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.     

Age of first breeding, dispersal and survival of Red Kites Milvus milvus in Wales

Twenty-nine Red Kites, that were tagged as nestlings in Wales, bred for the first time at 2–7 years old (mean 3–6). A minimum of 41% of tagged young had entered the breeding population by the 7th year. Individuals moved up to 22 km between birthplace and breeding place and, having bred, most stayed in the same locality from year to year. No differences in these various respects were found between the sexes, though two long movements by territorial birds both involved females. Annual survival of adult breeders was estimated at around 95%.     

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.     

The function of aggressive chases by breeding Black and Red Kites Milvus migrans and M. milvus during the post-fledging dependence period

Black and Red Kites Milvus migrans and M. milvus chase other raptors approaching their nests. The study of this behaviour during the post-fledging period suggested that it reflects mainly, but not only, anti-predator behaviour. The frequency of vigilance and aggressive chases decreased through the post-fledging period as predicted by theoretical models of nest defence. Although predation risks were similar, Black Kites invested more time chasing intruders than did Red Kites. Black Kites, unlike Red Kites, chased away intruding juveniles, which may be interpreted as a behaviour to avoid investment in unrelated fledglings. Black Kites usually nest in loose colonies where the risk of, and selection pressures against, accidental investment in unrelated fledglings is likely to be greater than for Red Kites. Differences in aggressive chases by Black and Red Kites are better related to this than to different predation risks.     

Preen oil chemical composition encodes individuality,seasonal variation and kinship in black kites Milvus migrans

Evidence that bird odour can encode social information that can be used in chemical communication is growing, but is restricted to a few taxonomic groups. Among birds, diurnal raptors (i.e. birds from the Accipitriformes and Falconiformes order) have always been considered as mainly relying on their visual abilities. Although they seem to have a functional sense of smell, whether their odour can convey social information has yet to be determined. Combining gas‐chromatography‐mass‐spectrometry (GCMS) and microsatellite data, we tested whether chemical compounds from preen gland secretions can encode sex, age, individuality, seasonal differences and genetic relatedness in the gregarious accipitriform black kite Milvus migrans. While no differences in preen oil composition were found between age classes, an individual signature was detected. While a seasonal variation was found in both sexes, compounds differ between sexes in the non‐breeding season. Finally, a significant correlation between chemical proximity and genetic proximity was detected in male–male dyads and male–female dyads but not in female–female dyads. Our study provides the first evidence in raptors that preen secretion can convey information that may potentially be used in individual recognition, reproductive synchronization and inbreeding avoidance, and suggests that raptors may rely upon their olfactory abilities more than previously thought. This study opens promising avenues for further studies in raptor chemical communication.     

Growth of nestling black kites Milvus migrans: effects of hatching order, weather and season

The growth of black kites Milvus migrans was studied in 1981, 1982 and 1984 at the Doñana Biological Reserve (south-west Spain). Fifteen variables were submitted to a principal component analysis in order to characterize the growth pattern of the young. In general, in the nestlings that fledged, rapid weight gain corresponded to rapid growth of the tarsus and feathers. For nestlings that starved, tarsus growth took priority over weight gain. Within the same brood, the last-hatched sibling grew more slowly than its older sibling and the differences between them increased in proportion to the hatching asynchrony. Brood size did not affect growth as clearly as hatching order and hatching asynchrony. Increased rainfall and lower maximum temperature, factors that seem to make the search for food more difficult, had negative effects on the youngest sibling in a multiple brood and on single nestlings. The only environmental variable that had an effect on all of the nestlings was the hatching date.     

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.     

Poisoning of reintroduced red kites (<Emphasis Type="Italic">Milvus Milvus</Emphasis>) in England

Programmes to reintroduce predatory birds are resource intensive and expensive, yet there are few long-term studies on the health of these reintroduced birds following release. A total of 326 red kites (Milvus milvus) were released at four sites in England between 1989 and 2006 as part of efforts to reintroduce this species to England and Scotland, resulting in the establishment of several rapidly expanding populations in the wild. Detailed post-mortem examinations were carried out on 162 individuals found dead between 1989 and 2007, involving both released and wild-fledged birds. Toxicological analysis of one or more compounds was performed on 110 of the 162 birds. Poisoning was diagnosed in 32 of these 110 kites, 19 from second-generation anticoagulant rodenticides, 9 from other pesticides and 6 from lead. Criteria for diagnosing anticoagulant rodenticide poisoning included visible haemorrhage on gross post-mortem examination and levels of anticoagulant rodenticide exceeding 100 ng/g, but levels were elevated above 100 ng/g in a further eight red kites without visible haemorrhages, suggesting poisoning may have occurred in more birds. The anticoagulant rodenticides difenacoum and bromadiolone were the most common vertebrate control agents involved during this period. Poisoning of red kites may be slowing their rate of population recovery and range expansion in England. Simple modifications of human activity, such as best practice in rodent control campaigns, tackling the illegal use of pesticides and the use of non-toxic alternatives to lead ammunition, can reduce our impact on red kites and probably other populations of predatory and scavenging species.     

Relationship between laying dates of black kites Milvus migrans and spring temperatures in Italy: rapid response to climate change?

Many Holarctic bird species have been shown to be laying progressively earlier in response to global warming. Most studies have been conducted on small-bodied species at northern latitudes. In Italy, black kite Milvus migrans populations are concentrated near large pre-Alpine lakes, whose water temperatures have been increasing steadily in recent decades. Annual black kite laying dates have become 10–11 days earlier during the last nine years. This shift was related to temporal variations in spring air temperatures, with earlier laying in warmer springs. The progressive advancement in laying dates was not associated with an increase in population level productivity, despite the fact that productivity declined with laying date at the individual territory level within years. Avian response to climate change may be occurring more rapidly than previously thought, even in relatively long-lived species, and not only at the most northern latitudes or highest elevations. Because this change has occurred so rapidly, and because laying dates match annual variations in spring temperatures, the response must be facultatively driven by prevailing conditions, rather than genetic in response to natural selection.     

Year-to-year patterns of circulating levels of testosterone and corticosterone in relation to breeding density, experience, and reproductive success of the polygynous red-winged blackbird.

To determine across-year patterns in plasma testosterone (T) and corticosterone (B) levels in free-living birds, we took blood samples in the same 2-week breeding period during 4 consecutive years from territorial male red-winged blackbirds. We used our data to test predictions of the "challenge hypothesis" of T secretion and also to examine hormonal correlations with age, breeding experience, breeding density, and reproductive success. Average T and B levels across years were not significantly different. Within individuals, T levels between years were, in general, highly variable. T levels of males with territories in high-density breeding areas were significantly higher than those of males with territories in low-density areas. T levels were positively correlated with harem size and there was a trend for males with high T levels to fledge more offspring. We found no relationships between T and B levels and male age or breeding experience. Some results of the study were consistent with the predictions of the challenge hypothesis. This study constitutes one of the few examinations of across-year patterns in circulating hormone levels in wild populations.     

From individual behaviour to population pattern: weather-dependent foraging and breeding performance in black kites

The links between weather and animal behaviour in population processes have received relatively little attention. I studied the effect of weather conditions on foraging and breeding performance of a medium-sized raptor, the black kite, Milvus migrans. The frequency of prey capture attempts and their likelihood of success increased with temperature and declined with rainfall. Kites used flight styles involving a higher energy expenditure in less favourable weather. Nestling provisioning rates declined during rain spells. More kites hunted during periods of favourable weather and after periods with a high frequency of successful prey capture attempts by conspecifics; this result suggested that individuals may fine-tune their foraging effort to the expected foraging reward. Such compensatory behavioural adjustments may increase species resilience to climate change. The behaviourally mediated effects of weather on prey availability translated into population effects. Yearly weather conditions during the last stage of the prelaying period affected population-level productivity, probably through an effect on female body condition mediated by male provisioning capability and hunting yield. The predicted effects of climate change on kites may already be occurring, with progressively earlier laying and northward range expansion. These results confirm the need to pay greater attention to behaviourally mediated effects of climate on populations, particularly when individuals make compensatory adjustments that may enhance resilience to climate change. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Red kite (Milvus milvus) and black kite (M. migrans): minute genetic interspecies distance of two raptors breeding in a mixed community (Falconiformes: Accipitridae)

Allozyme analysis of 24 loci in 154 red kites (Milvus milvus) and 36 black kites (Milvus migrans) from die Hakel forest (Sachsen-Anhalt, Germany) revealed a Nei's interspecies genetic distance of D = 0.009. Of die observed genetic variance of four polymorphic enzymes, 15.4% referred to die differentiation between die kite species, but 84.6% were contributed by the ingroup polymorphism widiin diese species. Allozymes permit the identification of some 78% of samples as orignating from M. milvus , but only of 5.4% of samples obtained from M. migrans. Akhough the genetic distance is slight, die Milvus kites are valid biospecies which, despite occasional instances of hybridization, coexist sympatrically and may breed in mixed breeding aggregations. Mate choice in die largely separate winter quarters of diese migratory birds or chromosomal incompatibility are hypodietical isolation mechanisms stabilizing die species boundary. Moreover, the range sympatry could have developed fairly recendy widi die spread of human agriculture.     

Territorial occupancy and breeding performance in a migratory raptor do not follow ideal despotic distribution patterns

It has commonly been argued that many territorial species select their breeding sites following an ideal despotic distribution model, in which the most productive, high-quality territories are more frequently occupied. Theoretical and empirical studies have shown that this occupancy pattern may have population regulatory consequences, leading to density dependence in heterogeneous habitats. During a 9-year research project in a forested area of south-eastern Spain, we tested some of the predictions of the ideal despotic distribution model and the site-dependent population regulation theory in a migratory raptor species, the booted eagle Hieraaetus pennatus . Contrary to the predictions of the despotic model, our results showed that the temporal pattern of territorial occupation did not differ from randomness, and that the territory occupancy rate was not significantly related to the reproductive parameters considered. At population level, the breeding variables were density independent, suggesting the absence of site-dependent regulation. In addition, we were unable to find significant differences in the habitat characteristics between high-quality and low-quality territories, classified according to the criteria of both occupancy frequency and average productivity. Overall, our results suggest that booted eagles select their territories at random, probably due to the lack of strong environmental heterogeneity, and that occupancy rate is not a good measure of territory quality for the population studied.     

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.     

Space and habitat use by Red Kites Milvus milvus during winter in the Guadalquivir marshes: a comparison between resident and wintering populations

Differences in habitat selection, diet and behaviour of resident and wintering Red Kites Milvus milvus were studied in Donana National Park, southwest Spain. Adult resident Red Kites roosted at their nests, while immature residents and wintering birds gathered at communal roosts. Individuals remained on average for two consecutive nights (range 1–8) at the same communal roost. Wintering kites spent significantly more time foraging gregariously than residents. The use of the marsh was greater by wintering kites while residents more often used the forest. Wintering birds consumed more goose carrion than the residents. Wintering kites had larger core areas, moved farther from their roost sites to feed and changed foraging areas more frequently than residents. Among residents, adult females had the smallest home ranges (core area, distances travelled and time spent flying). We suggest that Red Kites wintering in Donana occupy the marsh because it provides high food availability, and is vacated by Black Kites M. migrans which exploit this profitable habitat during the breeding season.     

Effects of territorial intrusions, courtship feedings and mate fidelity on the copulation behaviour of the osprey

We studied copulation behaviour of the osprey, Pandion haliaetus, a semicolonial, fish-eating raptor, in Corsica (Mediterranean). Pairs copulated over a long period (45 days) and at a high rate, with, on average, 288 within-pair copulations (WPCs) for a clutch. Pairs breeding at higher density faced more frequent territorial intrusions than others and were potentially at an increased cuckoldry risk. However, and contrary to predictions of the ‘paternity assurance’ hypothesis for frequent copulations, we found that WPC rate decreased with increasing frequency of territorial intrusions. Male territory attendance increased with territorial intrusion frequency, to the detriment of the food provisioning of the female. Both attempted and successful WPC rates were positively related to the amount of food delivered by the male. Thus, the more frequent the territorial intrusions, the more time the male spent within his territory, the less he courtship fed and the smaller the fish he delivered, and the less the pair copulated successfully. WPC rate was also higher in newly formed pairs than in established pairs, and decreased with increasing pair bond length. The results suggest that males rely on mate guarding rather than frequent copulations to ensure paternity, and do not support the idea that sperm competition is the main cause of frequent WPCs. Nonfertilization functions of frequent copulations, such as pair bonding, mate assessment and mate retention, were likely early in the prelaying period. The findings that WPC rate decreased with mate fidelity and that females traded copulations for food suggest that mate retention was a possible function of frequent copulations in this species. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Home-range use, activity, and density of caracal in relation to prey density

Two male and three female caracal Felis caracal were radio-tracked over a 1-year period in arid shrub on the west coast of South Africa, by day and night over at least 130 days for each caracal, and uninterrupted for up to 120 h at a time. These results, on short-term use of space, were related to concurrent availability of prey. The use by caracal of specific plant communities showed a significant positive correlation to prey biomass of rodents. Males had much larger home-ranges (26.9±0.75 km²) than females (7.39±1.68 km²). Male home-ranges overlapped completely with those of females, whereas female ranges overlapped between 0 and 19%. Caracal were active by night and day; onset of activity was affected more by ambient temperature (T_A) than photoperiod. Caracal were active significantly longer on nights colder than 20°C. Females ceased activity at T_A > 20°C, males at T_A > 22°C. Males foraged faster than females (667 vs. 312 m h⁻¹) and moved more than twice the distance of females during an active period. Calculated density of caracal was between 0.23 and 0.47 km⁻².