Nest‐site characteristics,breeding success and competitive interactions between two recently sympatric apex predators

An influential period in avian life‐cycles is the annual breeding season, when competition over suitable nesting sites and territories is a key factor that can determine fitness and distribution, especially for species that are highly selective in their nesting habitats. We analysed nest‐site characteristics, breeding success and competitive interactions between two apex predator populations. Whereas the Short‐toed Eagle Circaetus gallicus has nested in the Judean Foothills (Israel) for a long time, the Long‐legged Buzzard Buteo rufinus has only invaded the nesting habitat of the Short‐toed Eagle during their breeding season in the last two decades. These two recently sympatric species have similar nesting ecology and frequently use the same nests. They are therefore expected to compete over nesting sites and territories. We analysed interspecific interactions between these two species by combining information from comprehensive observational, experimental, GIS analysis and remote sensing data, deriving 65 variables to characterize the nest‐sites used and the breeding success in 381 breeding attempts over four consecutive breeding seasons. To assess interspecific and intraspecific territorial behaviour and aggressiveness, stuffed Long‐legged Buzzards and Short‐toed Eagles were presented close to nests. Nest‐site characteristics overlapped substantially between species, and Long‐legged Buzzards occupied 21% of all Short‐toed Eagle nests. Intraspecific aggression rates among Long‐legged Buzzards were higher than their interspecific aggression rates with Short‐toed Eagles and also higher than intraspecific aggression among Short‐toed Eagles. Long‐legged Buzzard and Short‐toed Eagle breeding densities (1.59 ± 0.11 and 2.96 ± 0.11 pairs per 10 km², respectively) are likely to be the highest across their respective breeding distributions, with a maximum productivity of 0.96 ± 0.01 and 0.56 ± 0.05 (young fledged/breeding pair) for Long‐legged Buzzard and Short‐toed Eagle, respectively. Intraspecific interactions among both species play an important role in determining their breeding success and the spatial distribution of nesting sites. Our results suggest that interspecific competition over nesting sites and territories between both species, and the potential dominance of Long‐legged Buzzard, has both direct and indirect impacts on the spatial and demographic distribution of Short‐toed Eagles due to the recent establishment of Long‐legged Buzzard territories in the Judean breeding area.     

Spatial relationships and mechanisms of coexistence between dominant and subordinate top predators

Most forest ecosystems contain a diverse community of top‐level predators. How these predator species interact, and how their interactions influence their spatial distribution is still poorly understood. Here we studied interactions among top predators in a guild of diurnal forest raptors in order to test the hypothesis that predation among competing predators (intraguild predation) significantly affects the spatial distribution of predator species, causing subordinate species to nest farther away from the dominant ones. The study analyzed a guild in southwestern Europe comprising three raptor species. For 8 years we studied the spatial distribution of used nests, breeding phenology, intraguild predation, territory occupancy, and nest‐builder species and subsequent nest‐user species. The subordinate species (sparrowhawk Accipiter nisus) nested farther away from the dominant species (goshawk A. gentilis), which preyed on sparrowhawks but not on buzzards Buteo buteo, and closer to buzzards, with which sparrowhawks do not share many common prey. This presumably reflects an effort to seek protection from goshawks. This potential positive effect of buzzards on sparrowhawks may be reciprocal, because buzzards benefit from old sparrowhawk nests, which buzzards used as a base for their nests, and from used sparrowhawk nests, from which buzzards stole prey. Buzzards occasionally occupied old goshawk nests. These results support our initial hypothesis that interspecific interactions within the raptor guild influence the spatial distribution of predator species in forest ecosystems, with intraguild predation as a key driver. We discuss several mechanisms that may promote the coexistence of subordinate and dominant predators and the spatial assembly of this raptor guild: spatial refuges, different breeding phenology, spatial avoidance, low territory occupancy between neighboring nesting territories, nest concealment and protection, and diet segregation.     

Interactions with Great Skuas Stercorarius skua as a factor in the long‐term decline of an Arctic Skua Stercorarius parasiticus population

Many of the UK’s seabird species have displayed high variation in breeding success since the 1980s, largely due to changes in the availability of Lesser Sandeels Ammodytes marinus, their main prey. During this time, Arctic Skuas Stercorarius parasiticus experienced a rapid decline in the UK and the species has subsequently been placed on the Red List of birds of conservation concern. Although shortage of Lesser Sandeels is likely to be an influential factor, the Arctic Skua’s breeding range overlaps with that of the Great Skua Stercorarius skua, a larger bird with a more varied diet, and interspecific interactions for nesting habitat may exert an additional pressure on Arctic Skua breeding populations. Results from four censuses, spanning 21 years, were used to model habitat use and analyse distributional change in nesting Arctic Skuas at a major colony located on Fetlar, Shetland, Scotland. The decline in Arctic Skuas was not uniform across the island and competition with Great Skuas for nest‐sites appears to have influenced localized breeding distribution. By 2006, Arctic Skuas had been almost entirely excluded from shrub heath, blanket bog and coastal heath habitats, which were identified as preferred habitat in 1986. In 2006, Arctic Skua breeding territories were mainly restricted to one core area of preferred habitat where over 90% nested in high density as this habitat became increasingly occupied by Great Skuas. The more generalist foraging habit of the Great Skua allowed the population to grow rapidly as numbers of the more specialist Arctic Skua decreased during times of low sandeel availability. Our model suggests that both interspecific competition for territories with Great Skuas and food limitation have played important roles in the decline of Arctic Skuas on Fetlar.     

Reproductive responses of two related coexisting songbird species to environmental changes: global warming, competition, and population sizes

Comparative analyses of interspecific differences in response to climate change can provide important insights into the factors initiating seasonal onset of reproduction in various species and subsequent fitness consequences. We present a comparative analysis based on a 30-year breeding survey of two related migratory songbird species [Reed Warbler Acrocephalus scirpaceus (RW) and Great Reed Warbler A. arundinaceus (GRW)], which coexist in reedbeds by means of various interspecific interactions. The RW advanced breeding by 15 days and shortened its breeding time window, which is a combined effect of higher temperatures, alleviated competition as a consequence of population declines in the dominant GRW, and reduced RW population. Although the breeding period of GRW changed only slightly, its clutch initiation was likewise related to temperature. Most probably, advanced breeding in RW is favoured by changes in food supply and accelerated reed growth, which provides the necessary nest concealment earlier, whereas this does not affect the GRW, a species less vulnerable to nest predation. Clutch size decreased later in the season, so that earlier breeding produced a net increase in both species. An additional increase of clutch size in GRW can be explained by the use of higher-quality territories in today’s smaller population. The main causes of nest losses were predation in RW and adverse weather in GRW, but reproductive success increased over the study period in both species, which was a consequence of larger clutches in RW, but of favourable weather during rearing and fewer total losses in GRW. Our results document that different causal mechanisms are involved in the reproductive changes of the two congeneric species living in the same habitat: RW breeding earlier by making use of competitive release and other ecological improvements, and GRW by benefiting from better rearing conditions. As species respond differentially to climate change depending on ecosystem and biotic interactions, predictions of population dynamics will remain vague until the specific response mechanisms have been elucidated.     

Competitive interactions among raptors in boreal forests

We examined inter-specific interactions among goshawks (Accipiter gentilis), common buzzards (Buteo buteo) and honey buzzards (Pernis apivorus) in western Finland in 1983–1996. Because goshawks are among the largest birds of prey species in boreal forests they may take over the nest of smaller and less-competitive forest-dwelling raptors when searching for suitable places for breeding. Accordingly, more than half of newly established goshawk territories were found on the territories previously occupied by the common buzzard and the honey buzzard. Otherwise, territory sharing between these species was rare. Fledgling production of honey buzzards was not associated with the presence of goshawks, probably owing to the almost 2 months later onset of breeding. This probably decreases competitive interactions between these two species. An intensive interference competition, instead, seemed to be evident between common buzzards and goshawks, because the fledgling production of common buzzards was decreased by 20% as a result of failures during incubation and nestling period in the vicinity (<1 km) of occupied goshawk nests. Similarly, territory occupancy of common buzzards till the next breeding season was significantly reduced in the presence of goshawks. Relatively high proportions of occupied buzzard territories (17%) in the study area were shared by breeding goshawks on the same territory. This suggests that although their diets are dissimilar they inhabit similar habitats and might compete for the available prime nesting habitats within forest landscapes. In addition, goshawks benefit from taking over the complete nests of other raptors, imposing upon the original owners of the nest, because building a large stick nest is probably energetically costly. As a large raptor, the goshawk apparently has a competitive advantage over smaller ones, and may have an ever-increasing impact on smaller birds of prey, if there is a lack of sheltered forests inducing competition for the available nest sites.     

Investigations of density interactions among breeding birds in ponderosa pine forests: correlative and experimental evidence

Summary We assessed the importance of interspecific competition among insectivorous birds breeding in northcentral Arizona's ponderosa pine forests. We examined density interactions among species using two analytic approaches; correlative and experimental. The correlative approach examined patterns of change in breeding densities over four years at the community level and within two foraging guilds; picker-gleaners and aerial feeders. The relationships between morphological and behavioral similarity with pairwise density interactions were also assessed. Our experimental approach involved placement of nest boxes on two treatment plots to increase breeding densities of secondary cavity nesting birds that were in foraging guilds with open nesting insectivores.We found little evidence of interspecific competition. Patterns of density fluctuations indicated large positive covariances among species at both the community level and within guilds. Pairwise density interactions were independent of morphological or behavioral similarity. Nest boxes significantly increased breeding densities of the secondary cavity nesters. However, these increases did not induce reprocal density changes in the open nesting species. Interspecific competition for food during the breeding season appears to be unimportant in ponderosa pine bird communities.     

Interspecific aggression declines seasonally in breeding great tits Parus major

During the breeding season, great tits show aggression to protect their nest from intra‐ and interspecific intruders. Aggression is a labile trait that can be plastically expressed as a result of individual differences (e.g., personality), seasonal gradients in the costs and benefits of aggression, or other environmental components (e.g., number of competitors). Competitors may try to take over great tit nests, because the number of suitable nesting sites is limited, and great tits may guard high quality territories. Taking over a great tit nest may be especially fruitful in early phenological stages (egg laying) when great tits frequent their nests less often. However, great tits may compensate for this vulnerability by being more aggressive toward intruders during early nesting stages, a pattern that has already been established in an intraspecific context. Previous studies have shown that interspecific intruders were most likely to die from great tit aggression during great tit egg laying, suggesting great tits may also be more aggressive during this phase in an interspecific context. Here, I tested this hypothesis with simulated territorial intrusions in great tit territories using taxidermized blue tits Cyanistes caeruleus (hereafter called blue tit models). Great tit aggression (number of calls and approach distance toward blue tit model) was assayed during egg laying, incubation, and chick rearing in the breeding season of 2014. Although sample size was low due to a high fraction of non‐responders (n = 44 out of 89 assays across 26 out of 35 individuals), I found that great tits showed a seasonal decline in aggressiveness, which is congruent with intraspecific results on this study species. I discuss my findings in the context of differential adjustment to climate change between interspecific competitors.     

Interspecific interactions and co-existence in dabbling ducks: observations and an experiment

We studied the possible role of resource limitation and interspecific competition in assemblages of dabbling ducks on breeding lakes in Finland and Sweden with observational and experimental data. After initial vegetation mapping and yearly censuses of ducks in 1985–1990, we collected observational data in 1991–1994 from 28 lakes with natural populations of mallard Anas platyrhynchos and teal A. crecca. Mallard and teal co-occur over vast areas in the Holarctic and they are the only breeding dabbling ducks on many oligotrophic lakes. Both species are migratory in our study regions, teal arriving later in spring than mallards. Log-linear model analysis of observational presence/absence data revealed a positive, not a negative, association between the species. This association was independent of habitat diversity as well as of lake size. Mallard-teal interaction was also studied in a cross-over introduction experiment in 32 other lakes in two years. Wing-clipped mallards were introduced to breeding lakes before the arrival of teal to induce resource limitation and interspecific competition, hypothesized to reduce lake use by teal. The density of mallard pairs on experimental lakes was 2.9–8.0 times higher than on controls, but there was no negative response by teal to the treatment. This is the first combined observational-experimental demonstration of lack of interspecific competition in waterfowl. Our results indicate that heterospecific attraction may affect species co-existence in dabbling ducks. Received: 28 October 1996 / Accepted: 13 January 1997     

Demography and habitat availability in territorial occupancy of two competing species

Although metapopulation dynamics have become the focus of considerable theoretical research, little attention has been paid to its role when examining the coexistence of species. When two or more species live in the same patch network, interspecific interactions may affect their dispersal, colonization and extinction rates, and it may be possible to incorporate competition affecting these parameters in metapopulation models. Here, we extend the territorial occupancy model proposed by Lande to competing species. Our model estimates an equilibrium proportion of habitat occupancy as a function of life‐history parameters, dispersal behavior, habitat suitability and interspecific interactions. Moreover, it could prove to be useful as a tool in the assessment of potential management decisions. We apply the model to the golden Aquila chrysaetos and the Bonelli's eagle Hieraaetus fasciatus, two territorial raptors that coexist in the Mediterranean region, sharing food and nesting habitats. Over the last twenty years, while the golden eagle has maintained and, in some cases, increased its breeding numbers, Bonelli's eagle has suffered a marked decline, with many territories abandoned by the latter now occupied by the former. This suggests that the dynamics of these species could be influenced by interspecific competition. The model identified the relative importance of competition (stable equilibrium that allows long‐term coexistence) and predicted that, when habitat overlap is slight as in the study area, intraspecific dynamics are much more important for the persistence of each species than interspecific ones. Our results suggest that the improvement of territorial bird survival and productivity are the most urgently needed actions to be undertaken in the case of the golden eagle, while for Bonelli's eagle efforts should be focused on improving territorial and non‐territorial bird survival. As habitat conservation measures, the proportion of suitable exclusive habitat should be increased for both species.     

Does interference competition explain why White Terns of Aride Island,Seychelles, breed predominantly when marine productivity is lower?

White Terns Gygis alba breed throughout the year on Aride Island but show a marked preference to nest during the northwest monsoon (November–March), when interspecific competition and also marine productivity are considered to be lower. In this study we investigated and compared breeding and foraging parameters of White Terns between the northwest and southeast (May–September) monsoons to assess whether interspecific competition affected foraging activity and explained the timing of breeding of White Terns. Goatfish (Mullidae) dominated the diet of White Terns and no significant differences were found in diet composition between the northwest and southeast monsoons. Similar diets and patterns of food provisioning of White Terns between the two monsoons do not support the hypothesis of strong exploitation competition during the southeast monsoon. On the other hand, foraging behaviour and frequency of capture attempts of White Terns differed between the two monsoons. The percentages of multi‐species flocks with White Terns and of large flocks with more than 25 noddies Anous and terns Onychoprion were significantly higher during the southeast monsoon. Moreover, capture attempt frequencies of White Terns foraging in multi‐species flocks were lower than those of solitary individuals, and both solitary and flocking White Terns had significantly lower capture attempt frequencies during the southeast monsoon in comparison with the northwest monsoon. These results suggest that interference competition depresses foraging activity of flock‐foraging White Terns, particularly during the southeast monsoon, which could potentially explain why White Terns breed preferentially during the northwest monsoon. However, and despite avoiding interference competition, White Terns nesting during the northwest monsoon did not show higher breeding success than those nesting during the southeast monsoon. Other hypotheses that might explain the preference of White Terns for breeding during the northwest monsoon, such as avoidance of agonistic interactions with Lesser Noddies Anous tenuirostris at nest sites, should also be investigated.     

Nest site availability and niche differentiation between two cavity‐nesting birds in time and space

Niche differentiation is a key concept in the field of ecology and refers to the process by which competing species within an ecological community partition utilization of environmental resources to achieve coexistence. The existence of niche differentiation is uniquely difficult to prove on account of the fact that historical interaction among species, which plays a key role in elucidating the current state of coexistence among species, is not well known. We created continuous niche gradients in nest‐site resources between two sympatric secondary cavity‐nesting birds, the green‐backed tit (Parus monticolus) and the russet sparrow (Passer cinnamomeus), and investigated whether nesting site is a factor contributing to limiting breeding overlap by regular inspection and 388,160 min of film recording. Our results indicate that although we manipulated nest site availability to be uniformly high along the habitat gradient, the two bird species have little overlap in nest sites and rarely compete for them. Furthermore, the green‐backed tit possessed a wide range of fundamental niche that covered that of the russet sparrow, while their reproductive time was largely segregated. The sparrow was more aggressive and outcompeted the tit in their overlapped range. These results suggest that even though nesting sites are crucial to the reproduction of cavity‐nesting birds, some other factor plays a more important role in limiting niche overlap between sparrows and tits in space and time. Given that these two cavity‐nesting birds continued to use different habitats and breed in segregated time after our manipulation, their relationship is better explained by the ghost of competition past theory.     

Territory occupancy rate of goshawk and gyrfalcon: no evidence of delayed numerical response to grouse numbers

Two recent studies on territory occupancy rates of goshawk Accipiter gentilis and gyrfalcon Falco rusticolus report a 2–3-year-delayed numerical response to grouse numbers, which is a requirement for a hypothesis of predator-generated grouse cycles. The time lags were assumed to reflect the average age of sexual maturity in the raptor species. In southern Norway, however, subadult (two-year-old) goshawk hens occupied only 18–25% of territories where occupancy was not recorded in the preceding year, and there was no significant relationship between the proportion of subadults among recruits and grouse indices two years earlier. We argue that territory occupancy rates are not appropriate indices of total raptor population levels, but rather reflect the proportion of territorial pairs that attempt to nest. Because this depends on the body condition of the hens, fluctuations in other important winter resident prey species (most important for the goshawk) and winter weather (most important for the gyrfalcon) should also be addressed. During 1988–2006, the annual proportion of goshawk territories with recorded nesting attempts in southern Norway was most closely related to the preceding autumn’s population indices of black grouse Tetrao tetrix and mountain hare Lepus timidus, whereas the annual proportion of gyrfalcon territories with observations of falcons or with confirmed breeding attempts in central Norway were best explained by population indices of willow grouse Lagopus lagopus and ptarmigan L. mutus from the previous autumn, and by December temperatures. Hence, our studies do not support the predation hypothesis for grouse cycles.     

Male brood provisioning rates provide evidence for inter‐age competition for mates in female Cooper's Hawks Accipiter cooperii

Life history theory predicts that individuals should maximize lifetime reproductive success (LRS) by breeding as soon as they reach sexual maturity, yet many species delay breeding, either because there are insufficient available mates or breeding sites, or because delayed breeding yields higher LRS. Accipitriform species, such as Cooper's Hawk Accipiter cooperii, exhibit both delayed breeding and delayed plumage maturation. However, in certain circumstances, first‐year females in non‐definitive plumage do breed and apparently compete with older females for high‐quality breeding territories. We predicted that these young females are at a competitive disadvantage compared with older females and that older females would have both higher reproductive success and be able to acquire higher quality nesting territories. We conducted brood counts and measured prey delivery rates by male Cooper's Hawks in an expanding urban population located in Albuquerque, New Mexico (USA), to assess our prediction. We found that older females had higher reproductive success, fledging 1.6 more offspring than younger females, and that they occupied territories where males provisioned at higher rates of 0.37 more prey items per 2‐h period. Our results showed that older females fared better than first‐year females but it is unclear if this is the result of passive or active competition. Older females initiated nesting 14.3 days sooner than first‐year females and thus may have filled vacant, high‐quality territories before first‐year females began seeking mates. Additionally, first‐year females were never observed persistently to confront older females for breeding territories, but they did actively compete against each other. First‐year females may defer to older females who, in a direct competitive interaction, would be most likely to prevail. Thus, delayed plumage maturation in Cooper's Hawks may serve to focus competition for nesting territories within age classes.     

Bioko: critically important nesting habitat for sea turtles of West Africa

We evaluate the conservation status and threats faced by sea turtle nesting populations at Bioko Island, Equatorial Guinea (Central Africa). Beaches were monitored to obtain a detailed sea turtle nest census and, where possible, tagging of adult females was undertaken. Four sea turtle species were found nesting in the area: the green turtle (Chelonia mydas), the leatherback (Dermochelys coriacea), the olive ridley (Lepidochelys olivacea) and the hawksbill (Eretmochelys imbricata); with the former two species nesting in regionally important numbers. Nesting activity was concentrated between November and February, with a peak in December–January. Tagging and recapture of green turtles in two consecutive seasons suggested an estimated 560 (interquartile range: 420–1,681) and 414 (interquartile range: 190–1,255) nesting females in the area, respectively. Estimated numbers of nesting leatherbacks ranged from 123 to 215 and 243 to 293 in the first and second season, respectively. The other two species were less abundant (olive ridley: 19–29 and 28–43; hawksbill: 4–10 and 2 turtles). Data were compared with more recent surveys in the area and contextualised with information on human related threats. Despite the size of nesting stocks, ongoing permitted and illegal take of adult turtles at the nesting site constitutes a serious threat for these breeding aggregations. Additionally, tag returns from throughout the Gulf of Guinea suggest that the level of take in regional fisheries may also be a major threat.     

Niche Segregation Between Two Alpine Rosefinches: to Coexist in Extreme Environments

Alpine habitats are expected to support fewer, morphologically distinct congeners that diverge in niche more fully to facilitate stable coexistence. We tested this prediction with a granivorous avian guild consisting of only two Carpodacus members that bred in a Tibetan alpine zone at 4,200–4,600 m a.s.l. These two species, Streaked rosefinch C. rubicilloides weighing 40 g and Pink-rumped rosefinch C. eos 20 g, represent the extreme of body size of their genus. Both rosefinch species as the dominant members of local passerine breeders had similar breeding densities. They did not segregate in clutch initiation date, nesting altitude, habitat patches or nesting plant species, but in nesting plant height, with C. rubicilloides using those taller than the average of local bushes and C. eos using those lower than the average. No overlap in diet fed to nestlings between the two rosefinch species. C. rubicilloides with large and strong beaks primarily exploited the leguminous seeds, whereas C. eos mainly collected small seeds at ground level from a variety of plant species in accord with their smaller beaks. Interspecific aggression and territoriality were lack. The mechanisms underlying stable coexistence of the two rosefinch congeners should contribute to their success in the harsh abiotic environments.     

Ein Fall interspezifischer Kompetition bei Weberv?geln(Ploceus)

Summary A case of interspecific competition between the Spectacled Weaver(Ploceus ocularis) and the Vieillot's Black Weaver(P. nigerrimus), observed in Uganda, is described in detail.A pair of Spectacled Weavers had begun to construct a nest in a palm, when several male Black Weavers from a distant colony started to strip leaves of the palm for nesting material and also to rob the Spectacleds'nest. Soon several Black Weavers decided to make their own nests in the palm. One of these males instantly attacked the building pair of Spectacleds, and systematically removed their nest. Finally after a last aggressive attempt to reconquer their own nest the Spectacled Weavers disappeared, while the Black continued to build his nest unmolested.This is a case of displacement of an inferior species by a dominant one, without the slightest ecological need, which usually may be present in struggles for nesting holes in European passerines.     

Salt marsh elevation is a strong determinant of nest‐site selection by Clapper Rails in Georgia,USA

As sea levels rise, birds nesting in coastal marshes will be particularly vulnerable to increased tidal inundation. Understanding how marsh birds select their nesting habitat along the elevational gradient of these marshes will provide insight into how these species might be affected by rising sea levels. Clapper Rails (Rallus crepitans) are coastal marsh‐nesting birds whose nests are vulnerable to flooding, but it is not clear if they select for habitat along the elevational gradient or only use other habitat cues. Our objective was to determine if Clapper Rails select higher‐elevation nest sites, while also controlling for selection of other habitat variables at both landscape and territory levels, by comparing nest habitat to habitat in other areas of territories and at random points in the marsh landscape. At the landscape level, Clapper Rails did not exhibit selection for the elevational gradient, with nests and random points at similar elevations. At the territory level, however, nest‐site selection was most influenced by elevation and plant height, with Clapper Rails selecting nest sites with higher elevations and in areas with taller plants. However, the strength of the elevation effect was uncertain, suggesting the importance of precise elevation measurements in the field. Given this selection for higher‐elevation nest sites, Clapper Rails may be somewhat resilient to increased tidal inundation. However, the potential for increased intra‐ and interspecific competition for high‐elevation marshes should make conservation of these habitats a priority.     

Possible functions of alternative nests in raptors: the case of Bonelli’s Eagle

Four non-exclusive hypotheses have been proposed to explain the alternative nest-building behaviour of raptors: (1) nests as an advertising signal in territories, (2) frustration nests, (3) competition avoidance by nest-site and (4) reduction of nest ectoparasites. We report here data collected during an observational study of a population of Bonelli’s Eagle Hieraaetus fasciatus in southeastern Spain. Our data does not support the first two hypotheses based on the closeness of nests within territories, the lack of correlation between the number of nests and the distance to neighbours and the construction of secondary nests at similar frequencies after breeding failures as after successes. In contrast, the construction of alternative nests to avoid competition with other cliff-nesting raptor species breeding nearby was clearly important since 30% of the pairs obtained some direct benefit from the existence of alternative nests within their territories. We also found abundant Coleoptera (Dermestidae) and Diptera (Carnidae, Calliphoridae and Phoridae) as ectoparasites in nests, and the alternative use of nests, synergetically with the presence of greenery as mechanisms for avoiding ectoparasites, was an important factor for the breeding success of the eagle. Our data suggest that competition avoidance and, in particular, the reduction of nest ectoparasite hypotheses are the more plausible explanations for the maintenance of alternative nests in raptors.     

Interactions between common buzzard Buteo buteo and goshawk Accipiter gentilis: trade-offs revealed by a field experiment

I examined the behavioural interactions between common buzzard Buteo buteo and goshawk Accipiter gentilis and their effects on buzzard breeding success and brood defence with a two-year field experiment using dummies and playback calls. A priori I showed through an extensive nest site analysis that there is considerable nesting habitat overlap between the two species and hence potential for interspecific competition for prime nesting habitat. Buzzards had a significantly lower breeding success when presented with a goshawk dummy compared to control broods but there was no effect of buzzard dummies on reproductive success. Buzzards failing with their breeding attempt tended to select another nest site while successful buzzards more frequently used the same nest again. Buzzard pairs were less often attacked by common crows Corvus corone while exposed to goshawk dummies compared to buzzard dummies. The decision to desert a nest seems to be a trade-off between predation risk on the one hand and protection against crows on the other. Goshawks proved far more aggressive against an intraspecific dummy than buzzards. Buzzards adjusted their level of brood defence against both intra- and interspecific dummies according to the age of offspring but not offspring number, with an increasing brood defence level with increasing offspring age. Thus the behaviour of buzzards towards goshawks is a result of a complex system of trade-offs between predation risk, competition for prime nesting habitat and protection from crows on which brood value acts as a temporal modifier.     

Green plant material versus ectoparasites in nests of Bonelli's eagle

The greenery and arthropod nest composition has been studied in a population of Bonelli's eagle Hieraaetus fasciatus in south-east Spain, relating them to the plant availability within territories and breeding success of pairs. Greenery was invariably from trees and shrubs, with pine and oak species accounting for 78% of the nest composition in weight. All eagle pairs with Pinus pinaster availability in territories actively selected it for nest greenery. This pine species is characterized by a high level of aromatic compounds, particularly β -pinene, highly repellent for insects. The amount of pine greenery in the nest was correlated with a lower presence of ectoparasites in that nest (blow fly larvae, Protocalliphora) , and higher breeding success of pairs. We discuss whether a coevolutionary process between parasites and their hosts has guided a particular nesting strategy of the eagles to improve their breeding success.