The influence of territory characteristics and food supply on the breeding performance of the Red-backed Shrike (<Emphasis Type="Italic">Lanius collurio</Emphasis>) in an extensively farmed region of eastern Poland

The breeding performance of the Red-backed Shrike (Lanius collurio) in relation to the territory characteristics was investigated. The study was conducted in an area of high agricultural land use, characterised by small field size, a more mosaic habitat and low use of mineral fertilisers in comparison to Western European countries. The influence of habitat structure and composition in the territory on the date of clutch initiation, nest predation and clutch size in the Red-backed Shrike was not found, but such an influence on the number of nestlings was shown. The minimum adequate model explained 18.3% of the variation between territories in nestling’s number. The number of fledglings was positively correlated with the area of pastures, meadows, orchards and fallows within territories, and negatively correlated with length of overhead transmission lines. The number of fledglings was correlated with food abundance within territories, based on to the results from pitfall traps. There was no correlation between territory food abundance and the size of clutch. The presence of pastures, meadows and fallows, which are foraging places, thus seems to be crucial for the protection of the Red-backed Shrike.     

Between land abandonment and agricultural intensification: habitat preferences of Red-backed Shrikes Lanius collurio in low-intensity farming conditions

Capsule The conservation of Red-backed Shrikes on farmland habitats depends on extensive farming conditions.

Aims To evaluate breeding density and habitat preferences of the declining Red-backed Shrike and relate its occurrence to an environmental gradient ranging from land abandonment to intensive farming.

Methods The study was carried out in the Apennines (northern Italy), at nine 21-ha study plots. We identified factors affecting breeding density and habitat preferences at two spatial scales (landscape and territory) and analysed the variation in territory density according to the relative farming intensity in a low-intensity agricultural landscape.

Results The presence of shrubs and cultivated/grazed land positively influenced the number of Red-backed Shrike territories per plot, while the species' settlement within plots was related to higher values of shrub cover and the presence of hedges. Shrike occurrence was associated with land-use categories intermediate between land abandonment and intensive agriculture.

Conclusions This study provides a first detailed assessment of Red-backed Shrike habitat requirements in southern Europe. Favoured habitats were pasture/cultivation mosaics flanked by or interspersed with shrubs/hedges (15–20% of the surface of the 1-ha medium-sized territory). Thus, the conservation of Red-backed Shrikes in low-intensity southern European farmland appears to reflect a trade-off between agricultural intensification and long-term land abandonment.     

An ‘ecological trap’ for yellow warbler nest microhabitat selection

Contrary to assumptions of habitat selection theory, field studies frequently detect ‘ecological traps’, where animals prefer habitats conferring lower fitness than available alternatives. Evidence for traps includes cases where birds prefer breeding habitats associated with relatively high nest predation rates despite the importance of nest survival to avian fitness. Because birds select breeding habitat at multiple spatial scales, the processes underlying traps for birds are likely scale‐dependent. We studied a potential ecological trap for a population of yellow warblers Dendroica petechia while paying specific attention to spatial scale. We quantified nest microhabitat preference by comparing nest‐ versus random‐site microhabitat structure and related preferred microhabitat features with nest survival. Over a nine‐year study period and three study sites, we found a consistently negative relationship between preferred microhabitat patches and nest survival rates. Data from experimental nests described a similar relationship, corroborating the apparent positive relationship between preferred microhabitat and nest predation. As do other songbirds, yellow warblers select breeding habitat in at least two steps at two spatial scales; (1) they select territories at a coarser spatial scale and (2) nest microhabitats at a finer scale from within individual territories. By comparing nest versus random sites within territories, we showed that maladaptive nest microhabitat preferences arose during within‐territory nest site selection (step 2). Furthermore, nest predation rates varied at a fine enough scale to provide individual yellow warblers with lower‐predation alternatives to preferred microhabitats. Given these results, tradeoffs between nest survival and other fitness components are unlikely since fitness components other than nest survival are probably more relevant to territory‐scale habitat selection. Instead, exchanges of individuals among populations facing different predation regimes, the recent proliferation of the parasitic brown‐headed cowbird Molothrus ater, and/or anthropogenic changes to riparian vegetation structure are more likely explanations.     

Protective nesting association between the Barred Warbler Sylvia nisoria and the Red-backed Shrike Lanius collurio: an experiment using artificial and natural nests

The pressure of predators may significantly affects the distribution pattern of nesting birds. Some individuals may reduce the risk of predation by nesting near other species with an aggressive nest defence. In the present study I tested the predator protection hypothesis using experimental (artificial nests) and observational (real nests) approaches on two ecologically similar passerine birds–the Barred Warbler Sylvia nisoria and the Red-backed Shrike Lanius collurio. Studies have been conducted in eastern Poland in two types of habitat: river valley and farmland. The main predators of natural and artificial nests were birds, and to a lesser extent, also mammals. I found wide variation level of predation of both types of nests in different years. Nest survival rate of artificial nests was significantly lower in the farmland than in the river valley and in natural nests I observed reverse pattern. According to the predictions of the predator protection hypothesis the survival rates of the natural and artificial nests were significantly higher in territories of individuals breeding in the protective nesting association. This type of interspecific positive interaction between two associate species can be classified as facultative mutualism.     

The reproductive success of Blackbirds Turdus merula in relation to habitat structure and choice of nest site

The reproductive success of a population of Blackbirds Turdus merula occupying farmland and woodland was studied over 3 years to investigate the effects of habitat on breeding success. Territory distribution was patchy in both farmland and woodland; some areas were unoccupied, while other areas were occupied at variable densities. Habitat structure appeared to influence occupation: the index of habitat complexity ("cover score") was higher in occupied areas than in unoccupied areas and high-density territories had higher cover scores than low-density territories. However, habitat structure had no significant effect on reproductive success because the cover scores of territories where pairs were successful did not differ significantly from those of territories where there were no successful breeding attempts. There was no evidence of differential mortality rates in adults according to habitat. The height, bulk and exposure of c. 430 nests were measured to determine the effect of nest and nest-site characteristics on reproductive success. Nest exposure was the only feature that differed between successful and failed nests, successful nests being less exposed than failed nests. The major cause of breeding failure was nest predation, but the effect of nest exposure operated only during the laying and incubation period and not during the nestling period. The significance of habitat structure for variation in population densities between habitats is discussed.     

Nesting success and within-season breeding dispersal in the Orange-breasted Sunbird Anthobaphes violacea

Nest predation is a primary cause of nesting mortality for many bird species, particularly passerines. Nest location can affect predation, and it has also been demonstrated that predation risk can alter nest site selection. Birds can limit predation risk by selecting specific habitat characteristics; by changing nest site characteristics between attempts; and by dispersing between nesting events. Here we report breeding data from a population of Orange-breasted Sunbirds Anthobaphes violacea (L.), for a single breeding season in the Jonkershoek Nature Reserve, South Africa. Neither shrub type nor nest height was found to affect the outcome of a nest. For subsequent breeding attempts, birds were not more likely to change the type of shrub in which they nested after a predation event than when the attempt was successful, nor did they change the height of their nest. However, we found that the distance between a nest and the subsequent one was significantly shorter after a successful nest than after an unsuccessful one. We interpret this as an adaptive strategy to avoid predation.     

Reproductive success of Tree Pipits Anthus trivialis in relation to habitat selection in conifer plantations

Aspects of the reproductive success of Tree Pipits Anthus trivialis were examined in relation to broad‐scale habitat and nest‐site selection in Thetford Forest, a coniferous plantation forest in eastern England. Three habitat classes were defined corresponding to previously reported densities of Tree Pipits: clearfell and recently planted stands (habitat class A: low density), stands 2–5 years old (B: high density) and stands 6 years or older (C: low density). The preference for 2–5‐year‐old stands indicated by higher densities was supported by the timing of territory settlement. Tree Pipits also showed distinct preferences for nest‐site characteristics that were relatively consistent across habitat classes and throughout the breeding season. At the ‘habitat scale’, results were consistent with the predictions of the ideal despotic distribution model. First clutches were laid significantly earlier in the preferred habitat class B. Overall nesting success (i.e. the proportion of nests producing fledglings), but not clutch size, also varied between habitats, being greater in habitat classes B and C than in habitat class A. The variation in overall nesting success between habitats was primarily driven by low nest survival rates during the laying/incubation period in clearfell and recently planted stands. Nest survival rates during the nestling period were lower in the preferred 2–5‐year‐old (and older) stands and declined over the course of the study. Preferences for nest‐site characteristics (at least for those that were measured) provided no apparent benefit to nest survival rates. Overall nesting success thus appeared to be determined at the habitat scale, perhaps because the broad differences in cover between habitats affected the likelihood of nest predation (the main cause of nest failure). It is suggested that the very low nesting success experienced by Tree Pipits in clearfell and new stands may be one factor in the species’ relative avoidance of this habitat and preference for 2–5‐year‐old stands.     

Habitat selection responses of parents to offspring predation risk: an experimental test

The ability of nest predation to influence habitat settlement decisions in birds is widely debated, despite its importance in limiting fitness. Here, we experimentally manipulated nest predation risk across a landscape and asked the question, do migratory birds assess and respond to variation in nest predation risk when choosing breeding habitats? We examined habitat preference by quantifying the density and settlement date of eight species of migratory passerines breeding in areas with and without intact nest predator communities. We found consistently more individuals nesting in areas with reduced nest predation than in areas with intact predator assemblages, although predation risk had no influence on settlement or breeding phenology. Additionally, those individuals occupying safer nesting habitats exhibited increased singing activity. These findings support a causal relationship between habitat choice and nest predation risk and suggest the importance of nest predation risk in shaping avian community structure and breeding activity.     

Habitat selection and polygyny in breeding Corn Buntings Miliaria calandra

Unlike many other polygynous passerine species, female Corn Buntings Miliaria calandra apparently do not suffer costs by pairing polygynously, yet it is unclear whether this is because polygynous males hold the highest quality territories or because pairing with polygynous males is unimportant in determining female reproductive success. Male Corn Buntings on North Uist, Scotland, consistently defended territories which contained nesting habitat, and females often foraged outside male territories when provisioning nestlings. Females showed strong preferences for nesting in uncultivated land, and 80% of nests were under Hogweed Heracleum sphondylium, possibly because this provided cover against predation and the weather. When provisioning nestlings, females showed strong preferences for foraging in cereal crops, probably because this habitat provided better food resources and/or better cover from predators. Males were unpaired or paired with one to three females per breeding season, but variation in territory size or vegetation composition did not explain differences in the number of females paired with individual males. We suggest that when females neither gain benefits nor suffer costs by breeding polygynously, and males do not differ greatly in the areas of habitat selected, polygyny can arise through random female settlement within the nesting habitat.     

Habitat selection by breeding Curlews Nwnenius arquata on mosaic farmland

Territory establishment and habitat use by breeding Curlews Numenius arquata were studied during 1987 and 1988 on mosaic farmland (dominated by dry tillage) at two sites in central Sweden. Curlews preferred to breed in areas with a high proportion of grassland, close to rivers, while dry tillage was avoided. Territories at my study site were larger (mean = 4 5.2 ha) than in areas consisting entirely of grassland. Territory size seemed to depend on the spatial distribution of grasslands, which suggests that habitat fragmentation forces Curlews to establish larger territories in modern farmland than in areas of grassland. The number of territories in patches of grassland was correlated with patch area, and unoccupied patches were more isolated than occupied patches. However, patch area was a more important factor than isolation, since large patches (> 3 5 ha) were always occupied. Sown grassland was used significantly more than expected for foraging early in the season, possibly indicating the strong influence of the nutritional requirements in the pre-breeding period on territory establishment. Habitat selection when foraging seemed to be less important late in the season, since there was no significant habitat preference then. During this period distance to the nest site seemed to be more important than habitat, sinced the preferred foraging fields (including fields of all habitats used more than expected by area) were situated closer to nests than the less preferred fields, probably an adaptation to the high nest predation risk. The same fields were mostly preferred in the pre-breeding period also, suggesting that nests were built close to good foraging areas.
My results indicate that the decline of the Swedish Curlew population since 1950 is caused by changes in land use, resulting in decreased grassland area and increased habitat fragmentation, which probably have affected both breeding and foraging possibilities negatively.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

Avoid nest predation when predation rates are low,and other lessons: testing the tropical–temperate nest predation paradigm

Nest predation is the most important cause of nest failure in most birds and latitudinal differences in nest predation rates and life histories suggest that nest predation has been influential in life history evolution. All else equal, natural selection should favor reduction of nest predation, yet evidence is equivocal. We used Monte Carlo simulations to examine the combined effects of variation in nest predation rates, breeding season length and renesting intervals on the annual number of young fledged. Simulations suggest that selection most strongly favors a reduction in nest predation when breeding seasons are short and predation rates are low (temperate characteristics). Conversely, selection favors shorter renesting intervals when breeding seasons are long and nest predation rates are high (tropical characteristics). Reducing already low rates provides a proportionately greater increase in annual nesting success than does the same reduction when nest predation rates are higher. In some tropical species, individuals increase reproductive success not by avoiding predation in subsequent nesting attempts, which is largely beyond their control, but rather by reducing renesting intervals. We suggest that the emphasis on nest predation avoidance has biased our perspectives for alternative hypotheses of how birds should respond to nest predation and the consequences of those alternatives for life history theory. Similarly to the need to control for phylogenetics in examining life history strategies, future studies must also control for differences in breeding season lengths and renesting intervals to better understand the influence of nest predation on avian life histories.     

Predation risk determines breeding territory choice in a Mediterranean cavity-nesting bird community

Non-direct effects of predation can be an important component of the total effect of predation, modulating animal population and community dynamics. The isolated effects of predation risk on the spatial organisation of the breeding bird community, however, remains poorly studied. We investigated whether an experimentally increased predation risk prior to reproduction affected breeding territory selection and subsequent reproductive strategies in three Mediterranean cavity-nesting birds, i.e., the little owl Athene noctua, European roller Coracias garrulus and scops owl Otus scops. We found that territories used the previous year were more likely to be re-occupied when they belonged to the safe treatment rather than to the risky treatment. The first choice of breeders of all three species was for safe territories over risky ones. When all breeding attempts in the season (i.e., final occupation) were considered, breeders also preferred safe to risky sites. In addition, little owls laid larger eggs in risky territories than in safe territories. Our study provides experimental evidence of a rapid preventive response of the three most abundant species in a cavity-nesting bird community to a short-term manipulation of predation risk. This response highlights the key role of the non-direct effects of predation in modulating avian community organisation.     

Nesting in an uncertain world: information and sampling the future

Seasonal fecundity is strongly influenced by the number of nests attempted (including renests following nest failure) in a season. This number is often assumed to be set by the length of the breeding season or through some predetermined maximum. Instead, the decision to renest likely results from a cost–benefit analysis honed by natural selection where the ultimate components of fitness, i.e. reproduction and survival, tradeoff with one another. Moreover, reproductive decision making should not occur in an information‐vacuum. In a world where habitat quality (i.e. likelihood of nest success) is uncertain females should use nest failures to update the probability their renest nest will succeed, and this estimate in turn can affect the decision to renest. We develop a model of renesting behavior based on these conceptual ideas using the framework of statistical decision theory (SDT) and the process of Bayesian updating. We assume that renesting incurs 1) a cost to future reproductive success and 2) a reduction in expected reproductive success for each successive replacement nest. We show, all else equal, birds should curtail renesting with (1) increased residual reproductive success, (2) increased cost to future reproduction, (3) declines in current reproduction with successive attempts, and (4) increasing nest predation rates. We also explore several ecological implications of the model. First, uncertainty in habitat quality and the process of information updating, based on nest outcomes, link changes in the quality or proportion of one habitat type to the behavior in the other habitat. Second, females may use their estimate of habitat quality, based on a sample of nesting attempts, to decide whether to return or disperse to a new site between years. We compare this to the win‐stay: lose‐switch rule for dispersal and discuss the implications for population dynamics.     

Do parasitic common goldeneye Bucephala clangula females choose nests on the basis of host traits or nest site traits?

Conspecific brood parasitism (CBP) is an important alternative breeding strategy for gaining reproductive output in birds. While interactions between hosts and parasites and consequences of CBP to breeding success of both parties have been studied a lot, the roles of host characteristics and nest site characteristics in CBP have received less attention. We studied the relative importance of host‐related traits, such as female condition and breeding experience, and nest‐site‐related factors, such as overall nest site preference and occupation rate, in explaining the occurrence of CBP in a common goldeneye Bucephala clangula population. We used spatially and temporally extensive data sets, analysed the data with generalized linear mixed models that allowed us to account for the non‐independency of individual nesting attempts across females and nesting sites, and used an information theoretic approach in model selection and inference. About half of the nests were parasitized annually during the seven year study period. The occurrence of CBP decreased with advancement of the breeding season but late nests were also frequently parasitized. We found that the occurrence of CBP was better explained by nest‐site characteristics than host traits, implying that parasitic females target a given nest based on factors related to the nest site itself rather than on the host. Our results suggest that more attention should be paid to factors associated with nest site attractiveness and quality when studying laying decisions of parasites and the occurrence of CBP in general.     

Experimental evidence of environmental effects on age-specific reproductive success: the importance of resource quality.

Age-specific access to high-quality resources (e.g. territory or nest site) might be an important determinant for improved reproductive performance with increasing age. I experimentally investigated the effects of territory quality versus other age-related improvements in breeding competence (e.g. foraging skills, breeding experience and local knowledge) on age-specific reproductive success. Territory quality (i.e. territory field layer height) was manipulated in year 2 of northern wheatears (Oenanthe oenanthe) that were breeding in the same territory in two consecutive years. Changing territory quality by changing field layer height had a strong effect on within-individual change in the reproductive success of wheatears. This effect was mainly due to a corresponding change in nest predation risk. When territory quality was kept constant (i.e. no between-year change in territory field layer height), within-individual reproductive success did not change between subsequent years. Thus, age-related improvements in foraging skills, breeding experience and local familiarity had no significant effect on within-individual changes in reproductive success. Increased reproductive success with increased age in northern wheatears is therefore mainly explained by an improved access to high-quality territories with increasing age. I conclude that age-dependent access to high-quality breeding resources might be a widespread phenomenon in nature.     

Group composition and reproductive success of cooperatively breeding white‐winged choughs (Corcorax melanorhamphos) in urban and non‐urban habitat

Abstract: White‐winged choughs (Corcorax melanorhamphos, Corcoracidae) are a common, breeding resident in and around the city of Canberra in the Australian Capital Territory. We compared five measures of reproductive success between the urban and non‐urban populations of choughs to investigate the effect of urbanization on this cooperatively breeding species. Urban choughs initiated breeding earlier than their non‐urban counterparts and were more likely to suffer nest failures. However, there was no difference in the number of successful nests in a season or the number of fledglings produced per successful nesting attempt. A greater proportion of fledglings survived their first 12 months in the non‐urban habitat. We suggest that increased rates of nest predation and fledgling mortality in the urban environment may have a negative effect on reproductive success and remove any advantage that might be gained through a longer breeding season. Possible effects of urbanization on the social and genetic structure of white‐winged choughs are also discussed.     

Should I Stay or Should I Go Now: Late Establishment and Low Site Fidelity as Alternative Territorial Behaviors

In species in which males defend territories for breeding, males may differ in territorial behavior; alternative behaviors among territorial males are not well understood. In our long‐term study of partially‐migratory song sparrows, we have observed that most territorial males establish territories before females begin nesting and remain site‐faithful both within and between breeding seasons; however, some males establish territories later in the season (late establishers) and/or change territory locations either within or between seasons (movers). Whether late establishment or moving are equally successful strategies for territory defense, or best‐of‐bad‐job options, is not known. Here, we compare the frequencies of these behaviors to demographic variables over a 9‐yr period and compare lifetime tenure and early season nesting success for males who differ in site fidelity and timing of territory establishment. Across years, late establishing was negatively correlated with the return rate of previously territorial males; moving was positively correlated with the number of occupied territories at the start of the breeding season (territory density). While moving was independent of number of years on territory, late territory establishment only occurred in a male’s first year as a territory holder. Of 88 males, 25% established their first territory late, primarily in undefended space; 31% moved. Late and early establishers did not differ in lifetime tenure; movers, however, had longer lifetime tenure than site‐faithful males. Among early establishers, movers and non‐movers did not differ in the number of successful early nests/year or number of young fledged/year; among late establishers, however, movers had significantly higher early nesting success by both measures. Late establishers who moved had higher early season nesting success and higher early season nesting success/year than site‐faithful early establishers. Thus, individual variation in the timing of territory establishment and site fidelity may be facultative alternative territorial strategies.     

Survival rates, causes of failure and productivity of Skylark Alauda arvensis nests on lowland farmland

This paper analyses data from 995 Skylark Alauda arvensis nests found on lowland farms in southern England from 1996 to 1998. The majority of recorded nest failures were caused by predation except in agricultural grass, where trampling and agricultural operations were equally important. Nest survival rates varied between crop types, nests in cereals being around twice as likely to succeed as nests in grass or set-aside. In cereals, nest survival rates increased with increasing distance from the nearest tramline and declined over the course of the breeding season. Predator control also had a significant independent effect on nest survival rates. On one farm where many other factors were held constant, a highly significant increase in nest survival rates from 12.3% to 40.7% coincided with the introduction of intensive predator control, which also appeared to bring forward mean laying dates. Most environmental factors explaining significant variation in nest survival rates did so only at the chick stage. The mean number of chicks produced per nesting attempt was 1.26 in cereals, 0.78 in set-aside and 0.63 in grass, the differences being due primarily to variation in nest survival rates. Low densities of Skylark territories in cereal crops are not therefore the consequence of low breeding success at the scale of the individual nest and probably reflect limitations on the number of attempts made in a season. Measures taken to improve the attractiveness of cereal crops as a nesting habitat for Skylarks, and beneficial changes in grassland management, are likely to increase overall productivity.     

Nonideal nest box selection by tree swallows breeding in farmlands: Evidence for an ecological trap?

Animals are expected to select a breeding habitat using cues that should reflect, directly or not, the fitness outcome of the different habitat options. However, human‐induced environmental changes can alter the relationships between habitat characteristics and their fitness consequences, leading to maladaptive habitat choices. The most severe case of such nonideal habitat selection is the ecological trap, which occurs when individuals prefer to settle in poor‐quality habitats while better ones are available. Here, we studied the adaptiveness of nest box selection in a tree swallow (Tachycineta bicolor) population breeding over a 10‐year period in a network of 400 nest boxes distributed along a gradient of agricultural intensification in southern Québec, Canada. We first examined the effects of multiple environmental and social habitat characteristics on nest box preference to identify potential settlement cues. We then assessed the links between those cues and habitat quality as defined by the reproductive performance of individuals that settled early or late in nest boxes. We found that tree swallows preferred nesting in open habitats with high cover of perennial forage crops, high spring insect biomass, and high density of house sparrows (Passer domesticus), their main competitors for nest sites. They also preferred nesting where the density of breeders and their mean number of fledglings during the previous year were high. However, we detected mismatches between preference and habitat quality for several environmental variables. The density of competitors and conspecific social information showed severe mismatches, as their relationships to preference and breeding success went in opposite direction under certain circumstances. Spring food availability and agricultural landscape context, while related to preferences, were not related to breeding success. Overall, our study emphasizes the complexity of habitat selection behavior and provides evidence that multiple mechanisms may potentially lead to an ecological trap in farmlands.