Family break-up in Black and Red Kites Milvus migrans and M. milvus: is time of independence an offspring decision?

I observed the natural process of family break-up in 13 families of Black Kites Milvus migrans and five families of Red Kites M. milvus in which fledglings had been individually marked. In other broods, I performed experiments which modified the parental investment fledglings received by supplementing nests of both species with food or transferring Black Kite chicks to nests with younger or older chicks of the same species.
The time of family break-up in the Black Kite is mainly an offspring decision which is not affected by an artificial increase of parental investment. The duration of the post-fledging period was not increased in Black Kites that were given supplementary food. Chicks transferred to nests with a younger chick did not extend the post-fledging period, nor did chicks transferred to nests with an older chick shorten the post-fledging period. In Red Kites, parental investment seemed to have more influence on the timing of the family break-up. Red Kite adults invested less as parents than did Black Kites during the post-fledging dependence period, and in nests where supplementary food was given, Red Kite fledglings stayed attached for a longer period.     

Geographic variation of the winter diet of the Red Kite Milvus milvus in the Iberian Peninsula

This paper presents data on the winter diet of the Red Kite Milvus milvus in the Iberian Peninsula. The diet composition and its relationship with food abundance and habitat characteristics were studied in two areas (northern and southern plateaux) with different food availability and habitat composition. Red Kites showed a clear difference in diet between the two plateaux, with a higher consumption of carrion on the northern plateau and of game species on the southern plateau. Common Voles Microtus arvalis , despite being a very abundant prey on the northern plateau, did not appear in the diet in high numbers, but their frequency was related to the proportion of land occupied by nonirrigated (cereal) crops. High frequencies of "small" carrion (e.g. farmed Rabbits Oryctolagus cuniculus , poultry Gallus sp.) in the diet were correlated with a decrease in the trophic diversity which suggested an active choice of the food, at least in years with low vole densities. We suggest that the use of carrion is influenced not only by its abundance but by its availability to Red Kites. Refuse tips and muladares (dumps for dead livestock) and their management during the winter are important for the conservation of the species.     

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.     

Successful breeding at one year of age by Red Kites Milvus milvus in southern England

This paper presents the first data on individually tagged male and female Red Kites Milvus milvus breeding at 1 year old and is validated by genetic fingerprinting. The data were collected during 1989 to 1994 when Red Kite nestlings were translocated to, and released in, southern England as part of a re-establishment project. A total of 23 1-year-old Red Kites attempted to breed in southern England from 1991 to 1994, of which three pairs, each composed of at least one yearling bird, reared five young. We conclude that habitat quality is ultimately the main influence that determines whether Red Kites can breed and how well they breed. However, individual experience, climate and competition with other Red Kites for nesting territories probably influence the age at which an individual can breed in a particular habitat.     

Intraspecific variation in prey quality: a comparison of nutrient presence in prey and nutrient extraction by predators

Prey quality can have large impacts on the survival, growth and behavior of predators. A number of studies have examined how different species of prey vary in quality. However, far less is known about intraspecific variation in the quality of prey for predators and even less about what nutrients are extracted from prey by predators. We examined how the sex, feeding level and developmental status of prey affected the quantities of nutrients present in prey bodies and the quantities of nutrients that could be extracted from prey by spiders. Female and well‐fed prey were larger and had more nutrients than male and food‐limited prey, respectively. After taking into account differences in prey size, spiders extracted relatively more lipid and less protein from female and well‐fed prey than from male and food‐limited prey, respectively. Mealworms were of higher quality than adult mealworm beetles; spiders were able to extract more lipid, protein and other nutrients from larvae than adults. While lipid present in prey was a good predictor of lipid consumed, protein present in prey was not a reliable predictor of protein consumed. The variation in prey quality that we observed within a single species of prey (i.e. well‐fed vs food‐limited crickets) was as large as variation in quality among the three species of prey used in these experiments. Intraspecific variation in prey quality may be an important factor affecting predatory arthropods, especially in habitats or at times of year when one species of prey is abundant. Further studies are needed to examine the consequences of intraspecific variation in prey quality on the life history and behavior of predators.     

The significance of seed food in chick development re‐evaluated by tracking day‐to‐day dietary variation in the nestlings of a granivorous passerine

The dietary adjustment of nestlings of granivorous birds to a seed diet and the different morphological characteristics of ingested food have rarely been examined in natural conditions. It has been suggested that the provision of cereal grains to nestlings of some seed‐eating bird species in modern agroecosytems is the result of poor food conditions after agricultural intensification. We analysed the abundance of invertebrate prey in the main foraging habitat of parent birds, daily changes (from hatching to fledging) in the efficiency of cereal seed digestion, and the dietary characteristics, diet composition and prey type delivered to nestlings of the Yellowhammer Emberiza citrinella. Analysis of faecal sacs from nests located in breeding habitat with an abundant invertebrate fauna revealed no relationship between the proportion of cereal seeds in the diet of nestlings and the food supply in the main foraging sites of the parents. Neonate nestlings (1 day old) exclusively received weakly chitinized invertebrate prey (arachnids and flies), but from the second day of life the nestlings were fed a variety of highly chitinized invertebrate prey, the percentage biomass of which did not change for the remainder of the nesting period. Cereal grains started to be delivered to 3‐day‐old nestlings and were already efficiently digested, and the percentage biomass of this food type increased progressively with nestling age. We suggest that the provisioning of cereal grains to nestlings is not forced by external factors, such as modern agricultural intensification; rather, it is an intentional behaviour of parent birds aimed at achieving physiological adjustment to seed food in the early stages of ontogeny.     

Stability in prey abundance may buffer Black Sparrowhawks Accipiter melanoleucus from health impacts of urbanization

As the global trend towards urbanization continues, the need to understand its impact on wildlife grows. Species may have different levels of tolerance to urban disturbance; some even appear to thrive in urban areas and use human‐subsidized resources. However, the physiological costs and trade‐offs faced by urban‐dwelling species are still poorly understood. We assess the evidence for a negative impact of urbanization on the Black Sparrowhawk Accipiter melanoleucus, a raptor that recently colonized Cape Town, South Africa, and explore the potential mechanisms behind any such effect. We predicted that birds in more urbanized areas may be in poorer health and that this may be partially driven by differences in prey quantity and quality along an urban habitat gradient. The health of Black Sparrowhawk nestlings was evaluated through measures of their physiological stress (heterophil/lymphocyte ratio), body condition and blood parasite infection (infection risk and intensity of Haemoproteus and Leucocytozoon). Diet composition was determined through an analysis of prey remains collected around nests, and prey abundance was determined through point counts in different habitat types. We could find no negative effects of urbanization on nestling health, with no significant relationships with heterophil/lymphocyte ratio, body condition, risk and intensity of infection by Haemoproteus or intensity of infection by Leucocytozoon. Risk of infection by Leucocytozoon did, however, decline with increasing urban cover, perhaps because urbanized areas contain less habitat for blackflies, the vectors of this parasite, which require moving fresh water. We found no change in diet breadth or composition with increasing urban cover. Although some prey species were abundant or less abundant in certain habitat types, all habitat types contained ample prey for Black Sparrowhawks. The widespread abundance of food resources and resulting lack of nutritional stress may explain why Black Sparrowhawks are seemingly free of the negative health impacts expected to arise from urbanization. These findings may explain the success of the species in Cape Town and suggest that for urban‐dwelling, bird‐eating raptors the abundance of prey in cities may override any potential negative impacts of urbanization on health due to disturbance or other sources of stress.     

Food acquisition by competing surfperch on a patchy environmental gradient

Synopsis Black surfperch, Embiotoca jacksoni, and striped surfperch, Embiotoca lateralis, coexisted along steep sloping rocky habitats at Santa Cruz Island, California. The range of depths occupied (to 15 m) was characterized by a strong gradient in abundance of prey and a changing mosaic of substrate types from which surfperch harvested food. Availability of prey and diversity of benthic substrates were greatest in shallowest areas and both declined with increasing depth. Individuals of both surfperch species were residential within a narrow range of depths, with the result that different segments of their populations were consistently exposed to different foraging environments. These two phenomena (residential behavior combined with a gradient in availability of resources) resulted in variation in foraging behaviors and diets among individuals that resided at different depths. The pattern of within-population variation differed between the surfperch species. Black surfperch individuals achieved similar taxonomic diets and expended similar foraging effort at all depths, but deep-water foragers captured much less prey biomass per unit effort. The taxonomic composition of striped surfperch diets differed among depths, and although similar amounts of prey biomass were captured everywhere, individuals in deep areas expended much greater effort to obtain that level of food return. For both species, habitat profitability (food return to foraging effort) declined with depth. The difference in habitat profitability appeared to influence fitness components of both surfperches. Individuals occupying deep habitats were about 5% shorter in standard length than conspecifics of the same chronological age living in shallow areas; the disparity in body size resulted in an estimated difference in clutch size of 10–18%.     

The diet of migrant Red Kites Milvus milvus during a Water Vole Arvicola terrestris outbreak in eastern France and the associated risk of secondary poisoning by the rodenticide bromadiolone

Poisoning by pesticides is considered one of the primary threats to the Red Kite Milvus milvus. We studied the diet of this raptor in an area of eastern France where the rodenticide bromadiolone is widely used to control Water Vole Arvicola terrestris outbreaks. A high degree of specialization for Water Voles was noted, as their remains were identified in all 119 pellets collected in autumn 2008, whereas other small rodent species and insects occurred in 27 and 9% of pellets, respectively. We estimated that Water Voles constituted 94% of the total biomass ingested by Red Kites under these conditions. Based on these data, the risk of secondary poisoning due to feeding on poisoned voles was assessed. Acute exposure on a single day was not considered a risk for Kites, but exposure to poisoned voles over 1 week represented the maximal risk for the Red Kite; the calculated dose of bromadiolone ingested by a Red Kite was 137 times higher than the toxicological benchmark for birds. A field survey in the studied area detected four dead Red Kites and one moribund bird in autumn 2008 but did not confirm that the cause of death was bromadiolone poisoning. We suggest that professional monitoring is needed to assess the impact of rodenticides on Red Kites in areas where voles are controlled.     

Storing prey in larders affects nestling haematological condition in the Red-backed Shrike (Lanius collurio)

Many animals, including shrikes (Laniidae), are known to store food. This behaviour ensures food availability during inclement weather, for rearing nestlings or signalling territory quality. Unlike most other shrike species, which accumulate food before breeding, Red-backed Shrikes Lanius collurio store food in larders, mainly during the nestling and fledgling periods. If this is an adaptive behaviour, variation in this behaviour should affect either or both the number and condition of nestlings, which may benefit from such larders. We assessed the haematological condition of Red-backed Shrike nestlings in east-central Poland in 2021 using a suite of blood parameters relating to sites where shrikes did or did not store food. Haemoglobin concentrations were positively correlated with larders, being significantly higher in nestlings of pairs that stored prey compared with those whose parents did not do so. White blood cell counts were lower in nestlings whose parents kept larders. Red blood cell counts and glucose concentrations in nestlings were not related to whether parents had larders. Food storage probably ensures a more constant access to food, which results in a superior haematological condition in nestlings.     

Importance of niche quality for Yellowhammer Emberiza citrinella nestling survival, development and body condition in its native and exotic ranges: the role of diet

The niche hypothesis predicts that some introduced species establish and spread successfully because their new environment provides expanded niche opportunities compared with their native environments. By investigating nestling survival, growth and body condition in relation to diet composition and prey abundance, we tested the prediction that the success of Yellowhammers Emberiza citrinella in New Zealand could be explained by the availability of better quality food resources in its introduced range compared with its native range. We found that Yellowhammer nestlings in New Zealand were larger and heavier than those in Britain, but that nestling survival and body condition did not differ between countries. The preferred prey items, which had some influence on nestling development and survival, were more prevalent in the diet of nestlings in New Zealand, suggesting that nestlings in the introduced range had a superior quality diet. However, there was little evidence to support the prediction that the preferred prey were more abundant in the introduced range, implying that some other factor such as prey accessibility or adult fitness may account for the differences in diet between countries.     

Prothonotary warbler nestling growth and condition in response to variation in aquatic and terrestrial prey availability

Aquatic prey subsidies entering terrestrial habitats are well documented, but little is known about the degree to which these resources provide fitness benefits to riparian consumers. Riparian species take advantage of seasonal pulses of both terrestrial and aquatic prey, although aquatic resources are often overlooked in studies of how diet influences the reproductive ecology of these organisms. Ideally, the timing of resource pulses should occur at the time of highest reproductive demand. This study investigates the availability of aquatic (mayfly) and terrestrial (caterpillar) prey resources as well as the nestling diet of the prothonotary warbler (Protonotaria citrea) at two sites along the lower James River in Virginia during the 2014 breeding season. We found large differences in availability of prey items between the two sites, with one having significantly higher mayfly availability. Nestling diet was generally reflective of prey availability, and nestlings had faster mean growth rates at the site with higher aquatic prey availability. Terrestrial prey were fed more readily at the site with lower aquatic prey availability, and at this site, nestlings fed mayflies had higher mean growth rates than nestlings fed only terrestrial prey. Our results suggest that aquatic subsidies are an important resource for nestling birds and are crucial to understanding the breeding ecology of riparian species.     

Supplementary feeding of the reintroduced Red Kite Milvus milvus in UK gardens

Capsule The provision of meat for garden birds is unusual in the UK but a reintroduced raptor, the Red Kite Milvus milvus, is now regularly fed in some areas. A questionnaire of garden kite feeders revealed that people were most often motivated to feed by a desire to see kites close up and that most provisioning falls within available guidelines. We estimated the median amount of food thought to be taken by kites per kite-feeding garden per day as 21?g, sufficient to support 0.12–0.26 individuals.     

Feeding ecology and life history variation of the blue tit in Mediterranean deciduous and sclerophyllous habitats

Summary High variation in laying date and clutch size of the blue tit between a Mediterranean mixed habitat on the mainland, southern France, and a sclerophyllous habitat on the island of Corsica is hypothesized to be related to differences in the food supply. The diet of the nestlings and feeding frequencies were studied using camera nestboxes and electronic chronographs. Food items brought to the nestlings were much more diverse on Corsica than on the mainland, including many fewer caterpillars and a wider range of taxa. However, when expressed as a volume index, prey items were on average larger on Corsica than on the mainland. Feeding frequencies were significantly lower on Corsica. A good correlation was found in both habitats between laying date and the caterpillar peak date, although both the leafing development of oaks and the peak of abundance of caterpillars occurred 3 weeks later in the Corsican sclerophyllous trees than in the mainland deciduous ones. Differences in the feeding ecology of tits between the two habitats are discussed in the light of the evergreen habit, which means that only 30% of leaves are available for phyllophagous insects instead of 100% in deciduous trees. the combination of a late and low food supply in evergreen trees is the best explanation for the differences in breeding traits betwen the two populations.     

Widespread supplementary feeding in domestic gardens explains the return of reintroduced Red Kites Milvus milvus to an urban area

Reintroductions are commonly used to mitigate biodiversity loss. One prominent example is that of the Red Kite Milvus milvus, a charismatic raptor of conservation concern. This species has been reintroduced across the UK over the last 25 years following its near extinction after centuries of persecution. The species was not expected to recolonize urban areas; its historical association with human settlements is attributed to scavenging on human waste and refuse, a resource now greatly reduced on the streets of modern European cities. However, the species has become a common daytime visitor to a large conurbation centred on the town of Reading, southern England, approximately 20 km from the first English reintroduction site. Given a near‐absence of breeding and roost sites, we investigated foraging opportunities and habitat associations that might explain use by Red Kites of this urban area. Surveys of discarded human foods and road‐kill suggested that these could support at most 13–29 Kites per day. Face‐to‐face surveys of a cross‐section of residents revealed that 4.5% (equivalent to 4349 households) provided supplementary food for Red Kites in their gardens. Using estimates of per‐household resource provision from another study, we calculated that this is potentially sufficient to feed 142–320 Kites, a substantial proportion of the total estimated to visit the conurbation each day (between 140 and 440). Road transects found positive associations between Red Kites and residential areas. We suggest that the decision made by thousands of householders to provide supplementary food for Red Kites in their gardens is the primary factor explaining their daytime abundance in this urban area.     

SYNCHRONOUS FLUCTUATIONS IN POPULATIONS OF SOME RAPTORS AND THEIR PREY

Raptorial birds which depend on a small number of prey species, especially those living in open habitats in the tundra, semi-desert and desert belts of the Palaearctic, undergo local fluctuations in numbers which are synchronous with the fluctuations in numbers of their main prey species. In this they contrast with the asynchronous or lagging type of predator-prey oscillation which has received wider attention in the literature.
Evidence from ringing recoveries shows that the species which fluctuate locally in synchrony with their prey tend to move greater distances from their birth-place or previous breeding place than those with more stable populations. The former species also tend to be split into fewer sub-specifically distinct forms than the latter. It is argued that the species which are subject to a fluctuating food supply have evolved, as an important adaptation, the ability to undertake more or less extensive 'searching migrations', which enable them to find and settle in areas of adequate food supply, often far removed from the previous breeding area. In the longer evolutionary aspect, this ability may be a factor promoting food specialization.
The synchronous population fluctuations of raptorial birds with their prey are compared with the asynchronous or lagging oscillations of carnivorous mammals.     

Nest orientation and hatching success of Black Kites Milvus migrans in Spain

An analysis of the orientation of 182 nest sites of the Black Kite Milvus migrans in two areas in south and central Spain found that orientations toward the east were preferred. In southern Spain an analysis of the relationship between nest orientation and hatching success showed that success was lower among the nests located in non-preferred orientations, with some inter-annual variation. The tendency to orientate nests toward the east was greater among the early nesting pairs and the effect that the orientation had on the hatching success was also greater than in late nesting pairs. The data suggest that rainfall and prevailing winds were the main factors conditioning nest orientation.     

Differential mortality of wintering shorebirds on the Banc d'Arguin, Mauritania, due to predation by large falcons

Predators may influence many aspects of the daily life and seasonal movements of their prey. Here we quantify direct, and evaluate indirect effects of predation by three falcon species (Lanner Falcon Falco biarmicus , Barbary Falcon Falco pelegrinoides and Peregrine Falcon Falco peregrinus ) on coastal shorebirds wintering on the Banc d'Arguin, Mauritania, an area hosting approximately 30% of the East Atlantic Flyway population of shorebirds. On the basis of 754 h of observation over five winters, 97 witnessed attacks and 585 collected prey remains, we show that shorebirds were safer in larger flocks, which tended to be attacked less often. Furthermore, species that forage relatively close to shore and in small flocks were depredated more often than expected from their relative abundance. In three species, Red Knot Calidris canutus canutus , Bar-tailed Godwit Limosa lapponica taymyrensis and Dunlin Calidris alpina , the juveniles were more vulnerable than adults. We estimated that on average 1% of the juvenile and 0.1% of the adult Red Knots present were killed by large falcons each winter. For Red Knots we simultaneously quantified annual survival on the basis of an individual colour-marking programme: mortality due to predation by falcons accounted for an estimated 6.2% (juveniles) and 0.8% (adults) of annual mortality. We suggest that juvenile Red Knots are 10 times as likely to be killed by falcons because they use riskier habitats, i.e. early and late tide foraging areas closer to shores where surprise attacks are both more common and more successful. These results indicate that the strength of indirect effects of predation operating in a shorebird population largely outweigh the effects of mortality per se .     

Supplementary feeding increases nestling feather corticosterone early in the breeding season in house sparrows

Several studies on birds have proposed that a lack of invertebrate prey in urbanized areas could be the main cause for generally lower levels of breeding success compared to rural habitats. Previous work on house sparrows Passer domesticus found that supplemental feeding in urbanized areas increased breeding success but did not contribute to population growth. Here, we hypothesize that supplementary feeding allows house sparrows to achieve higher breeding success but at the cost of lower nestling quality. As abundant food supplies may permit both high‐ and low‐quality nestlings to survive, we also predict that within‐brood variation in proxies of nestling quality would be larger for supplemental food broods than for unfed broods. As proxies of nestling quality, we considered feather corticosterone (CORT_f), body condition (scaled mass index, SMI), and tarsus‐based fluctuating asymmetry (FA). Our hypothesis was only partially supported as we did not find an overall effect of food supplementation on FA or SMI. Rather, food supplementation affected nestling phenotype only early in the breeding season in terms of elevated CORT_f levels and a tendency for more variable within‐brood CORT_f and FA. Early food supplemented nests therefore seemed to include at least some nestlings that faced increased stressors during development, possibly due to harsher environmental (e.g., related to food and temperature) conditions early in the breeding season that would increase sibling competition, especially in larger broods. The fact that CORT_f was positively, rather than inversely, related to nestling SMI further suggests that factors influencing CORT_f and SMI are likely operating over different periods or, alternatively, that nestlings in good nutritional condition also invest in high‐quality feathers.     

Snow tracking reveals different foraging patterns of red foxes and pine martens

Red fox (Vulpes vulpes) shares similar prey preferences and co-occurs with several other carnivores, and is together with pine marten (Martes martes), the most common mesocarnivore in the northern boreal forest. Voles are important prey for both species, but it is unclear to what extent they compete for the same food resources in winter. Here, we use 2139 km and 533 km of meticulous snow tracking of red foxes and pine martens to evaluate their food niches. We measured hunting and digging behaviour, whether successful or not, and the effect of snow depth and temperature. Pine martens were restricted to forested habitats, whereas red foxes used a wide range of habitats. Red foxes were found to dig more often than pine martens, 0.67 vs. 0.39 digging events per kilometre. Hunting was less common and similar in both species, about 0.1 hunting event per kilometre. Pine martens were more efficient in hunting and finding food remains compared to red foxes. Increasing snow depth reduced hunting success and also reduced dig success of red foxes. Food niche overlap was small. Red foxes used mostly voles and carrion remains of ungulates, whereas pine martens used cached eggs and small birds. We suggest that caching eggs is an important strategy for pine martens to survive winter in northern latitudes. Snow depth was important for capturing voles, and thick snow cover appeared to mask the effect of vole peaks. Intensified land use, as clear-cutting and leaving slaughter remains from harvest, will benefit red foxes on the expense of pine martens. The ongoing climate change with warmer winters and less snow will likely further benefit the red fox.