Hacking techniques improve health and nutritional status of nestling White-tailed Eagles.

Birds of prey frequently feature in reintroductions and the hacking technique is typically used. Hacking involves removing large nestlings from donor populations, transferring them to captivity, feeding them ad libitum. Potentially, via the hacking method, the stress of captivity and disruption of parental feeding may be detrimental. Alternatively, the provision of ad libitum food may be advantageous. Although hacking has underpinned reintroduction project successes there has been no research on how the method may affect the health and nutritional status of translocated birds during captivity. We compared blood chemistry data from 55 young White-tailed Eagles, translocated from Norway as part of the species' reintroduction to Scotland, from sampling soon after arriving in captivity and again (≈42 days later) before their release. Numerous significant differences between the first and second samples were found, but no significant interactions showed that the sexes responded similarly to captivity. According to hematological and biochemical metrics, individuals showed several changes during captivity, including in red blood cell parameters, plasma proteins, and white cellular parameters related to the immune system, that indicated improved health status. Captivity with ad libitum food was associated with decreased urea and uric acid values: high values can indicate nutritional stress. Urea values became more normally distributed before release, indicating that ad libitum food had reduced nutritional differences between early nestlings in the season and later ones. Despite plentiful food, both sexes lost body mass before release, suggesting an inherent physiological mechanism to improve flight performance in fledglings. We conclude that hacking improved the health and nutritional status of released eagles which is likely to enable birds to cope with greater costs of exploratory behavior which they may require in reintroduction projects. In this context, we note the absence of survival differences between hacked and wild raptors in previous research.

Non-territorial GPS-tagged golden eagles Aquila chrysaetos at two Scottish wind farms: Avoidance influenced by preferred habitat distribution, wind speed and blade motion status.

Wind farms can have two broad potential adverse effects on birds via antagonistic processes: displacement from the vicinity of turbines (avoidance), or death through collision with rotating turbine blades. These effects may not be mutually exclusive. Using detailed data from 99 turbines at two wind farms in central Scotland and thousands of GPS-telemetry data from dispersing golden eagles, we tested three hypotheses. Before-and-after-operation analyses supported the hypothesis of avoidance: displacement was reduced at turbine locations in more preferred habitat and with more preferred habitat nearby. After-operation analyses (i.e. from the period when turbines were operational) showed that at higher wind speeds and in highly preferred habitat eagles were less wary of turbines with motionless blades: rejecting our second hypothesis. Our third hypothesis was supported, since at higher wind speeds eagles flew closer to operational turbines; especially-once more-turbines in more preferred habitat. After operation, eagles effectively abandoned inner turbine locations, and flight line records close to rotor blades were rare. While our study indicated that whole-wind farm functional habitat loss through avoidance was the substantial adverse impact, we make recommendations on future wind farm design to minimise collision risk further. These largely entail developers avoiding outer turbine locations which are in and surrounded by swathes of preferred habitat. Our study illustrates the insights which detailed case studies of large raptors at wind farms can bring and emphasises that the balance between avoidance and collision can have several influences.

Clinging on to alpine life: Investigating factors driving the uphill range contraction and population decline of a mountain breeding bird.

Climate change and anthropogenic nitrogen deposition are widely regarded as important drivers of environmental change in alpine habitats. However, due to the difficulties working in high-elevation mountain systems, the impacts of these drivers on alpine breeding species have rarely been investigated. The Eurasian dotterel (Charadrius morinellus) is a migratory wader, which has been the subject of uniquely long-term and spatially widespread monitoring effort in Scotland, where it breeds in alpine areas in dwindling numbers. Here we analyse data sets spanning three decades, to investigate whether key potential drivers of environmental change in Scottish mountains (snow lie, elevated summer temperatures and nitrogen deposition) have contributed to the population decline of dotterel. We also consider the role of rainfall on the species' wintering grounds in North Africa. We found that dotterel declines-in both density and site occupancy of breeding males-primarily occurred on low and intermediate elevation sites. High-elevation sites mostly continued to be occupied, but males occurred at lower densities in years following snow-rich winters, suggesting that high-elevation snow cover displaced dotterel to lower sites. Wintering ground rainfall was positively associated with densities of breeding males two springs later. Dotterel densities were reduced at low and intermediate sites where nitrogen deposition was greatest, but not at high-elevation sites. While climatic factors explained variation in breeding density between years, they did not seem to explain the species' uphill retreat and decline. We cannot rule out the possibility that dotterel have increasingly settled on higher sites previously unavailable due to extensive snow cover, while changes associated with nitrogen deposition may also have rendered lower lying sites less suitable for breeding. Causes of population and range changes in mountain-breeding species are thus liable to be complex, involving multiple anthropogenic drivers of environmental change acting widely across annual and migratory life cycles.

Sex-specific patterns of reproductive senescence in a long-lived reintroduced raptor.

For many species, there is evidence that breeding performance changes as an individual ages. In iteroparous species, breeding performance often increases through early life and is expected to level out or even decline (senesce) later in life. An individual's sex and conditions experienced in early life may also affect breeding performance and how this changes with age. Long-term monitoring of individuals from reintroduced populations can provide unique opportunities to explore age-related trends in breeding performance that might otherwise be logistically challenging. We used a unique dataset from a reintroduced population of white-tailed eagles Haliaeetus albicilla in Scotland, which has been intensively monitored since their initial reintroduction in 1975, to study age- and sex-specific trends in two measures of breeding performance. This monitoring provided data on the breeding performance of known individuals ranging in age from 3 to 26 years. We also explored changes in breeding performance in relation to early life experience (i.e., whether they were released or fledged in the wild). Breeding performance increased with age in early life in a similar manner for both sexes. We found stronger evidence for senescence in breeding performance in males than females. However, late-life female breeding success was associated with early life experience, while male senescent trends were not apparently impacted by conditions experienced during early life. Sexual differences in senescence mean that older males are less likely to breed successfully compared to older females, and this may influence females' mate changes later in life. This difference may suggest a linked sexual difference in survival rates or the possibility of proactive partner change by females in later life in this typically monogamous biparental species.

Ranging behaviour and habitat preferences of the Martial Eagle: Implications for the conservation of a declining apex predator.

Understanding the ranging behaviours of species can be helpful in effective conservation planning. However, for many species that are rare, occur at low densities, or occupy challenging environments, this information is often lacking. The Martial Eagle (Polemaetus bellicosus) is a low density apex predator declining in both non-protected and protected areas in southern Africa, and little is known about its ranging behaviour. We use GPS tags fitted to Martial Eagles (n = 8) in Kruger National Park (KNP), South Africa to describe their ranging behaviour and habitat preference. This represents the first time that such movements have been quantified in adult Martial Eagles. Territorial eagles (n = 6) held home ranges averaging ca. 108 km2. Home range estimates were similar to expectations based on inter-nest distances, and these large home range sizes could constrain the carrying capacity of even the largest conservation areas. Two tagged individuals classed as adults on plumage apparently did not hold a territory, and accordingly ranged more widely (ca. 44,000 km2), and beyond KNP boundaries as floaters. Another two territorial individuals abandoned their territories and joined the 'floater' population, and so ranged widely after leaving their territories. These unexpected movements after territory abandonment could indicate underlying environmental degradation. Relatively high mortality of these wide-ranging 'floaters' due to anthropogenic causes (three of four) raises further concerns for the species' persistence. Habitat preference models suggested Martial Eagles used areas preferentially that were closer to rivers, had higher tree cover, and were classed as dense bush rather than open bush or grassland. These results can be used by conservation managers to help guide actions to preserve breeding Martial Eagles at an appropriate spatial scale.

A balanced solution to the cumulative threat of industrialized wind farm development on cinereous vultures (Aegypius monachus) in south-eastern Europe.

Wind farm development can combat climate change but may also threaten bird populations' persistence through collision with wind turbine blades if such development is improperly planned strategically and cumulatively. Such improper planning may often occur. Numerous wind farms are planned in a region hosting the only cinereous vulture population in south-eastern Europe. We combined range use modelling and a Collision Risk Model (CRM) to predict the cumulative collision mortality for cinereous vulture under all operating and proposed wind farms. Four different vulture avoidance rates were considered in the CRM. Cumulative collision mortality was expected to be eight to ten times greater in the future (proposed and operating wind farms) than currently (operating wind farms), equivalent to 44% of the current population (103 individuals) if all proposals are authorized (2744 MW). Even under the most optimistic scenario whereby authorized proposals will not collectively exceed the national target for wind harnessing in the study area (960 MW), cumulative collision mortality would still be high (17% of current population) and likely lead to population extinction. Under any wind farm proposal scenario, over 92% of expected deaths would occur in the core area of the population, further implying inadequate spatial planning and implementation of relevant European legislation with scant regard for governmental obligations to protect key species. On the basis of a sensitivity map we derive a spatially explicit solution that could meet the national target of wind harnessing with a minimum conservation cost of less than 1% population loss providing that the population mortality (5.2%) caused by the operating wind farms in the core area would be totally mitigated. Under other scenarios, the vulture population would probably be at serious risk of extinction. Our 'win-win' approach is appropriate to other potential conflicts where wind farms may cumulatively threaten wildlife populations.

Food availability and predation risk, rather than intrinsic attributes, are the main factors shaping the reproductive decisions of a long-lived predator.

Deciphering the causes of variation in reproductive success is a fundamental issue in ecology, as the number of offspring produced is an important driver of individual fitness and population dynamics. Little is known, however, about how different factors interact to drive variation in reproduction, such as whether an individual's response to extrinsic conditions (e.g. food availability or predation) varies according to its intrinsic attributes (e.g. age, previous allocation of resources towards reproduction). We used 29 years of reproductive data from marked female tawny owls and natural variation in food availability (field vole) and predator abundance (northern goshawk) to quantify the extent to which extrinsic and intrinsic factors interact to influence owl reproductive traits (breeding propensity, clutch size and nest abandonment). Extrinsic and intrinsic factors appeared to interact to affect breeding propensity (which accounted for 83% of the variation in owl reproductive success). Breeding propensity increased with vole density, although increasing goshawk abundance reduced the strength of this relationship. Owls became slightly more likely to breed as they aged, although this was only apparent for individuals who had fledged chicks the year before. Owls laid larger clutches when food was more abundant. When owls were breeding in territories less exposed to goshawk predation, 99·5% of all breeding attempts reached the fledging stage. In contrast, the probability of breeding attempts reaching the fledging stage in territories more exposed to goshawk predation depended on the amount of resources an owl had already allocated towards reproduction (averaging 87·7% for owls with clutches of 1-2 eggs compared to 97·5% for owls with clutches of 4-6 eggs). Overall, our results suggested that changes in extrinsic conditions (predominantly food availability, but also predator abundance) had the greatest influence on owl reproduction. In response to deteriorating extrinsic conditions (fewer voles and more goshawks), owls appeared to breed more frequently, but allocated fewer resources per breeding attempt. However, intrinsic attributes also appeared to have a relatively small influence on how an individual responded to variation in extrinsic conditions, which indicates that owl reproductive decisions were shaped by a complex series of extrinsic and intrinsic trade-offs.

Age and sex-selective predation moderate the overall impact of predators.

Currently, there is no general agreement about the extent to which predators impact prey population dynamics and it is often poorly predicted by predation rates and species abundances. This could, in part be caused by variation in the type of selective predation occurring. Notably, if predation is selective on categories of individuals that contribute little to future generations, it may moderate the impact of predation on prey population dynamics. However, despite its prevalence, selective predation has seldom been studied in this context. Using recoveries of ringed tawny owls (Strix aluco) predated by 'superpredators', northern goshawks (Accipiter gentilis) as they colonized the area, we investigated the extent to which predation was sex and age-selective. Predation of juvenile owls was disproportionately high. Amongst adults, predation was strongly biased towards females and predation risk appeared to increase with age. This implies age-selective predation may shape the decline in survival with age, observed in tawny owls. To determine whether selective predation can modulate the overall impact of predation, age-based population matrix models were used to simulate the impact of five different patterns of age-selective predation, including the pattern actually observed in the study site. The overall impact on owl population size varied by up to 50%, depending on the pattern of selective predation. The simulation of the observed pattern of predation had a relatively small impact on population size, close to the least harmful scenario, predation on juveniles only. The actual changes in owl population size and structure observed during goshawk colonization were also analysed. Owl population size and immigration were unrelated to goshawk abundance. However, goshawk abundance appeared to interact with owl food availability to have a delayed effect on recruitment into the population. This study provides strong evidence to suggest that predation of other predators is both age and sex-selective and that selective predation of individuals with a low reproductive value may mitigate the overall impact of predators on prey population dynamics. Consequently, our results highlight how accounting for the type of selective predation occurring is likely to improve future predictions of the overall impact of predation.

When do young birds disperse? Tests from studies of golden eagles in Scotland.

BACKGROUND: Dispersal comprises three broad stages - departure from the natal or breeding locations, subsequent travel, and settlement. These stages are difficult to measure, and vary considerably between sexes, age classes, individuals and geographically. We used tracking data from 24 golden eagles, fitted with long-lived GPS satellite transmitters as nestlings, which we followed during their first year. We estimated the timing of emigration from natal sites using ten previously published methods. We propose and evaluate two new methods. The first of these uses published ranging distances of parents as a measure of the natal home range, with the requirement that juveniles must exceed it for a minimum of 10 days (a literature-based measure of the maximum time that a juvenile can survive without food from its parents). The second method uses the biggest difference in the proportion of locations inside and outside of the natal home range smoothed over a 30 day period to assign the point of emigration. We used the latter as the standard against which we compared the ten published methods. RESULTS: The start of golden eagle dispersal occurred from 39 until 250 days after fledging (based on method 12). Previously published methods provided very different estimates of the point of emigration with a general tendency for most to apparently assign it prematurely. By contrast the two methods we proposed provided very similar estimates for the point of emigration that under visual examination appeared to fit the definition of emigration much better. CONCLUSIONS: We have used simple methods to decide when an individual has dispersed - they are rigorous and repeatable. Despite one method requiring much more information, both methods provided robust estimates for when individuals emigrated at the start of natal dispersal. Considerable individual variation in recorded behaviour appears to account for the difficulty capturing the point of emigration and these results demonstrate the potential pitfalls associated with species exhibiting complex dispersal behaviour. We anticipate that coupled with the rapidly increasing availability of tracking data, our new methods will, for at least some species, provide a far simpler and more biologically representative approach to determine the timing of emigration.