Experimental food supplementation affects the physical development,behaviour and survival of Little Owl Athene noctua nestlings

In birds, energy supply during growth is a major predictor of the fledglings' physical condition and survival prospects. Differential quantity and quality of fledglings produced under varying nestling food supplies are likely to affect the number of offspring that recruit into the breeding population. However, the underlying mechanisms and associated consequences are still poorly known. Using a partial cross‐fostering and food supplementation experiment, we estimated the effect of variation in food supply during growth on nestling survival and fledgling phenotypic traits of Little Owls Athene noctua. Survival to fledging was much higher in food‐supplemented nestlings (98.6%) than in control nestlings (82.4%). Furthermore, supplemented nestlings were on average 8.9 g heavier and were more likely to develop subcutaneous fat deposits (99.4 vs. 73.7% of treatment and control nestlings, respectively). Supplemented nestlings also had on average longer wings than control nestlings, but tarsi and culmen did not differ significantly. Furthermore, experimentally supplemented fledglings struggled more when handled and emerged sooner from tonic immobility than control fledglings. The irises of supplemented fledglings were less intensely coloured. The experimentally induced changes in nestling development probably affect individual performance beyond fledging. Nestlings from orchard‐dominated habitats were larger than those from habitats dominated by arable land. As nestling food supply is largely determined by natural food availability, we conclude that habitat quality affects Little Owl productivity and offspring quality, and ultimately, population dynamics.     

Happy to breed in the city? Urban food resources limit reproductive output in Western Jackdaws

Urban areas expand worldwide, transforming landscapes and creating new challenging habitats. Some bird species, mainly omnivorous feeding on human waste and cavity nesters, commonly breed in these habitats and are, therefore, regarded as urban‐adapted. Although urban areas may provide new nesting sites and abundant human waste, the low breeding success found in some of these species suggests that the poor protein content in human waste might limit breeding parameters. We investigated whether the breeding success of a cavity nester and omnivorous species commonly breeding in urban areas, the Western Jackdaw (Corvus monedula), depended on the availability of good‐quality non‐urban food. We approached the objective by combining a literature review and experiments in the field. With the literature review, we compared jackdaw populations in different habitats across Europe and found that clutch size and number of fledglings per pair decreased with distance to non‐urban foraging grounds, even after controlling for the effect of colony size, latitude, and climate. In two experiments, we tested whether the breeding success of urban pairs could be increased by supplementing high‐quality food, first only during egg formation and second also until chick fledging. Food supplementation during egg formation led to larger eggs and higher hatching success than in urban control nests, but this did not result in higher chick survival. However, when food supplementation was prolonged until fledging in the second experiment, we observed a significant increase of nestling survival. These findings highlight that research and management actions should not only focus on species displaced by urbanization, but also on “urban‐adapted” species, as they might be suffering from a mismatch between availability of nesting sites in buildings and adequate non‐urban food resources. In these cases, nest sites should be provided in or close to adequate food resources.     

Testing sources of variation in nestling‐stage nest success of Florida Scrub‐Jays in suburban and wildland habitats

Human modification of habitats can reduce reproductive success by providing novel cues to which birds may respond with behaviors that are actually maladaptive in those environments. Ad libitum human‐provided foods may provide the perception that urban habitats are food‐rich even as natural food availability decreases. Similarly, human activity may increase the perception that predation risk is high even as natural predators may decrease in abundance. In response, birds may reduce parental care with a subsequent cost to successful reproduction. Florida Scrub‐Jays (Aphelocoma coerulescens) in suburban areas have lower nest success during the nestling period than do wildland jays, possibly the result of such maladaptive responses, but maybe because of ecological differences with wildlands. We manipulated adult perception of predation risk and the availability of nestling foods in suburban and wildland areas to determine if these factors influenced parental care and nestling begging, and if the behavioral responses of adults influence nest survival during the nestling stage. Experimentally increasing perception of predation risk reduced parental care by both suburban and wildland females, but did not influence care by males. Increasing food availability, but not predation risk, had little influence on parental care, but resulted in decreased nestling begging rates and an increase in the frequency (pitch) of begging calls in both habitats. However, neither parental care nor food availability influenced nest survival during the nestling stage. Instead, the presence of helpers was the most important variable in nest survival analyses, suggesting that habitat‐specific differences in nest survival during the nestling stage were not simply the result of maladaptive parental behavior or shortage of nestling food resources in the suburban habitat. The lack of helpers combined with ecological differences, such as the abundance of nest predators, may be why fewer nests of Florida Scrub‐Jays survive during this stage in suburban areas.     

Breeding time,food supply and fitness components of Blue Tits Parus caeruleus in Mediterranean habitats

We analysed the relationship between the timing of food availability and within-season variation of both reproductive success and nestling body size of Blue Tits Parus caeruleus in Mediterranean habitats. Synchronization between food supply and reproduction was expected to be positively related to fitness components. We measured deviation from maximum food supply using a parameter that we called “time-lag”, which quantifies the degree of synchronization between the date of maximum food requirements by the nestlings and the date of maximum caterpillar supply in the habitat. This parameter was expected to be related to reproductive success as measured by the number and body-condition of fledglings. The predictions were that time-lag should be negatively correlated with the proportion of nestlings raised to fledging and the size of the fledglings. These predictions have been tested in different types of habitat. The results demonstrate that caterpillar supply during a critical nestling period can have a strong influence on fitness components. As predicted, synchronization with caterpillar supply is positively related to the number and body size of fledglings. Since there is large between-habitat variation in the timing of food supply, the key issue seems to be the adjustment to local patterns of food availability.     

Reproductive success and nestling diet in the Lesser Spotted Woodpecker ( Picoides minor ): the early bird gets the caterpillar

Populations of Lesser Spotted Woodpecker (Picoides minor) are decreasing in size, necessitating management strategies. However, data on the reproductive biology of this species are scarce. The aim of this study was to investigate the influence of timing of breeding and nestling diet on the reproductive success of the Lesser Spotted Woodpecker and thereby contribute to an understanding of the determinants influencing its reproductive success. During 6 study years between 1996–2003, we investigated various variables of reproductive success in a Lesser Spotted Woodpecker population in the Taunus low mountain range, Germany. We observed nestling feeding at breeding holes to assess the composition of nestling diet and how it changes during the season. Clutch size, number of fledglings as well as body mass of the nestlings declined with the start of egg laying. Pairs composed of individuals that had breed together in previous years started egg laying earlier than newly established pairs. Nestling diet consisted mainly of aphids, caterpillars, craneflies and wood-living larvae. The composition changed considerably within the breeding season in both early and late broods. However, early broods were provided with caterpillars more frequently, whereas late broods received wood-dwelling larvae more often. Our results suggest that Lesser Spotted Woodpeckers time their breeding so as to coincide with the early nestling stage with the highest availability of caterpillars, since this is the time of highest energy demand. Moreover, food availability declines during the breeding season, and the decline in reproductive performance seems to be an effect of this development.     

Geographical variation in reproductive ageing patterns and life‐history strategy of a short‐lived passerine bird

We investigated differences in ageing patterns in three measures of breeding performance in populations of barn swallows Hirundo rustica L. from Spain and Denmark differing in breeding latitude and hence migration distance and duration of the breeding season. We found differences in ageing patterns between populations. Generally, young (i.e. yearling) and old females (i.e. ≥ 5 years of age) laid their first eggs later and produced smaller clutches than middle‐aged females (i.e. 2–4 years of age) in both populations. The southernmost population (i.e. Spanish) showing the shorter migratory distance experienced a greater within‐individual increase in timing of breeding and clutch size in early life and a greater within‐individual decrease in laying date but not in clutch size during senescence compared with the northernmost population (i.e. Danish). We also found that the number of fledglings produced annually was related to the age of the two members of the breeding pairs with pairs composed of young and old females performing less well than breeding pairs composed of middle‐aged females. We did not find reproductive senescence for the age of the male while controlling for the age of the female on the number of fledglings produced annually by the breeding pair. Differential survival between individuals did not explain age effects on laying date or annual clutch size in neither population. However, the increase in the number of fledglings produced annually with age was partly explained by the disappearance of poor‐quality members of the pairs, mainly poor‐quality males. Age‐related breeding success (i.e. number of fledglings) was similar for barn swallows from Spain and Denmark. Therefore, the study of ageing patterns and life‐history strategies in free‐ranging animals from more than a single population can throw new light on life‐history theory, population dynamics and evolutionary studies of senescence.     

Comparing food limitation among three stages of nesting: supplementation experiments with the burrowing owl

Food availability is an important limiting factor for avian reproduction. In altricial birds, food limitation is assumed to be more severe during the nestling stage than during laying or incubation, but this has yet to be adequately tested. Using food‐supplementation experiments over a 5‐year period, we determined the degree and timing of food limitation for burrowing owls (Athene cunicularia) breeding in Canada. Burrowing owls are an endangered species and food limitation during the nestling stage could influence reproductive performance of this species at the northern extent of their range. Supplemented pairs fledged on average 47% more owlets than unfed pairs, except during a year when natural food was not limiting (i.e., a prey irruption year). The difference in fledgling production resulted from high nestling mortality in unfed broods, with 96% of all nestling deaths being attributed to food shortage. Supplemental feeding during the nestling period also increased fledgling structural size. Pairs fed from the start of laying produced the same number of hatchlings as pairs that received no supplemental food before hatch. Furthermore, pairs supplemented from egg laying to fledging and pairs supplemented during the nestling period alone had the same patterns of nestling survival, equal numbers of fledglings, and similar fledgling mass and structural size. Our results provide empirical support for the hypothesis that the nestling period is the most food‐limited phase of the breeding cycle. The experimental design we introduce here could be used with other altricial species to examine how the timing of food limitation differs among birds with a variety of life‐history strategies. For burrowing owls, and other species with similar life histories, long‐term, large‐scale, and appropriately timed habitat management increasing prey abundance or availability is critical for conservation.     

The importance of food quantity and quality for reproductive performance in alpine water pipits (Anthus spinoletta)

Studies relating reproduction to food availability are usually restricted to food quantity, but ignore food quality and the effects of habitat structure on obtaining the food. This is particularly true for insectivorous birds. In this study we relate measures of reproductive success, time of reproduction and nestling size of water pipits (Anthus spinoletta) to biomass, taxonomic composition and nutritional content of available food, and to vegetation structure and distance to feeding sites. Clutch size was positively correlated with the proportion of grass at the feeding sites, which facilitates foraging. This suggests that water pipits adapt their clutch size to environmental conditions. Also, pipits started breeding earlier and produced more fledglings when abundant food and a large proportion of grass were available, probably because these conditions allow the birds to gain more energy in less time. The number of fledglings was positively correlated with the energy content of available food. No significant relationships were found between feeding conditions and nestling size or the time that nestlings took to fledge. This suggests that water pipits do not invest more in individual nestlings when food conditions are favourable but rather start breeding earlier and produce more young. Taxonomic composition and nutritional content of prey were not correlated with any of the reproductive parameters, indicating that profitability rather than quality of food affects reproductive success. Received: 31 May 1996/Accepted: 12 August 1996     

Urbanization,nestling growth and reproductive success in a moderately declining house sparrow population

Ecological conditions are likely to change with increasing urbanization, influencing the demography and size of animal populations. Although one of the most tightly linked species to humans, the house sparrow has been suffering a significant decline worldwide, especially in European cities. Several factors have been proposed to explain this conspicuous loss of urban sparrows, but studies evaluating these factors are usually restricted to Britain where the decline was very drastic, and it is unclear whether similar or different processes are affecting urban populations of the species elsewhere. In this study we investigated the reproductive success of urban and rural sparrows in a central European country, Hungary where our census data indicate a moderate decline during the last decade. We found that rural pairs produced more and larger fledglings than suburban pairs, and the difference remained consistent in two years with very contrasting meteorological conditions during breeding. This difference is likely explained by habitat differences in nestling diet, because we found that 1) rural parents provided large prey items more often than suburban parents, 2) birds from differently urbanized habitats produced fledglings of similar number and size in captivity under identical rearing conditions with ample food for nestlings, and 3) in a cross‐fostering experiment, nestlings tended to grow larger in rural than in suburban nests irrespective of their hatching environment. These results agree with those found in a recent British study, indicating that poor nestling development and survival due to inadequate diet may be widespread phenomena in urbanized habitats.     

Double-brooding and annual breeding success of great tits in urban and forest habitats

Urban areas differ from natural habitats in several environmental features that influence the characteristics of animals living there. For example, birds often start breeding seasonally earlier and fledge fewer offspring per brood in cities than in natural habitats. However, longer breeding seasons in cities may increase the frequency of double-brooding in urban compared with nonurban populations, thus potentially increasing urban birds’ annual reproductive output and resulting in lower habitat difference in reproductive success than estimated by studies focusing on first clutches only. In this study, we investigated 2 urban and 2 forests great tit Parus major populations from 2013 to 2019. We compared the probability of double-brooding and the total number of annually fledged chicks per female between urban and forest habitats, while controlling for the effects of potentially confounding variables. There was a trend for a higher probability of double-brooding in urban (44% of females) than in forest populations (36%), although this was not consistent between the 2 urban sites. Females produced significantly fewer fledglings annually in the cities than in the forest sites, and this difference was present both within single- and double-brooded females. Furthermore, double-brooded urban females produced a similar number of fledglings per season as single-brooded forest females. These results indicate that double-brooding increases the reproductive success of female great tits in both habitats, but urban females cannot effectively compensate in this way for their lower reproductive output per brood. However, other mechanisms like increased post-fledging survival can mitigate habitat differences in reproductive success.     

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.     

Supplementary feeding increases Common Buzzard Buteo buteo productivity but only in poor‐quality habitat

Temporal heterogeneity in the effects of food supply during the breeding season on the productivity of the Common Buzzard Buteo buteo was investigated in a supplementary feeding experiment. Pairs were fed artificially (1) before egg‐laying, (2) after chicks hatched and (3) continuously throughout the season, and compared with (4) unfed controls. Pairs fed before egg‐laying had marginally larger clutches than those not fed, but lay date, egg volume and weight, brood size and hatching success were unaffected. Territorial quality had far greater effects, with pairs nesting in low‐quality habitats (bog, scrub and semi‐natural grassland) laying later and having lower hatching success, smaller broods and fewer fledglings than those in more productive agricultural landscapes. Supplementary feeding after egg hatching neutralized the negative effect of poor habitat, resulting in fed birds having significantly more fledglings. This study emphasizes the importance of food availability when provisioning chicks in suboptimal habitats and has implications for the success of diversionary feeding in reducing game losses to Buzzards.     

Quantitative genetics of plumage color: lifetime effects of early nest environment on a colorful sexual signal

Phenotypic differences among individuals are often linked to differential survival and mating success. Quantifying the relative influence of genetic and environmental variation on phenotype allows evolutionary biologists to make predictions about the potential for a given trait to respond to selection and various aspects of environmental variation. In particular, the environment individuals experience during early development can have lasting effects on phenotype later in life. Here, we used a natural full‐sib/half‐sib design as well as within‐individual longitudinal analyses to examine genetic and various environmental influences on plumage color. We find that variation in melanin‐based plumage color – a trait known to influence mating success in adult North American barn swallows (Hirundo rustica erythrogaster) – is influenced by both genetics and aspects of the developmental environment, including variation due to the maternal phenotype and the nest environment. Within individuals, nestling color is predictive of adult color. Accordingly, these early environmental influences are relevant to the sexually selected plumage color variation in adults. Early environmental conditions appear to have important lifelong implications for individual reproductive performance through sexual signal development in barn swallows. Our results indicate that feather color variation conveys information about developmental conditions and maternal care alleles to potential mates in North American barn swallows. Melanin‐based colors are used for sexual signaling in many organisms, and our study suggests that these signals may be more sensitive to environmental variation than previously thought.     

Time and travelling costs during chick‐rearing in relation to habitat quality in Little Owls Athene noctua

In many bird species, parents adjust their home‐ranges during chick‐rearing to the availability and distribution of food resources, balancing the benefits of energy intake against the costs of travelling. Over recent decades, European agricultural landscapes have changed radically, resulting in the degradation of habitats and reductions in food resources for farmland birds. Lower foraging success and longer foraging trip distances that result from these changes are often assumed to reduce the reproductive performance of parents, although the mechanisms are not well understood. We tested the behavioural response of chick‐rearing Little Owls Athene noctua to variation in habitat diversity in an agricultural landscape. We equipped females with GPS loggers and received adequate range‐use data for 19 individuals (6063–14 439 locations per bird). In habitats dominated by homogeneous cropland habitats, home‐ranges were over 12 ha in size, whereas in highly diverse habitats they were below 2 ha. Large home‐ranges were associated with increased flight activity (117% of that of birds in small home‐ranges) and distances travelled per night (152%), increased duration of foraging trips (169%) covering larger distances (246%), and reduced nest visiting rates (81%). The study therefore provides strong correlative evidence that Little Owls breeding in monotonous farmland habitats expend more time and energy for a lower benefit in terms of feeding rates than do birds in more heterogeneous landscapes. As nestling food supply is the main determinant of chick survival, these results suggest a strong impact of farmland characteristics on local demographic rates. We suggest that preserving and creating islands of high habitat diversity within uniform open agricultural landscapes should be a key target in the conservation of Little Owl populations.     

Do baseline glucocorticoids simultaneously represent fitness and environmental quality in a declining aerial insectivore?

Glucocorticoids (GCs) are often interpreted as indicators of disturbance, habitat quality, and fitness in wild populations. However, since most investigations have been unable to examine habitat variability, GC levels, and fitness simultaneously, such interpretations remain largely unvalidated. We combined a quantification of two habitat types, a manipulation of foraging ability (feather‐clipping just prior to nestling rearing), multiple baseline plasma GC measures, and multi‐year reproductive monitoring to experimentally examine the linkages between habitat quality, GCs, and fitness in female tree swallows Tachycineta bicolor. Control females experiencing the higher early‐season food resources of inland–pasture habitat laid larger clutches, but fledged an equal number but lower mass offspring compared to those in riparian–cropland habitat. Despite these differences in reproductive success, females nesting in the two habitat types did not differ in baseline GC levels at the early‐ or late‐breeding stage. Feather‐clipping reduced provisioning rate in both habitat types. However, baseline GC levels were affected in a habitat‐specific way; only individuals in inland–pasture habitats showed an increase in GCs. Despite this difference in GC levels, the manipulation did not influence offspring mass, reproductive output, adult return rate (a proxy for survival) to the following year, or reproductive success in the subsequent year. Nonetheless, regardless of treatment, individuals with higher GC levels during the late breeding stage returned in the following year with higher GC levels at incubation, indicating a long‐term effect on future GC levels. Our results indicate that environmental changes (e.g. foraging conditions) can have consequences for body condition, behaviour, and current and future baseline GC levels without concomitant influences on fitness, and that differences in fitness components between habitats may not be reflected in baseline GC levels. These results illustrate that baseline GCs may not simultaneously reflect environmental quality and fitness, potentially limiting their application in ecological and conservation settings.     

Parental care of kestrels living in stable and varying environmental conditions

We studied parental behaviour of two populations of Eurasian kestrels (Falco tinnunculus) living in environments with different reproductive constraints: a northern Italy population, resident, with relatively long reproductive season and almost stable prey availability, and a western Finland population, long-distance migrant, with short reproductive season and fluctuating availability of main prey (voles). The Finnish population was studied during a low vole availability year. We aimed to test the supposition that in variable environments the few pairs capable of breeding during low prey availability years show high-quality parental care. To estimate the quality of parental care in both Finnish and Italian pairs we measured the difference in the number of prey delivered by the male and by the female within each pair. The parental care, measured by that parameter, was greater among Italian pairs during both the incubation and early nestling periods. However, this was not the case in the late nestling period, when food requirements of offspring are greater. Moreover, the Finnish and Italian pairs produced about the same number of fledglings. Therefore, our data supported the initial supposition and we conclude that the few Finnish pairs capable of breeding in a low vole availability year show parental care similar to Italian pairs.     

Climate change and micro-geographic variation in laying date

Climate change has been shown to affect the timing of reproduction, with earlier reproduction being associated with an increase in temperature. Changes in the timing of reproduction arise from changes in food availability as well as other factors, and differences in the timing of reproduction among sites may cause sites with early reproduction to contribute disproportionately to local recruitment. In this study, spatial variation in the laying date of barn swallows Hirundo rustica at 39 sites in a 45-km²study area during the period 1971–2004 was used to investigate micro-geographic patterns in the timing of breeding. I found that individuals breeding at sites with early reproduction had a disproportionately large reproductive success. Early sites were characterized by early plant phenology, as determined by the date of leafing of the broad-leaved elm Ulmus glabra and date of flowering of the snowdrop Galanthus nivalis during a single year. Such early sites showed greater advancement in laying date between 1971 and 2004 than the average site. Early sites were also generally occupied during more years by a larger number of breeders than were late sites. Breeders at early sites produced more fledglings, and breeders at such sites were characterized by a smaller adult body size and larger secondary sexual characters than individuals at the average site. These observations are consistent with the hypothesis that temporal changes in the timing of reproduction occur as a consequence of differential recruitment at phenologically early sites that produce disproportionately large numbers of offspring. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Urbanisation lowers great tit Parus major breeding success at multiple spatial scales

While numerous studies have reported negative effects of urbanisation on birds, few have examined the role of urban scale in influencing breeding success. Furthermore, many studies have relied on qualitative rather than quantitative assessments of urbanisation. This study sought to address these issues by testing the effects of urbanisation, measured at two spatial scales, on the breeding success of great tits Parus major. A nested study design, incorporating over 400 nestboxes, was used in study sites across northern Belgium with a priori quantified degrees of urbanisation at both local and regional scales. All measured breeding parameters were found to vary at one or both spatial scales of urbanisation; in more urbanised areas great tits displayed advanced laying dates but lower breeding success compared to rural areas, with smaller clutch sizes, lower nestling masses and fewer fledglings per egg. Importantly, urbanisation effects were not limited to big cities as birds breeding in gardens or parks in small towns also had comparatively low success. We found that both regional‐ and local‐scale urbanisation had consistent significant effects on laying date, clutch size and nestling mass, while the number of fledglings per egg was negatively influenced by local‐scale urbanisation only. Results of this study therefore highlight the importance of utilising multiple spatial scales in analysing urbanisation effects, as well as the potential negative impact of local urbanisation on breeding success. This calls for further investigation into mechanisms driving urbanisation effects and how these may vary at different scales.     

Benefits of extra food to reproduction depend on maternal condition

The amount of food resources available to upper‐level consumers can show marked variations in time and space, potentially resulting in food limitation. The availability of food resources during reproduction is a key factor modulating variation in reproductive success and life‐history tradeoffs, including patterns of resource allocation to reproduction versus self‐maintenance, ultimately impacting on population dynamics. Food provisioning experiments constitute a popular approach to assess the importance of food limitation for vertebrate reproduction. In this study of a mesopredatory avian species, the lesser kestrel Falco naumanni, we provided extra food to breeding individuals from egg laying to early nestling rearing. Extra food did not significantly affect adult body condition or oxidative status. However, it increased the allocation of resources to flight feathers moult and induced females to lay heavier eggs. Concomitantly, it alleviated the costs of laying heavier eggs for females in poor body condition, and reduced their chances of nest desertion (implying complete reproductive failure). Extra food provisioning improved early nestling growth (body mass and feather development). Moreover, extra food significantly reduced the negative effects of ectoparasites on nestling body mass, while fostering forearm (a flight apparatus trait) growth among highly parasitized nestlings. Our results indicate that lesser kestrels invested the extra food mainly to improve current reproduction, suggesting that population growth in this species can be limited by food availability during the breeding season. In addition, extra food provisioning reduced the costs of the moult–breeding overlap and affected early growth tradeoffs by mitigating detrimental ectoparasite effects on growth and enhancing development of the flight apparatus with high levels of parasitism. Importantly, our findings suggest that maternal condition is a major trait modulating the benefits of extra food to reproduction, whereby such benefits mostly accrue to low‐quality females with poor body condition.     

Hawk mimicry in cuckoos and anti‐parasitic aggressive behavior of barn swallows in Denmark and China

Hosts of brood parasites defend their nests against parasitism by aggression and subsequently, if parasitized, by rejection of the parasite egg or nestling. Cuckoos have evolved plumage mimicry with convergence towards the phenotype of Accipiter hawks that are common predators of cuckoo hosts. Here we tested two alternative hypotheses 1) whether barn swallows Hirundo rustica have evolved less aggressive behavior towards cuckoos in areas of sympatry with more abundant Accipiter hawks; and 2) whether barn swallows have evolved more aggressive anti‐parasite behavior in areas with a single species of cuckoo. We presented dummies of common cuckoo Cuculus canorus, sparrowhawk Accipiter nisus and Oriental turtledove Streptopelia orientalis (a benign control) at the nests of barn swallows during breeding, while recording intensity of response and proximity of barn swallows to the dummy. We demonstrated that cuckoos moved away when attacked aggressively and approached more closely by barn swallows showing that barn swallow behavior was efficient at driving away cuckoos. Barn swallows were significantly more aggressive and approached cuckoo and sparrowhawk dummies more closely in Denmark than in China, despite sparrowhawks being relatively more common in Denmark. Responses towards cuckoo dummies differed from responses towards sparrowhawk dummies, showing that barn swallows distinguished between the two different causes of danger. These findings are inconsistent with a less aggressive response towards cuckoo dummies in areas of sympatry with more abundant Accipiter hawks, but consistent with the alternative hypothesis that barn swallows have evolved more aggressive behavior towards cuckoos in areas with a single species of brood parasite, but not in areas with multiple species of parasites, where it is harder for hosts to tell the difference.