Pleiotropic effects of melanin pigmentation: haemoparasite infection intensity but not telomere length is associated with plumage morph in black sparrowhawks.

There is increasing recognition of the potential pleiotropic effects of melanin pigmentation, particularly on immunity, with reports of variation in haemoparasite infection intensity and immune responses between the morphs of colour-polymorphic bird species. In a population of the black sparrowhawk (Accipiter melanoleucus) in western South Africa, light morphs have a higher haemoparasite infection intensity, but no physiological effects of this are apparent. Here, we investigate the possible effects of haemoparasite infection on telomere length in this species and explore whether relative telomere length is associated with either plumage morph or sex. Using quantitative polymerase chain reaction analysis, we confirmed that dark morphs had a lower haemoparasite infection intensity than light morphs. However, we found no differences in telomere length associated with either the haemoparasite infection status or morph in adults, although males have longer telomeres than females. While differences in haemoparasite intensity between morphs are consistent with pleiotropic effects of melanin pigmentation in the black sparrowhawk, we found no evidence that telomere length was associated with haemoparasite infection. Further work is needed to investigate the implications of possible pleiotropic effects of plumage morph and their potential role in the maintenance of colour polymorphism in this species.

African savanna raptors show evidence of widespread population collapse and a growing dependence on protected areas.

The conversion of natural habitats to farmland is a major cause of biodiversity loss and poses the greatest extinction risk to birds worldwide. Tropical raptors are of particular concern, being relatively slow-breeding apex predators and scavengers, whose disappearance can trigger extensive cascading effects. Many of Africa's raptors are at considerable risk from habitat conversion, prey-base depletion and persecution, driven principally by human population expansion. Here we describe multiregional trends among 42 African raptor species, 88% of which have declined over a ca. 20-40-yr period, with 69% exceeding the International Union for Conservation of Nature criteria classifying species at risk of extinction. Large raptors had experienced significantly steeper declines than smaller species, and this disparity was more pronounced on unprotected land. Declines were greater in West Africa than elsewhere, and more than twice as severe outside of protected areas (PAs) than within. Worryingly, species suffering the steepest declines had become significantly more dependent on PAs, demonstrating the importance of expanding conservation areas to cover 30% of land by 2030-a key target agreed at the UN Convention on Biological Diversity COP15. Our findings also highlight the significance of a recent African-led proposal to strengthen PA management-initiatives considered fundamental to safeguarding global biodiversity, ecosystem functioning and climate resilience.

Interactive effects of rising temperatures and urbanisation on birds across different climate zones: A mechanistic perspective.

Climate change and urbanisation are among the most pervasive and rapidly growing threats to biodiversity worldwide. However, their impacts are usually considered in isolation, and interactions are rarely examined. Predicting species' responses to the combined effects of climate change and urbanisation, therefore, represents a pressing challenge in global change biology. Birds are important model taxa for exploring the impacts of both climate change and urbanisation, and their behaviour and physiology have been well studied in urban and non-urban systems. This understanding should allow interactive effects of rising temperatures and urbanisation to be inferred, yet considerations of these interactions are almost entirely lacking from empirical research. Here, we synthesise our current understanding of the potential mechanisms that could affect how species respond to the combined effects of rising temperatures and urbanisation, with a focus on avian taxa. We discuss potential interactive effects to motivate future in-depth research on this critically important, yet overlooked, aspect of global change biology. Increased temperatures are a pronounced consequence of both urbanisation (through the urban heat island effect) and climate change. The biological impact of this warming in urban and non-urban systems will likely differ in magnitude and direction when interacting with other factors that typically vary between these habitats, such as resource availability (e.g. water, food and microsites) and pollution levels. Furthermore, the nature of such interactions may differ for cities situated in different climate types, for example, tropical, arid, temperate, continental and polar. Within this article, we highlight the potential for interactive effects of climate and urban drivers on the mechanistic responses of birds, identify knowledge gaps and propose promising future research avenues. A deeper understanding of the behavioural and physiological mechanisms mediating species' responses to urbanisation and rising temperatures will provide novel insights into ecology and evolution under global change and may help better predict future population responses.

A utilization distribution for the global population of Cape Vultures (Gyps coprotheres) to guide wind energy development.

The rapid development of wind energy in southern Africa represents an additional threat to the already fragile populations of African vultures. The distribution of the vulnerable Cape Vulture Gyps coprotheres overlaps considerably with wind energy development areas in South Africa, creating conflicts that can hinder both vulture conservation and sustainable energy development. To help address this conflict and aid in the safe placement of wind energy facilities, we map the utilization distribution (UD) of this species across its distributional range. Using tracking data from 68 Cape Vultures collected over the last 20 years, we develop a spatially explicit habitat use model to estimate the expected UDs around known colonies. Scaling the UDs by the number of vultures expected to use each of the colonies, we estimate the Cape Vulture population utilization distribution (PUD) and determine its exposure to wind farm impacts. To complement our results, we model the probability of a vulture flying within the rotor sweep area of a wind turbine throughout the species range and use this to identify areas that are particularly prone to collisions. Overall, our estimated PUD correlates well with reporting rates of the species from the Southern African Bird Atlas Project, currently used to assess potential overlap between Cape Vultures and wind energy developments, but it adds important benefits, such as providing a spatial gradient of activity estimates over the entire species range. We illustrate the application of our maps by analyzing the exposure of Cape Vultures in the Renewable Energy Development Zones (REDZs) in South Africa. This application is a scalable procedure that can be applied at different planning phases, from strategic, nationwide planning to project-level assessments.

A global review of the temporal and spatial patterns of DDT and dieldrin monitoring in raptors.

Concentrations of organochlorine pesticides have been extensively monitored in birds, particularly from higher trophic guilds such as raptors. While monitoring of raptors has been ongoing for decades, patterns from monitoring activities have never been summarised on a global scale. In this study, we undertake a review to better describe the monitoring of two widespread organochlorine pesticides monitored globally in raptors, DDT and dieldrin. We provide a historical retrospective on the monitoring effort of a global environmental issue. Sampling was heavily biased geographically to the global north, with more than 90 % of studies conducted in this socio-geographic region, most from Europe and North America. Although monitoring occurred from at least 114 species, most samples came from relatively few species, with three species (Eurasian Sparrowhawk Accipiter nisus, Bald Eagle Haliaeetus leucocephalus, and Peregrine Falcon Falco peregrinus) comprising 50 % of samples. The types of raptors sampled have changed over time, with avian and mammal specialists dominating samples until the 1970s, but more diverse dietary guilds monitored in later decades, and greater proportions of samples coming from generalist species. The three most sampled tissues (egg, liver, and plasma) comprised 84 % of all samples. Eggs were the earliest tissue examined and the only tissue sampled in all decades. The geographical bias in monitoring effort and relatively narrow species focus, suggests that patterns in these pesticides are unlikely to be fully representative of all global environments occupied by raptors. While DDT has been banned throughout most of the global north, it remains in use in the global south, yet monitoring effort in the south, does not match that of the north. While monitoring remains prevalent in the global north, contemporary monitoring is limited in the global south with less than 10 % of raptors sampled in Asia, Africa, and South America, over the last 3 decades.

Innate immune function and antioxidant capacity of nestlings of an African raptor covary with the level of urbanisation around breeding territories.

Urban areas provide breeding habitats for many species. However, animals raised in urban environments face challenges such as altered food availability and quality, pollution and pathogen assemblages. These challenges can affect physiological processes such as immune function and antioxidant defences which are important for fitness. Here, we explore how levels of urbanisation influence innate immune function, immune response to a mimicked bacterial infection and antioxidant capacity of nestling Black Sparrowhawks Accipiter melanoleucus in South Africa. We also explore the effect of timing of breeding and rainfall on physiology since both can influence the environmental condition under which nestlings are raised. Finally, because urbanisation can influence immune function indirectly, we use path analyses to explore direct and indirect associations between urbanisation, immune function and oxidative stress. We obtained measures of innate immunity (haptoglobin, lysis, agglutination, bactericidal capacity), indices of antioxidant capacity (total non-enzymatic antioxidant capacity (tAOX) and total glutathione from nestlings from 2015 to 2019. In addition, in 2018 and 2019, we mimicked a bacterial infection by injecting nestlings with lipopolysaccharide and quantified their immune response. Increased urban cover was associated with an increase in lysis and a decrease in tAOX, but not with any of the other physiological parameters. Furthermore, except for agglutination, no physiological parameters were associated with the timing of breeding. Lysis and bactericidal capacity, however, varied consistently with the annual rainfall pattern. Immune response to a mimicked a bacterial infection decreased with urban cover but not with the timing of breeding nor rainfall. Our path analyses suggested indirect associations between urban cover and some immune indices via tAOX but not via the timing of breeding. Our results show that early-life development in an urban environment is associated with variation in immune and antioxidant functions. The direct association between urbanisation and antioxidant capacity and their impact on immune function is likely an important factor mediating the impact of urbanisation on urban-dwelling animals. Future studies should explore how these results are linked to fitness and whether the responses are adaptive for urban-dwelling species.

Using web-sourced photographs to examine temporal patterns in sex-specific diet of a highly sexually dimorphic raptor.

Traditional methods to study raptor diet are usually limited temporally, e.g. prey remains at nesting sites, and are unsuitable to examine dietary changes throughout the year. Using web-sourced photography, we explore temporal patterns in prey size and key prey species between sexes of the sexually dimorphic Eurasian sparrowhawk (Accipiter nisus) throughout the United Kingdom. We examined 666 photographs of sparrowhawk on prey and identified the prey species involved, together with sparrowhawk sex. Changes in prey size and proportions of key prey species over time (seasonally and monthly) were explored for each sex. Prey weight was substantially higher for females than males. However, on average, prey size for both sexes declined during the summer period (May-June) being the lowest in June, which is the main nestling-rearing month for both sparrowhawks and their prey. Compared with summer, rock doves (Columba livia) were more important prey for female sparrowhawk in winter. Whereas, for males, Eurasian blackbirds (Turdus merula) were more important in spring compared with autumn. Web-sourced photography can overcome several limitations of previous methods used to study raptor diet including the ability to quantify diet between the sexes throughout the entire year, however, may also introduce a prey-size bias toward larger prey items.

Causes, temporal trends, and the effects of urbanization on admissions of wild raptors to rehabilitation centers in England and Wales.

Data from wildlife rehabilitation centers (WRCs) can provide on-the-ground records of causes of raptor morbidity and mortality, allowing threat patterns to be explored throughout time and space. We provide an overview of native raptor admissions to four WRCs in England and Wales, quantifying the main causes of morbidity and mortality, trends over time, and associations between threats and urbanization between 2001 and 2019. Throughout the study period, 14 raptor species were admitted totalling 3305 admission records. The Common Buzzard (Buteo buteo; 31%) and Tawny Owl (Strix aluco; 29%) were most numerous. Relative to the proportion of breeding individuals in Britain and Ireland, Peregrine Falcons (Falco peregrinus), Little Owls (Athene noctua), and Western Barn Owls (Tyto alba) were over-represented in the admissions data by 103%, 73%, and 69%, respectively. Contrastingly Northern Long-eared Owls (Asio otus), Western Marsh Harriers (Circus aeruginosus), and Merlin (Falco columbarius) were under-represented by 187%, 163%, and 126%, respectively. Across all species, vehicle collisions were the most frequent anthropogenic admission cause (22%), and orphaned young birds (10%) were most frequent natural cause. Mortality rate was highest for infection/parasite admissions (90%), whereas orphaned birds experienced lowest mortality rates (16%). For one WRC, there was a decline in admissions over the study period. Red Kite (Milvus milvus) admissions increased over time, whereas Common Buzzard and Common Kestrel admissions declined. There were significant declines in the relative proportion of persecution and metabolic admissions and an increase in orphaned birds. Urban areas were positively associated with persecution, building collisions, and unknown trauma admissions, whereas vehicle collisions were associated with more rural areas. Many threats persist for raptors in England and Wales, however, have not changed substantially over the past two decades. Threats associated with urban areas, such as building collisions, may increase over time in line with human population growth and subsequent urban expansion.

Pulses of anthropogenic food availability appear to benefit parents, but compromise nestling growth in urban red-winged starlings.

The provision of anthropogenic food undoubtedly influences urban bird fitness. However, the nature of the impact is unclear, with both benefits and costs of urban diets documented. Moreover, the influence of short-term fluctuations in food availability, linked to urban weekday/weekend cycles of human presence, is largely unknown. We explored whether breeding red-winged starlings Onychognathus morio in Cape Town, South Africa, altered foraging and provisioning behaviour between days with high human presence (HHP) and days with low human presence (LHP)-i.e. weekdays versus weekends and vacation days. We investigated the relationship between starling diet, adult body mass and nestling development. Breeding adults consumed and provisioned the same quantity of food, but a significantly greater proportion of anthropogenic food on HHP compared to LHP days. Adults apparently benefited from the anthropogenic diet, experiencing significantly greater mass gain on HHP days. However, nestlings experienced a cost, with the number of HHP days during the nestling period associated negatively with nestling size. Adults may, therefore, benefit from the high calorie content of anthropogenic food, while nestlings may be negatively affected by nutrient limitation. The quantity of food available in urban environments may, therefore, benefit adult survival, while its quality imposes a cost to nestling growth.

Changes in surface water drive the movements of Shoebills.

Animal movement is mainly determined by spatial and temporal changes in resource availability. For wetland specialists, the seasonal availability of surface water may be a major determinant of their movement patterns. This study is the first to examine the movements of Shoebills (Balaeniceps rex), an iconic and vulnerable bird species. Using GPS transmitters deployed on six immature and one adult Shoebills over a 5-year period, during which four immatures matured into adults, we analyse their home ranges and distances moved in the Bangweulu Wetlands, Zambia. We relate their movements at the start of the rainy season (October to December) to changes in Normalized Difference Water Index (NDWI), a proxy for surface water. We show that Shoebills stay in the Bangweulu Wetlands all year round, moving less than 3 km per day on 81% of days. However, average annual home ranges were large, with high individual variability, but were similar between age classes. Immature and adult Shoebills responded differently to changes in surface water; sites that adults abandoned became drier, while sites abandoned by immatures became wetter. However, there were no differences in NDWI of areas used by Shoebills before abandonment and newly selected sites, suggesting that Shoebills select areas with similar surface water. We hypothesise that the different responses to changes in surface water by immature and adult Shoebills are related to age-specific optimal foraging conditions and fishing techniques. Our study highlights the need to understand the movements of Shoebills throughout their life cycle to design successful conservation actions for this emblematic, yet poorly known, species.

Malar stripe size and prominence in peregrine falcons vary positively with solar radiation: support for the solar glare hypothesis.

Many falcons (Falco spp.) exhibit a distinct dark plumage patch below the eye, termed the malar stripe. This stripe is hypothesized to reduce the amount of solar glare reflected into the eyes while foraging, thereby increasing hunting efficiency in bright conditions. Here, we use a novel, global-scale correlative approach to test this 'solar glare hypothesis' in peregrine falcons (Falco peregrinus), the most widespread falcon species, using web-sourced photographs from across the species' global range. We found that the size and prominence of the malar stripe were positively associated with average annual solar radiation, but not with other environmental variables, such as temperature and rainfall. Our results provide the first published evidence for the hypothesis that this plumage feature functions to reduce the amount of solar glare reflected into the falcon's eyes, thereby improving the ability to pinpoint and target agile prey in bright conditions.

Parental morph combination does not influence innate immune function in nestlings of a colour-polymorphic African raptor.

Conditions experienced during early life can have long-term individual consequences by influencing dispersal, survival, recruitment and productivity. Resource allocation during development can have strong carry-over effects onto these key parameters and is directly determined by the quality of parental care. In the black sparrowhawk (Accipiter melanoleucus), a colour-polymorphic raptor, parental morphs influence nestling somatic growth and survival, with pairs consisting of different colour morphs ('mixed-morph pairs') producing offspring with lower body mass indices, but higher local apparent survival rates. Resource allocation theory could explain this relationship, with nestlings of mixed-morph pairs trading off a more effective innate immune system against somatic growth. We quantified several innate immune parameters of nestlings (hemagglutination, hemolysis, bacteria-killing capacity and haptoglobin concentration) and triggered an immune response by injecting lipopolysaccharides. Although we found that nestlings with lower body mass index had higher local survival rates, we found no support for the proposed hypothesis: neither baseline immune function nor the induced immune response of nestlings was associated with parental morph combination. Our results suggest that these immune parameters are unlikely to be involved in providing a selective advantage for the different colour morphs' offspring, and thus innate immunity does not appear to be traded off against a greater allocation of resources to somatic growth. Alternative hypotheses explaining the mechanism of a low nestling body mass index leading to subsequent higher local survival could be related to the post-fledgling dependency period or differences in dispersal patterns for the offspring from different morph combinations.

Prevalence and drivers of poison use by South African commercial farmers and perceptions of alternative livestock protection measures.

The use of poison to eliminate predators is causing African vulture populations to collapse. To understand the prevalence and motivations of this practice we conducted an extensive survey with South African commercial farmers. Using a specialised questioning technique and ad hoc quantitative methods we found that an estimated 22% and 31% of farmers used poison over a 1-year and 5-year period, respectively. Poison use hotspots generally coincided with small stock farming areas. The strongest predictor of poison use was whether farmers believed the practice to be common amongst their peers. Our results suggest that farmers' attitudes to vultures are primarily positive, and farmers are less likely to use poisons if they frequently encounter vultures on their farm. Overall, our findings provide an understanding on poison use that provides leverage points to change farmers' behaviour and help avert the African vulture crisis and possible cascading ecosystem impacts.

Nest boxes buffer the effects of climate on breeding performance in an African urban raptor.

As the world's human population increases, transformation of natural landscapes into urban habitats continues to increase. In Africa, rates of human population growth and urbanisation are among the highest in the world, but the impacts of these processes on the continent's biodiversity remain largely unexplored. Furthermore, the effects of ongoing anthropogenic climate change are likely to be severe and to interact with urbanisation. Some organisms appear resilient to urbanisation, and even proliferate in human-modified environments. One such species is the peregrine falcon Falco peregrinus in Cape Town, South Africa. Using a long-term data set (1989-2014), we investigate the relationship between breeding attempts, timing of breeding and breeding performance under varying weather conditions. Exploring these issues along an urbanisation gradient, we focus on the role of artificially provided nest boxes, and their capacity to buffer against extreme weather events. Pairs in more urbanised areas, and particularly those in nest boxes, were more likely to breed and to commence breeding earlier. Additionally, pairs using nest boxes were more likely to breed in years with higher rainfall. Warm and dry weather conditions generally advanced the timing of breeding, although this relationship with weather was not seen for urban pairs using nest boxes. Furthermore, weather did not impact breeding performance directly (breeding success and fledged brood size), but timing of breeding did, with earlier breeders producing more fledglings. Our study shows that falcons breeding in specially provided nest boxes were less sensitive to local weather dynamics than pairs using more natural nest sites. This has important implications as it suggests that the managed provision of such nesting sites can help this key urban species to cope with extreme weather events, which are predicted to increase with climate change.

High prevalence and genetic diversity of Haemoproteus columbae (Haemosporida: Haemoproteidae) in feral pigeons Columba livia in Cape Town, South Africa.

In this study, we explore blood parasite prevalence, infection intensity, and co-infection levels in an urban population of feral pigeons Columba livia in Cape Town. We analyze the effect of blood parasites on host body condition and the association between melanin expression in the host's plumage and parasite infection intensity and co-infection levels. Relating to the haemosporidian parasite itself, we study their genetic diversity by means of DNA barcoding (cytochrome b) and show the geographic and host distribution of related parasite lineages in pigeons worldwide. Blood from 195 C. livia individuals was collected from April to June 2018. Morphometric measurements and plumage melanism were recorded from every captured bird. Haemosporidian prevalence and infection intensity were determined by screening blood smears and parasite lineages by DNA sequencing. Prevalence of Haemoproteus spp. was high at 96.9%. The body condition of the hosts was negatively associated with infection intensity. However, infection intensity was unrelated to plumage melanism. The cytochrome b sequences revealed the presence of four Haemoproteus lineages in our population of pigeons, which show high levels of co-occurrence within individual birds. Three lineages (HAECOL1, COLIV03, COQUI05) belong to Haemoproteus columbae and differ only by 0.1% to 0.8% in the cytochrome b gene. Another lineage (COLIV06) differs by 8.3% from the latter ones and is not linked to a morphospecies, yet. No parasites of the genera Leucocytozoon and Plasmodium were detected.

Response time of an avian prey to a simulated hawk attack is slower in darker conditions, but is independent of hawk colour morph.

To avoid predation, many species rely on vision to detect predators and initiate an escape response. The ability to detect predators may be lower in darker light conditions or with darker backgrounds. For birds, however, this has never been experimentally tested. We test the hypothesis that the response time of avian prey (feral pigeon Columbia livia f. domestica) to a simulated hawk attack (taxidermy mounted colour-polymorphic black sparrowhawk Accipiter melanoleucus) will differ depending on light levels or background colour. We predict that response will be slower under darker conditions, which would translate into higher predation risk. The speed of response of prey in relation to light level or background colour may also interact with the colour of the predator, and this idea underpins a key hypothesis proposed for the maintenance of different colour morphs in polymorphic raptors. We therefore test whether the speed of reaction is influenced by the morph of the hawk (dark/light) in combination with light conditions (dull/bright), or background colours (black/white). We predict slowest responses to morphs under conditions that less contrast with the plumage of the hawk (e.g. light morph under bright light or white background). In support of our first hypothesis, pigeons reacted slower under duller light and with a black background. However, we found no support for the second hypothesis, with response times observed between the hawk-morphs being irrespective of light levels or background colour. Our findings experimentally confirm that birds detect avian predators less efficiently under darker conditions. These conditions, for example, might occur during early mornings or in dense forests, which could lead to changes in anti-predator behaviours. However, our results provide no support that different morphs may be maintained in a population due to differential selective advantages linked to improved hunting efficiencies in different conditions due to crypsis.

Differential fitness effects of moonlight on plumage colour morphs in barn owls.

The Moon cycle exposes nocturnal life to variation in environmental light. However, whether moonlight shapes the fitness of nocturnal species with distinct colour variants remains unknown. Combining data from long-term monitoring, high-resolution global positioning system tracking and experiments using prey, we show that barn owls (Tyto alba) with distinct plumage colourations are differently affected by moonlight. The reddest owls are less successful at hunting and providing food to their offspring during moonlit nights, which associates with lower body mass and lower survival of the youngest nestlings and with female mates starting to lay eggs at low moonlight levels. Although moonlight should make white owls more conspicuous to prey, it either positively affects or does not affect the hunting and fitness of the whitest owls. We experimentally show that, under full-moon conditions, white plumage triggers longer freezing times in prey, which should facilitate prey catchability. We propose that the barn owl's white plumage, a rare trait among nocturnal predators, exploits the known aversion of rodents to bright light, explaining why, counterintuitively, moonlight has a lesser impact on the whitest owls. Our study provides evidence for the long-suspected influence of the Moon on the evolution of colouration in nocturnal species, highlighting the importance of colour in nocturnal ecosystems.

The relationship between wealth and biodiversity: A test of the Luxury Effect on bird species richness in the developing world.

The Luxury Effect hypothesizes a positive relationship between wealth and biodiversity within urban areas. Understanding how urban development, both in terms of socio-economic status and the built environment, affects biodiversity can contribute to the sustainable development of cities, and may be especially important in the developing world where current growth in urban populations is most rapid. We tested the Luxury Effect by analysing bird species richness in relation to income levels, as well as human population density and urban cover, in landscapes along an urbanization gradient in South Africa. The Luxury Effect was supported in landscapes with lower urbanization levels in that species richness was positively correlated with income level where urban cover was relatively low. However, the effect was reversed in highly urbanized landscapes, where species richness was negatively associated with income level. Tree cover was also positively correlated with species richness, although it could not explain the Luxury Effect. Species richness was negatively related to urban cover, but there was no association with human population density. Our model suggests that maintaining green space in at least an equal proportion to the built environment is likely to provide a development strategy that will enhance urban biodiversity, and with it, the positive benefits that are manifest for urban dwellers. Our findings can form a key contribution to a wider strategy to expand urban settlements in a sustainable way to provide for the growing urban population in South Africa, including addressing imbalances in environmental justice across income levels and racial groups.

Patterns of satellite tagged hen harrier disappearances suggest widespread illegal killing on British grouse moors.

Identifying patterns of wildlife crime is a major conservation challenge. Here, we test whether deaths or disappearances of a protected species, the hen harrier, are associated with grouse moors, which are areas managed for the production of red grouse for recreational shooting. Using data from 58 satellite tracked hen harriers, we show high rates of unexpected tag failure and low first year survival compared to other harrier populations. The likelihood of harriers dying or disappearing increased as their use of grouse moors increased. Similarly, at the landscape scale, satellite fixes from the last week of life were distributed disproportionately on grouse moors in comparison to the overall use of such areas. This pattern was also apparent in protected areas in northern England. We conclude that hen harriers in Britain suffer elevated levels of mortality on grouse moors, which is most likely the result of illegal killing.

Where eagles soar: Fine-resolution tracking reveals the spatiotemporal use of differential soaring modes in a large raptor.

Unlike smaller raptors, which can readily use flapping flight, large raptors are mainly restricted to soaring flight due to energetic constraints. Soaring comprises of two main strategies: thermal and orographic soaring. These soaring strategies are driven by discrete uplift sources determined by the underlying topography and meteorological conditions in an area. High-resolution GPS tracking of raptor flight allows the identification of these flight strategies and interpretation of the spatiotemporal occurrence of thermal and orographic soaring. In this study, we develop methods to identify soaring flight behaviors from high-resolution GPS tracking data of Verreaux's eagle Aquila verreauxii and analyze these data to understand the conditions that promote the use of thermal and orographic soaring. We use these findings to predict the use of soaring flight both spatially (across the landscape) and temporally (throughout the year) in two topographically contrasting regions in South Africa. We found that topography is important in determining the occurrence of soaring flight and that thermal soaring occurs in relatively flat areas which are likely to have good thermal uplift availability. The predicted use of orographic soaring was predominately determined by terrain slope. Contrary to our expectations, the topography and meteorology of eagle territories in the Sandveld promoted the use of soaring flight to a greater extent than in territories in the more mountainous Cederberg region. Spatiotemporal mapping of predicted flight behaviors can broaden our understanding of how large raptors like the Verreaux's eagle use their habitat and how that links to energetics (as the preferential use of areas that maximize net energy gain is expected), reproductive success, and ultimately population dynamics. Understanding the fine-scale landscape use and environmental drivers of raptor flight can also help to predict and mitigate potential detrimental effects of anthropogenic developments, such as mortality via collision with wind turbines.