Differences in the onset of laying between island and mainland Mediterranean Blue Tits Parus caeruleus: phenotypic plasticity or genetic differences?

Blue tits Parus caeruleus breeding in deciduous or in evergreen woodlands on the mainland of Europe start to lay 3–4 weeks earlier than Blue Tits on the island of Corsica breeding in evergreen woodlands. A similar difference in average laying date between the two Blue Tit populations has been found in a relatively small sample of captive birds held in outdoor aviaries on the mainland. To provide data on the habitat contrast in Corsica for comparison with that on the mainland, a new study area was started in one of the few deciduous woodlands on Corsica. In addition, new data on laying dates of captive Blue Tits were gathered to increase the sample size of independent pairs in aviaries. This study shows that the habitat effect (deciduous ν evergreen woodland) on the average laying date of Blue Tits is relatively small in comparison with the geography effect (European mainland ν Corsica). The laying date differences between mainland and insular Blue Tits persisted whatever the type of habitat (deciduous, evergreen or outdoor aviaries on the mainland). The data support the hypothesis that the observed differences in the onset of laying between the mainland and insular Blue Tits were mainly influenced by genetic effects and relatively little by non-genetic maternal effects of by genotype—environment interactions.     

What are the consequences of being a downy oak (quercus pubescens) or a holm oak (Q. ilex) for breeding blue tits (Parus caeruleus)?

By comparison with deciduous oaks, the lower yearly production of new leaves in sclerophyllous oaks is hypothesized to have several consequences on animal communities. In particular, the production of arthropod communities that feed upon the leaves should be lower in sclerophyllous than in deciduous oaks, this causing changes in breeding patterns and the demographic balance in insectivorous birds. Studies in both deciduous and sclerophyllous habitats in southern France have shown that: 1) the spring development of new leaves occurs later and more slowly in sclerophyllous than in deciduous oaks, 2) the biomass of caterpillars is much lower in sclerophyllous oak forests, and 3) there is a large variation in life history traits of the Blue Tit depending in which type of habitat they breed. Laying date occurs later and clutch size is lower in sclerophyllous habitats than in deciduous habitats. The evolution of life history traits is discussed according to whether poor sclerophyllous habitats are isolated (e.g. on Corsica) or are parts of landscapes including both habitat types.     

Nestling diet variation in an insular Mediterranean population of blue tits Parus caeruleus: effects of years,territories and individuals

Mediterranean evergreen forests of Corsica are characterized by relatively high species diversity of arthropods with low population densities. Food is never superabundant for Corsican blue tits Parus caeruleus. This study focused on the composition of the food of blue tit nestlings and especially on two main components, caterpillars and spiders. The nestling diet was studied for two years using 8-mm cameras that automatically took photographs of adult birds with food. The diet was composed of c. 50% caterpillars and c. 30% spiders. There were between-year and between-individual differences in these proportions. In both years of the study the proportion of caterpillars declined during the course of the breeding season. Individual and time effects on prey sizes were also observed. Pairs and individuals were fairly constant in the proportions of prey over the feeding period. Different food items were not brought in runs. These findings suggest that strong food limitation exists on Corsica, which can considerably influence life-history traits of the blue tit.     

Local specialization and maladaptation in the Mediterranean blue tit (Parus caeruleus)

Although the evolutionary interplay between gene flow and local adaptation of organisms in heterogeneous environments has been widely discussed from a theoretical point of view, few empirical studies have been designed to test predictions on the consequences of habitat patchiness on the evolution of life history traits. Using blue tits in Mediterranean habitat mosaics as a model, we defined two nested levels of habitat heterogeneity: an inter-regional level which compares two isolated landscapes (mainland, southern France vs the island of Corsica), and an intra-regional level which compares two habitat types within each landscape (deciduous vs evergreen trees). Deciduous habitats are more common than evergreen habitats on the mainland whereas the opposite is true on the island. Results suggest that: (1) on a regional scale, each population is specialized to the more common habitat, i.e. life history traits have evolved in such a way that breeding success is high; (2) in the less common habitats within each landscape, birds are clearly mistimed because they mismatch the best period of food availability, and hence their breeding success is lower; and (3) the density of the populations and the morphometry of the birds support the model of ideal despotic distribution. These results, which are supported by preliminary data on the genetic variation and gene flow of populations in the mainland landscape, are consistent with a source-sink model of population structure within each landscape. They are discussed in the framework of metapopulation theory and habitat selection models.     

Parental provisioning behaviour in Pied Flycatchers Ficedula hypoleuca is well adjusted to local conditions in a mosaic of deciduous and coniferous habitat

Capsule Pied Flycatchers are better able than Great Tits to adjust their feeding behaviour to varying conditions in the same area.

Aims Great Tits breeding in a mosaic of deciduous and coniferous forests in the northern temperate region exhibit consistently lower breeding success in their preferred deciduous habitat than in coniferous habitat. This was explained by the unexpectedly poor nestling feeding conditions in deciduous forests of this region. We studied whether the same paradox applies to Pied Flycatchers that occupy the same habitats in the same area.

Methods Parental provisioning behaviour was studied using video‐recording and experimental manipulation. Caterpillar abundance and basic breeding parameters were measured in different habitat types.

Results Parental provisioning frequency and the proportion of caterpillars in nestling diet was lower, while food objects were on average larger, in coniferous compared with deciduous habitat. However, the total volume of caterpillars and adult Lepidoptera delivered to nestlings did not differ between habitats. In contrast to Great Tits, offspring body parameters in Pied Flycatchers did not differ between habitat types.

Conclusions These results demonstrate how the relative suitability of particular habitat types varies between species and is dependent upon geographical location.     

On humans and wildlife in Mediterranean islands

Aim To investigate the effects of human‐induced landscape changes in Mediterranean islands on the ecological and evolutionary responses of bird communities and populations. The combination of mass extinction of large mammals and massive deforestation by humans was hypothesized to produce new selection regimes to which organisms were likely to respond. Habitat selection and niche breadth have been investigated at the scale of species, and phenotypic variation at the scale of local populations. Location The study was carried out along habitat gradients and in habitat mosaics at different spatial scales on the island of Corsica and in areas of similar size and structure in continental France. Methods Two sets of gradients have been used for investigating habitat selection and niche breadth: gradients of altitude, and gradients of vegetation structure. Population studies focused on the blue tit, Cyanistes caeruleus. Large samples of breeding attempts by this species in 10 habitats provided detailed data on phenotypic variation of fitness‐related traits both on Corsica and on the mainland. Results The extent of niche space used by birds differed substantially depending on which habitat gradient was considered. Many species have been found to contract their habitat niche along the elevation gradient on Corsica compared with the mainland, whereas all species in the vegetation gradient broadened their niche on the island. Breeding patterns of the blue tit differed considerably depending on whether they settle in deciduous oaks (Quercus humilis) or in evergreen sclerophyllous oaks (Quercus ilex). Phenotypic variation of breeding traits was much higher on the island, where more populations were correctly timed for the best breeding period than on the mainland, a pattern that is likely to result from lower dispersal of organisms on the island. Main conclusions The differences in observed niche breadth between the two series of habitat gradients is explained both by the species‐specific ecology of the species and the human‐induced environmental history of Corsica. Large‐scale landscape changes provided new opportunities for island colonization by non‐forest species, which are isolated as small, ‘fugitive’ local populations. In both gradients, forest species that are typical components of the Corsican bird fauna definitely expanded their niche and occupied a wider range of habitats on Corsica than on the mainland. At the population scale, landscapes included habitat patches with contrasted selection regimes, which resulted in high phenotypic variation for many fitness‐related traits. Reduced dispersal of birds on the island resulted in a much higher degree of local differentiation on Corsica than on the mainland.     

The effect of habitat quality on foraging patterns, provisioning rate and nestling growth in Corsican Blue Tits Parus caeruleus

Many bird species face seasonal and spatial variation in the availability of the specific food required to rear chicks. Caterpillar availability is often identified as the most important factor determining chick quality and breeding success in forest birds, such as tits Parus spp. It is assumed that parents play an important role in mediating the effect of environment on chick development. A reduction in prey availability should therefore result in increased foraging effort to maintain the amount of food required for optimal chick development. To investigate the capacity of adults to compensate for a reduction in food supply, we compared the foraging behaviour of Blue Tits Parus caeruleus breeding in rich and poor habitats in Corsica. We monitored the foraging effort of adults using radiotelemetry. We also identified and quantified prey items provided to nestlings by using a video camera mounted on the nest. We found that the mean travelling distance of adults was twice as great in the poor habitat as it was in the rich. Despite the marked difference in foraging distance, the proportion of optimal prey (caterpillars) in the diet of the chicks and the total biomass per hour per chick did not differ between the two habitats. We argue that relationships between habitat richness, offspring quality and breeding success cannot be understood adequately without quantifying parental effort.     

Simultaneous pituitary-gonadal recrudescence in two Corsican populations of male blue tits with asynchronous breeding dates

Animal populations living in geographically variable environments respond to different selection pressures. The adaptive character of the responses to environmental information determines the degree of synchrony of the breeding period with local optimal conditions. An example is provided by two populations of Mediterranean blue tits (Parus caeruleus) in Corsica, breeding in different habitats, with a 1-month difference in the onset of egg laying. This difference in the onset of lay is supposed to be adaptive because, although chicks from both populations are raised mostly on caterpillars, the timing of the appearance of caterpillars is earlier for populations of tits associated with deciduous oak trees than those associated with evergreen oak trees. Here, we show that, despite the difference in the timing of egg laying, males from these two populations start seasonal hypothalamo-hypophysial-testicular development at approximately the same time, in late winter. Specifically, the vernal recrudescence of brain GnRH-I perikarya and fibers, testes volume and song activity began around the same dates and proceeded at the same pace in late winter in both populations. Plasma testosterone and LH levels displayed seasonal variations that were shifted by less than 2 weeks compared to the 1-month difference in egg laying periods. We hypothesize that the strong selection pressures on these two populations to adapt the timing of their breeding seasons to their local environment may have acted mostly on the female egg laying dates, and not so much on the initiation and rate of seasonal recrudescence of the hypothalamo-hypophysial-testicular activity in males.     

短尾猴栖息地的季节变化

本文阐述了短尾猴栖息地的季节变化。春季,主要利用海拔570-1600米的常绿阔叶林、常绿、落叶阔叶混交林和落叶阔叶林;夏季,限于海拔1000米以上的落叶阔叶林。秋季的栖息地,在海拔650-1200米常绿阔叶林和常绿、落叶阔叶混交林;冬季仅利用海拔600-900米的常绿阔叶林。此外还报道了短尾猴四季的食物组成。并对栖息地的变化原因,做了初步探讨。     

基于地形因子的天童地区常绿树种和落叶树种共存机制研究

位于亚热带的浙江天童和古田山常绿阔叶林大样地分布有较高比例的落叶树种,那么它们与常绿树种的共存机制是什么?常绿树种和落叶树种生态习性差异较大,二者对生境的选择应有所不同,我们推测生境分化可能是两类植物实现共存的主要机制。为检验该假设,我们以天童20ha动态样地调查数据为依托,选择个体数≥20的55个常绿树种和42个落叶树种作为分析对象,用典范对应分析(CCA)研究了地形因子对二者分布的影响差异,用torus转换检验来分析常绿树种和落叶树种与各类地形生境的关联。结果如下:(1)CCA分析表明地形因子对常绿树种分布的解释量为19.2%,对落叶树种分布的解释量为7.0%。(2)torus转换检验结果表明,与沟谷成正关联的常绿树种和落叶树种的比例分别为16.4%和28.6%,成负关联的比例分别为40%和7%;与山脊成正关联的常绿树种和落叶树种的比例分别为41.8%和4.8%,成负关联的比例分别为10.9%和47.6%;与受干扰生境成正关联的常绿树种和落叶树种的比例分别为16.4%和42.9%。上述结果说明地形对常绿树种分布的影响大于落叶树种;两个植物类群对生境的选择多呈现相反格局,尤其是在沟谷生境和山脊生境,这进一步表明生境分化是常绿树种和落叶树种共存的重要机制之一,生态位理论在一定程度上能较好地解释亚热带常绿阔叶林物种多样性的维持。     

Influence of fine-scale habitat structure on nest-site occupancy,laying date and clutch size in Blue Tits Cyanistes caeruleus

Most birds have specific habitat requirements for breeding. The vegetation structure surrounding nest-sites is an important component of habitat quality, and can have large effects on avian breeding performance. We studied 13 years of Blue Tit Cyanistes caeruleus population data to determine whether characteristics of vegetation structure predict site occupancy, laying date and number of eggs laid. Measurements of vegetation structure included the density of English Oak Quercus robur, European Beech Fagus sylvatica, and other deciduous, coniferous and non-coniferous evergreen trees, within a 20-m radius of nest-boxes used for breeding. Trees were further sub-divided into specific classes of trunk circumferences to determine the densities for different maturity levels. Based on Principal Component Analysis (PCA), we reduced the total number of 17 measured vegetation variables to 7 main categories, which we used for further analyses. We found that the occupancy rate of sites and the number of eggs laid correlated positively with the proportion of deciduous trees and negatively with the density of coniferous trees. Laying of the first egg was advanced with a greater proportion of deciduous trees. Among deciduous trees, the English Oak appeared to be most important, as a higher density of more mature English Oak trees was associated with more frequent nest-box occupancy, a larger number of eggs laid, and an earlier laying start. Furthermore, laying started earlier and more eggs were laid in nest-boxes with higher occupancy rates. Together, these findings highlight the role of deciduous trees, particularly more mature English Oak, as important predictors of high-quality preferred habitat. These results aid in defining habitat quality and will facilitate future studies on the importance of environmental quality for breeding performance.     

Winter bird communities in the Åland archipelago: an island biogeographic point of view

The distribution of wintering land birds was examined in the archipelago of Åland (60°N, 20°E) along two ecological dimensions: (1) the island continuum from small skerries ((0.5 ha) to the mainland of Åland (97000 ha), and (2) the habitat range of the mainland of Åland. In the species-area relationship, both the exponential and power function models fitted well. The increase of the number of species with island size was interpreted as a consequence of increasing habitat diversity on larger islands. The smallest islands supported only few seed-eating species utilizing the islands in a fine-grained fashion; none of these birds belonged to the breeding fauna of the islands. With increasing island size and habitat diversity, insectivores and species tied to the human culture were added. On the mainland of Åland, more than half of the birds were observed near human habitations: bird communities wintering in pine forests were richer than in deciduous forests. Proportion of wintering species was greatest in the breeding communities of pine forests. The proportion of wintering species was presumably regulated by the degree of structural change in the habitat between summer and winter, this change being most drastic in deciduous habitats The species observed could be broadly classified into three feeding categories: opportunists ., omnivores and scavengers, dependent on the diverse food items provided by man; food specialists , species éating seeds and berries, ranging widely in the islands and habitats; and insectivorous habitat specialists , inhabiting coniferous habitats, and islands with coniferous (pine) forests. Opportunists, in particular, gel increasing wintering opportunities by the impact of man.     

The effects of spatial and temporal ecological variation on fatty acid compositions of wild great tits Parus major

In birds, fatty acids (FA) have three main functions; they are structural components of cell membranes, metabolic fuel, and inflammatory molecules. Environmental factors, such as diet and ambient temperature, affect FA composition, thereby function and ultimately fitness. Thus, variation in FA compositions can be the underlying mechanism for varying performance of birds in different habitats. Here we examine variation in plasma FA composition in nestling and adult great tits Parus major, between 1) deciduous and coniferous, and 2) sun‐exposed and shaded habitats. The main results revealed that nestlings had a higher proportion of α‐linolenic acid (α‐LNA) in deciduous habitats and arachidonic acid in coniferous habitats. This reflects a difference in caterpillar availability between habitats with the deciduous habitat being caterpillar‐rich, whereas the coniferous habitats are rich in spiders. In addition, α‐LNA increased with nestling body condition in the coniferous habitat, supporting the importance of caterpillars for fledging success in this species. In line with dietary intake, the proportion of the essential α‐LNA and linoleic acid (LA) increased over the course of the day for all birds. In the deciduous habitat, adult females showed a positive association between LA and body condition. Furthermore, habitat sun‐exposure showed significant interactions with body condition for polyunsaturated FAs in nestlings, and with saturated FA in adult males, which is in accordance with the homeoviscous hypothesis stating that the proportion of saturated FA should decrease with decreasing ambient temperature. Taken together, small‐scale heterogeneity in habitat structure significantly influences FA compositions of great tits. Many of the results can be linked to dietary, and possibly, ambient temperature differences between habitats. These habitat effects on FA compositions can lead to different capacities of individual birds to deal with infections and low temperatures, two stressors that cause major mortality among wild birds.     

Food supplementation in distinct Corsican oak habitats and the timing of egg laying by Blue Tits

ABSTRACT.   Resource constraints may impose physiological limitations on egg production and influence the timing of breeding in seasonally breeding birds. Food-supplementation experiments have demonstrated that food availability may influence the timing of egg laying, but the moderate response of birds in most studies suggests that the effect of food availability may be apparent only under certain ecological conditions. Experiments conducted in habitats that differ in ecological characteristics and natural availability of food sources are, therefore, needed to effectively examine the possible role of habitat-specific physiological constraints on the timing of breeding. We compared the response of Blue Tits ( Cyanistes caeruleus ) to supplemental feeding in four oak woodlands that differed in elevation, forest type, and food availability on the island of Corsica. We found that supplemental feeding advanced the date of egg laying (by about 1 week) at only one of four sites; a site dominated by evergreen holm oak where the availability of natural food was likely lower than at the other sites. Our results suggest that the response of Blue Tits to supplemental food depends on the natural level of resource abundance, but, in addition, that the effect of supplemental feeding on the timing of breeding appears to be small compared to the typical and, for our study sites, great (>1.5 mo) between-population variation in clutch initiation dates. Physiological limitations related to the acquisition of nutrients and energy may exert only a limited effect on the onset of breeding, and behavioral flexibility in the integration of fine-scale environmental signals (e.g., temperature and phenology) that predict future breeding conditions may be more important in explaining variation among populations in the timing of breeding.     

Forest fragmentation is associated with primary brood sex ratio in the treecreeper (Certhia familiaris)

We studied the primary brood sex ratio of an old-growth forest passerine, the Eurasian treecreeper (Certhia familiaris), along a gradient of forest fragmentation. We found evidence that male nestlings were more costly to produce, since they suffered twofold higher nestling mortality and were larger in body size than females. Furthermore, the proportion of males in the brood was positively associated with the provisioning rate and the amount of food delivered to the nestlings. During the first broods, a high edge density and a high proportion of pine forests around the nests were related to a decreased production of males. The densities of spiders, the main food of the treecreeper, were 38% higher on spruce trunks than on pine trunks. This suggests that pine-dominated territories with female-biased broods may have contained less food during the first broods. The observation was further supported by the fact that the feeding frequencies were lower in territories with high proportions of pines. In the second broods, territories with a high forest patch density produced female-biased broods, whereas high-quality territories with a large amount of deciduous trees and mixed forests produced male-biased broods. Our results suggest that habitat quality as measured by habitat characteristics is associated with sex allocation in free-living birds.     

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.     

Factors limiting the distribution of deciduous broadleaved trees in warm-temperate mountainous riparian forests

We examined differences in the responses of deciduous and evergreen broadleaved trees to fluvial disturbance and light environment near a river-facing forest edge in order to identify the factors limiting the distribution of deciduous trees, which are important components of warm-temperate mountainous riparian forests. Deciduous trees tended to be distributed on lower terraces of less than 2-m relative elevation from the water level, which corresponded with sites suffering from strong bank erosion due to high flood frequency. On the other hand, evergreen trees showed an opposite trend of distribution, indicating that high flood frequency associated with strong erosion-dominated soil disturbances might be a constraint for their establishment and/or survival. Furthermore, crowns of deciduous trees tended to be in the canopy layer at the forest edge, with fewer individuals observed beneath the canopy of evergreen trees, even at the forest edge. In contrast, evergreen trees were concentrated in the understory of the forest interior. These observations suggested that the light environment is the predominant factor affecting the establishment and survival of deciduous trees in warm-temperate mountainous riparian forests; however, the river-facing forest edge beneath the evergreen canopy is not a suitable habitat for deciduous trees. We concluded that erosion-dominated soil disturbance on lower terraces provides suitable habitats for deciduous trees by facilitating their reaching the canopy layer by limiting the establishment and/or survival of evergreen trees.

     

Timing of breeding and nestling diet of Wood Warbler Phylloscopus sibilatrix in relation to changing food supply

Capsule Wood Warblers did not match their reproduction to the caterpillar peak

Aims To study the timing of Wood Warblers’ breeding and nestling diet in relation to caterpillar abundance under primeval conditions in the Bia?owie?a National Park, Poland.

Methods Observations of food brought by parents. Inter‐year and seasonal changes in availability of folivorous caterpillars were assessed by direct counts and caterpillar frass collection.

Results Maximum food requirements occurred two weeks after the peak of caterpillar abundance. The mismatch had no effect on nestlings’ development. Diet varied little across years and habitats, but varied strongly within a season. Following the decline of ‘green’ caterpillars, their proportion in the Wood Warbler diet strongly declined. Caterpillars were replaced by winged insects. Small nestlings received more spiders than older ones.

Conclusion Timing of Wood Warbler breeding in Bia?owie?a National Park was constrained by the females’ arrival time. Birds did not match their reproduction to the caterpillar peak. ‘Green’ caterpillars were the preferred food for nestlings; birds responded to the caterpillar decrease by feeding smaller proportions of them and switching to alternative prey, namely winged insects.     

Tracing site‐specific isotopic signatures along a Blue Tit Cyanistes caeruleus food chain

Food chains culminating with temperate insectivorous passerines are well described, yet whether trophic webs can be site‐specific remains a largely unexplored question. In the case of site‐ or habitat‐specificity of food webs, stable isotope signatures of bird feathers may enable assignment of unmarked individuals to a site or a habitat of origin. We address this question in landscapes that include contrasting forest habitat patches with either deciduous Downy Oak Quercus humilis or evergreen Holm Oak Quercus ilex as dominant tree species. First, we examine the spatial variation across habitats and sites in the stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) along the oak leaf–Tortrix moth Tortrix viridana caterpillar–Blue Tit Cyanistes caeruleus food chain. Secondly, we assess whether the isotopic signatures allow for correct assignment of individual birds to their site or habitat of origin. At the scale of the landscape, stable isotope values enabled identification of the different components of the Blue Tit food chain: from oak leaves to Blue Tit nestlings and yearling birds. However, isotopic signatures were site‐specific (i.e. geographical) more than habitat‐specific (i.e. deciduous vs. evergreen oaks). Discriminant analyses correctly assigned 85% of nestlings and 83% of resident yearling birds, indicating a pronounced effect of site on Blue Tit feather isotopic signatures. We thereby demonstrate that isotopes reflect a stronger association of locally born birds to the local features of their habitat than that of un‐ringed yearling birds, whose plumage may have grown while in a wider geographical area. This study provides evidence of site‐specific isotopic signatures from oak leaves to Blue Tit feathers at a fine spatial scale.     

Food habits and the survival and growth of nestlings in two sympatric kites (Milvus milvus and Milvus migrans)

A comparative study of the Red Kite and Black Kite was conducted in the Doana National Park in southwestern Spain. Although the diets of both kites were similar, the Red Kite captured significantly more prey items from scrub habitats close to nesting areas than from marsh habitats, which were further away. Habitat differences in prey abundance and distance from breeding areas suggest that the Red Kite depended on a less abundant but more stable food supply. Nestling mortality from starvation was highest for the last hatched nestlings in both species. The growth rate of Black Kite nestlings was markedly higher than that of Red Kite nestlings. Moreover, growth rates increased with clutch size in the Red Kite, whereas they were constant in the Black Kite. Differences between species in reproductive traits are discussed mainly with regard to their feeding ecology. The slower growth and greater effect of sibling asymmetry on mortality of the Red Kite are consistent with our interpretation that the Red Kite's food supply is less abundant than the Black Kite's.