as chick survival, as well as examining the predator community, wherever possible.     

Habituation and habitat changes can moderate the impacts of human disturbance on shorebird breeding performance

Disturbance by humans is widely expected to reduce the reproductive fitness of nesting birds if disturbance reduces nest attentiveness, and unattended eggs experience increased risk of predation or exposure to potentially lethal temperature extremes. Yet, relatively few studies have examined the physiological or behavioural mechanisms whereby disturbance influences reproductive fitness, or the extent to which the costs of disturbance may be reduced through habituation. We compared the behavioural responses, egg temperatures and reproductive success of shore-nesting white-fronted plovers Charadrius marginatus to disturbance at two breeding sites experiencing low versus high human recreational activity, respectively. Daytime nest attentiveness decreased with increasing experimental disturbance at both sites, but this relationship differed between sites; for any given level of disturbance, incubating birds at the more disturbed site had greater nest attentiveness. They achieved this through habituation, allowing a closer human approach before leaving the nest, and returning to the nest faster after a disturbance event. Despite lower average daytime nest attentiveness at the more disturbed site, incubation temperatures did not differ significantly between sites. Nest mortality, mostly by natural mammalian and corvid predators, was significantly lower at the site experiencing high recreational activity. However, chick mortality was significantly greater at the more disturbed site, most likely because of predation by domestic dogs. Chick mortality may have been increased by the habitation of chicks, whose escape responses were much reduced at the more disturbed site. Nonetheless, annual fecundity was substantially higher at the more disturbed site, showing that the overall reproductive fitness of wild birds is not always compromised by human disturbance and urbanization.     

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.     

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。     

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.     

Differential visual predation on morphs of Timema cristinae (Phasmatodeae:Timemidae) and its consequences for host range

Timema cristinae is a herbivorous insect that exhibits polymorphism for body coloration (green, red and grey morphs) and for pattern (striped, expressed only in the green morph, and unstriped). The striped green morph is associated with ceanothus ( Ceanothus spinosus ) and the unstriped green morph is associated with chamise ( Adenostoma fasciculatum ). This study examines the relative vulnerabilities to predation of the different pattern and colour morphs on their natural backgrounds. The vulnerabilities of the striped and unstriped morphs on their two food plants were tested using uncaged wild birds (Scrub Jays) and captive western fence lizards. Strong differential predation was observed suggesting that each morph is most cryptic on the food plant on which it is most common. Furthermore, in a mark-recapture experiment in a patch of ceanothus the unstriped and red morphs were recaptured in higher proportion than the other morphs. The vulnerabilities of the grey and green morphs on the ground and foliage were tested using lizards. The grey morph was more vulnerable on the plants than the green morph, but the inverse was observed on the ground (where they drop after a disturbance). This may be why the grey morph is not associated with specific food plants. The striped and colour polymorphisms in T. cristinae appear to be an evolutionary consequence of differential predation on different backgrounds. The implications of differential predation to food-plant utilization are discussed.     

Nest site, laying period, and breeding success of the Woodchat Shrike (Lanius senator) in Mediterranean France

Summary Breeding habitats in Mediterranean France consist in open, dry grasslands with a low and discontinuous grass layer and scattered bushes and trees. The choice of nest supports was largely opportunistic, the most abundant suitable nest support type (bush or tree) and the most abundant species being present at each study site have been used. About 75% of the nests were placed at heights between 1 and 3 m. Laying period lasted from 9 May until 6 July (peak of first clutches: 21–30 May). Mean clutch size was 5.14±0.82 eggs (n=114). Breeding success was independent from the site, the height and the concealment of nests. Mean survival rate of nests from laying to fledgling was 0.365. The main cause of low breeding success was high nest predation exerted by a rich guild of predators. The species is mainly threatened through dramatic habitat shrinkage due to pastoralism reduction leading to closures of lightly bushed grasslands in Mediterranean France.

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Nistplatzwahl, Legeperiode und Bruterfolg des Rotkopfwürgers (Lanius senator) in Südfrankreich

Zusammenfassung Als Bruthabitat bevorzugt der Rotkopfwürgers im mediterranen Südfrankreich offene und trockene Graslandlandschaften mit niedrigem oder lückigem Grasbewuchs und einzelstehenden Büschen und/oder Bäumen. Häufigster Nestträger ist die jeweils im Brutterritorium häufigste Baum- oder Buschart. Ungefähr 75% der Nester standen zwischen 1 und 3 m über Boden. Die Legeperiode reichte vom 9. Mai bis 6. Juli (Gipfel der Erstgelege am 21.–30. Mai und der Ersatzgelege am 20.–24. Juni). Die Gelegegröße bestand aus 3 bis 7, meist 5 Eier (Erstgelege im Mittel 5,35±0,71 Eier, Ersatzgelege: 4,68±0,85). Der Bruterfolg war unabhängig von der Lage, der Höhe und der Verborgenheit des Nestes. Die mittlere Überlebensrate eines Nestes zwischen Bebrütung des Geleges und Ausfliegen der Jungen betrug 0,365. Dieser niedrige Bruterfolg wurde hauptsächlich durch Nesträuber verursacht. In Südfrankreich ist die Art aber vor allem durch die zunehmende Verschließung der Bruthabitate infolge des Rückgangs des extensiven Weidebetriebs gefährdet.