The direct and indirect effects of predation by Hen Harriers Circus cyaneus on trends in breeding birds on a Scottish grouse moor

Two phases of an experimental reduction in management at Langholm Moor in southwest Scotland provided an insight into factors that determined trends in the abundance of moorland birds. In 1992 the historical control of breeding Hen Harriers Circus cyaneus stopped and in 2000 grouse moor management was discontinued. Golden Plover Pluvialis apricaria , Lapwing Vanellus vanellus , Curlew Numenius arquata and Red Grouse Lagopus lagopus scoticus declined, whilst Carrion Crow Corvus corone and Snipe Gallinago gallinago increased. Hen Harriers increased from two to 20 breeding females, then declined back to two. Lapwing abundance was positively associated with that of Hen Harriers, and moorland passerines (Skylark Alauda arvensis , Meadow Pipit Anthus pratensis and Stonechat Saxicola torquata ) were negatively associated. Golden Plover, Lapwing, Curlew, Red Grouse, Skylark and Hen Harrier were more abundant when the moor was managed for grouse, whilst Carrion Crow, a common predator of clutches of ground-nesting birds, increased during the second half when management ceased. Increased Crow numbers, together with an increase in Red Foxes Vulpes vulpes , probably contributed to the observed bird declines.     

Numerical and functional responses of Common Buzzards Buteo buteo to prey abundance on a Scottish grouse moor

Predators will often respond to reductions in preferred prey by switching to alternative prey resources. However, this may not apply to all alternative prey groups in patchy landscapes. We investigated the demographic and aggregative numerical and functional responses of Common Buzzards Buteo buteo in relation to variations in prey abundance on a moor managed for Red Grouse Lagopus lagopus scotica in south‐west Scotland over three consecutive breeding and non‐breeding seasons. We predicted that predation of Red Grouse by Buzzards would increase when abundance of their preferred Field Vole Microtus agrestis prey declined. As vole abundance fluctuated, Buzzards responded functionally by eating voles in relation to their abundance, but they did not respond demographically in terms of either breeding success or density. During a vole crash year, Buzzards selected a wider range of prey typical of enclosed farmland habitats found on the moorland edge but fewer Grouse from the heather moorland. During a vole peak year, prey remains suggested a linear relationship between Grouse density and the number of Grouse eaten (a Type 1 functional response), which was not evident in either intermediate or vole crash years. Buzzard foraging intensity varied between years as vole abundance fluctuated, and foraging intensity declined with increasing heather cover. Our findings did not support the prediction that predation of Red Grouse would increase when vole abundance was low. Instead, they suggest that Buzzards predated Grouse incidentally while hunting for voles, which may increase when vole abundances are high through promoting foraging in heather moorland habitats where Grouse are more numerous. Our results suggest that declines in their main prey may not result in increased predation of all alternative prey groups when predators inhabit patchy landscapes. We suggest that when investigating predator diet and impacts on prey, knowledge of all resources and habitats that are available to predators is important.     

The importance of pre-thicket conifer plantations for nesting Hen Harriers Circus cyaneus in Ireland

Hen Harriers Circus cyaneus are threatened across much of their range and their conservation requires appropriate habitat management. The locations of 148 Hen Harrier nests found in the Republic of Ireland during national breeding surveys in 2000 and 2005 were used to assess nest-site selection. The distribution of these nests was compared to distributions of randomly located points to investigate selection at the scale of the nest-site and landscape. The main nesting habitats selected were pre-thicket stage of first and, particularly, second rotation plantations, mostly of exotic conifers. There was no evidence that the area of post-closure plantations negatively affected Hen Harrier nest distribution. There was a positive correlation across study areas between changes in numbers of Hen Harrier nests between 2000 and 2005 and changes in the area of pre-thicket second rotation plantations over the same period. The overall effect of plantation forests on breeding Hen Harriers in Ireland therefore appears to be positive. However, this study did not consider the effects of plantation habitats on breeding success. Improved grassland was strongly avoided as a nesting habitat. Furthermore, after controlling for the influence of nesting habitat on nest location, landscapes with a high percentage cover of improved grassland were also avoided. Further agricultural intensification of grassland in areas where Hen Harriers breed is likely to have a negative impact on this species. These results are required for the development of management strategies for the conservation of this species.     

Global dispersion of nesting Hen Harriers Circus cyaneus; implications for grouse moors in the U.K.

In the U.K., a full recovery of Hen Harrier Circus cyaneus breeding numbers is prevented by illegal culling by some gamekeepers who fear the species threatens the future of grouse moors. This study's main purpose was to estimate how many more Hen Harriers there would be in the U.K. if this culling were to cease. A review of 33 studies of the Hen Harrier in nine countries showed that the densities of nesting females decreased strongly as the size of the areas studied increased and also that such information could not be used to estimate the potential density over a large area, such as the U.K. Correcting for the size of the areas studied showed that the potential density of nesting Hen Harriers on grouse moors could be predicted from studies overseas. Estimates of the densities of Hen Harriers in occupied areas globally were therefore used to estimate the density appropriate for the U.K. If all potential habitats were occupied, present numbers could more than double, to an estimated 1660 nesting females. This estimate represents an average of one nesting female per 2 5 km² of habitat, a density which would cause little or no significant economic damage on grouse moors. However, because Hen Harriers tend to aggregate, they would not spread out evenly but would nest in relatively high densities on a number of moors. The economic impact on Red Grouse Lagopus lagopus would not be a function of overall numbers, rather it would arise from the uneven dispersion of nesting Hen Harriers.     

Hen harrier Circus cyaneus nest sites on the Isle of Mull are associated with habitat mosaics and constrained by topography

Capsule: Hen Harriers Circus cyaneus on the Isle of Mull, UK, are associated with habitat mosaics consisting of moorland, scrub and forestry but avoid grazed land, suggesting that forested habitats could be managed sympathetically for the Hen Harrier in the future should the current UK population increase.

Aims: To use distribution modelling to investigate nesting habitat associations using a long-term data set for the Hen Harrier on Mull.

Methods: We develop area-interaction models using a least absolute shrinkage and selection operator penalty to explore the distribution of 102 Hen Harrier nest sites in relation to habitat and topography. Our model is then successfully validated in tests using data for 70 nest sites from subsequent years.

Results: Our model is effective in predicting suitable areas for Hen Harrier nest sites and indicates that Hen Harriers on Mull are found in habitat mosaics below 200 m above sea level. Hen Harrier nest intensity is positively associated with increasing proportions of moorland and scrub, open canopy forestry and closed canopy forestry. Nest intensity is negatively associated with increasing proportions of grazed land.

Conclusion: Hen Harriers avoid grazed areas but are relatively tolerant of other habitat combinations. These findings are supported by previous observations of Hen Harrier habitat use and have implications for the recovery of some Hen Harrier populations in Special Protection Areas and future forest management. Open canopy forest and forest mosaics could potentially be incorporated into landscape-scale conservation plans for Hen Harriers using the population on Mull as an example.     

Factors influencing Hen Harrier Circus cyaneus territory site selection and breeding success

ABSTRACT

Capsule: Our findings regarding Hen Harrier Circus cyaneus territory site selection and breeding success in Ireland offer an opportunity for the development of initiatives and conservation actions aimed at enhancing the suitability of upland areas for breeding Hen Harriers and ensuring the long-term persistence of the species.

Aims: To investigate landscape-scale associations between habitat composition and Hen Harrier territory site selection, and to explore the influence of habitat and climate on breeding success.

Methods: We used multi-model inference from generalized linear models and Euclidean distance analyses to explore the influence of habitat, topographic, anthropogenic and climatic factors on Hen Harrier territory selection and breeding success in Ireland, based on data from national breeding surveys in 2010 and 2015.

Results: Hen Harrier territories were associated with heath/shrub, bog and pre-thicket coniferous forests. Comparisons between territories and randomly generated pseudo-absences (upland and lowland) showed that breeding pairs preferentially select for these habitats. Breeding success was negatively influenced by rainfall early in the breeding season and by climatic instability, and was positively influenced by the presence of heath/shrub and bog.

Conclusions: The results suggest that Hen Harrier breeding success is compromised by the synergistic effects of climate, landscape composition and management. Effective conservation of Hen Harriers in Ireland will therefore rely on landscape-scale initiatives.     

Diversionary feeding and nestling diet of Hen Harriers Circus cyaneus

Capsule: Diversionary feeding reduced Hen Harrier Circus cyaneus nestlings’ natural food intake by half. Red Grouse Lagopus lagopus scotica chicks constituted 0–4% of all nestling food items. Annually, this reduced annual grouse chick production by 0–6%.

Aim: To quantify proportions of diversionary and natural food (including grouse) delivered to Hen Harrier nestlings in relation to brood size, male status and natural prey abundance.

Methods: We recorded diversionary food provisioned to 25 Hen Harrier broods (2008–15) and studied the diet of 15 broods using observations from hides, nest cameras and regurgitated pellet analysis. Variation in nestling diet was analysed using compositional analysis.

Results: Hen Harriers took 76% of diversionary food provided. Depending on assessment method, average nestling diet was 44–53% diversionary food, 39–55% natural prey (including 24–45% passerines, 4–15% small mammals, 0–4% grouse chicks) and 0–9% unknown items. The amount of diversionary food consumed was not influenced by male status, brood size or natural prey abundance. The number of Red Grouse chicks delivered annually was 34–100% lower than expected under unfed conditions, however, the confidence intervals associated with these estimates were large.

Conclusion: Diversionary food influenced Hen Harrier nestling diet and reduced the number of Red Grouse chicks taken relative to modelled predictions. It may help reduce conflict between Hen Harrier conservation and Red Grouse shooting, but only if overall grouse productivity is thereby maintained or increased.     

Food-niche differentiation in sympatric Hen Circus cyaneus and Montagu's Harriers Circus pygargus

Hen Harriers Circus cyaneus and Montagu's Harriers Circus pygargus are medium-sized raptors that differ in size (Hen Harrier being slightly bigger than Montagu's Harrier) and breeding system (Montagu's Harriers are semi-colonial and Hen Harriers defend nesting–hunting territories). In contrast, the diets of the two species when in sympatry are very similar. We evaluated food-niche differentiation among these coexisting raptor species and how between-species differences in body size and social system influence interspecific relationships. We present data from a study conducted in 1997 and 1998 in northeastern Madrid province (central Spain). Diet of the two species largely overlapped (55–95%) during the breeding season, but Hen Harriers preyed more often on larger species. This segregation was observed both in the average size of the primary prey (lagomorphs) and in the alternative prey (birds for Hen Harriers vs. insects for Montagu's Harriers), and was particularly apparent late in the season. Accordingly, feeding frequency of Montagu's Harriers, but not of Hen Harriers, increased later in the season. Size differences between species in prey brought to the nest were apparent for both males and females. Foraging behaviour also differed, as Hen Harriers spent more time hunting close to the nest than did Montagu's Harriers. This implies that segregation in foraging areas may also exist. Observed niche partitioning may relax the potential for competition between these species.     

The status of the Hen Harrier,Circus cyaneus,in the UK and Isle of Man in 2010

Capsule The fourth national survey of Hen Harrier showed that the population in the UK and the Isle of Man declined significantly between 2004 and 2010.

Aim To estimate the size of the breeding Hen Harrier population (with associated 95% confidence intervals) in the UK and Isle of Man, constituent countries and Scottish regions, in 2010 and calculate population change since previous surveys in 1998 and 2004.

Methods Complete surveys were made of 10-km squares likely to be occupied by breeding Hen Harriers in England, Wales, Northern Ireland and the Isle of Man, using standard methods developed for previous national surveys. In Scotland, a ‘census’ area was non-randomly selected for survey by volunteers, and randomly selected squares were surveyed in two strata covering the rest of the known range.

Results The UK and Isle of Man Hen Harrier population was estimated at 662 territorial pairs (95% confidence interval (CI): 576–770), a significant decline of 18% since 2004. Scotland holds the bulk (76%) of the population (505 territorial pairs; 95% CI: 417–612), with smaller numbers in Northern Ireland (59 pairs), Wales (57 pairs), the Isle of Man (29 pairs) and England (12 pairs). Declines of 49% and 20% were observed in the Isle of Man and in Scotland, respectively, whereas the Welsh population increased by 33%. A significant decrease was recorded in numbers of pairs using young and mature plantation forest in Scotland.

Conclusion The breeding population of Hen Harriers in the UK and Isle of Man declined between 2004 and 2010. Notable decreases in Scotland and the Isle of Man may be related to habitat change and illegal persecution. Illegal persecution continues to limit the population size of harriers in England to very low levels.     

Mismatches between breeding success and habitat preferences in Hen Harriers Circus cyaneus breeding in forested landscapes

During the past century, the upland breeding areas of Hen Harriers in Ireland have been extensively afforested. There is no evidence that this species avoids breeding in heavily forested landscapes and, indeed, young commercial forests in their second rotation are often selected as nest‐sites. However, Hen Harriers have coexisted with these forested areas for only a few decades and it is possible that such landscapes are suboptimal. We examined the relationship between breeding success and habitat using a dataset spanning three years and four study areas in the south and west of Ireland. We assessed whether nest success and fledged brood size were related to habitat type, both at the nest‐site and in the surrounding landscape. Neither measure of breeding productivity was related to total forest cover or to percentage cover of closed canopy forest in the landscape. However, in a subset of areas, high cover of second‐rotation pre‐thicket (young forests planted on land from which a first rotation has already been harvested) in the surrounding landscape was associated with low levels of breeding success. This may be due to factors related to predation, disturbance or prey availability. The fact that second‐rotation pre‐thicket is a preferred habitat for nesting in Ireland suggests that Hen Harriers may be making suboptimal decisions in the landscapes available to them.     

Field Vole Microtus agrestis abundance and Hen Harrier Circus cyaneus diet and breeding in Scotland

In many parts of the global range, voles form an important part of the diet of Hen Harriers Circus cyaneus , and breeding numbers are correlated with the abundance of these small mammals. In Scotland, however, little information is available on harrier diet in the spring and our understanding of causes of variation in harrier breeding density is complicated by human interference. In this paper we explore the relationship between Field Vole Microtus agrestis abundance and harrier spring diet, density and productivity in southern Scotland. Over three years, voles occurred on average in 67% of pellets, and 79% in years of high and intermediate vole abundance. From 1992, the number of breeding harriers increased following protection from illegal persecution. After accounting for this trend, harrier numbers correlated strongly with vole abundance. Harrier clutch size was also correlated with vole abundance. Although fledging success tended to be greater in years of vole increase than in years of vole decline, fledging success was not significantly correlated with the relative abundance of voles, or with the abundance of Meadow Pipits or Red Grouse chicks.     

Status of the Hen Harrier Circus cyaneus in the UK and Isle of Man in 2016

Capsule: The fifth UK and the Isle of Man survey of Hen Harrier in 2016 showed varying trends by country and region.

Aim: To estimate the size of the breeding Hen Harrier population (with associated 95% confidence intervals) in the UK and Isle of Man, constituent countries and Scottish regions, in 2016 and calculate population change over the five surveys to date.

Methods: Complete surveys were made of all 10-km squares likely to be occupied by breeding Hen Harriers in England, Wales, Northern Ireland and the Isle of Man, using standard methods developed for previous national surveys. In Scotland, self-selected 10-km squares were surveyed by volunteers and a stratified random selection of the remaining 10-km squares within the known breeding range was surveyed.

Results: The UK and Isle of Man Hen Harrier population was estimated at 575 territorial pairs (95% confidence limits, 477–694), a non-significant decline of 13% since 2010 but a significant decline of 24% since 2004. Scotland held the majority (80%) of the population with 460 (359–573) territorial pairs. Elsewhere, 46 territorial pairs were recorded in Northern Ireland, 35 in Wales, 30 in the Isle of Man and four in England. Significant decreases were recorded in the number of pairs in Scotland using grouse moor (?57%) and young forest (?54%).

Conclusion: The combined breeding population of Hen Harriers in the UK and Isle of Man has shown a non-significant decline between 2010 and 2016. There were notable decreases in England, Northern Ireland and Wales since 2010.     

Comparative breeding biology of Hen Harrier and Montagu's Harrier: an 8-year study in north-eastern France

Hen and Montagu's Harriers breed in the same cultivated areas of eastern France. We present data from an extensive study conducted in three adjacent areas where 757 nests of the two harriers were monitored between 1993 and 2000, with the aim of comparing the breeding ecology of these two species and to evaluate their possible future trends. Breeding habitat for harriers consisted nearly exclusively of winter cereals, causing great conservation concern in this intensively farmed region. The Hen Harrier was almost absent from two of the study areas. This species showed significantly larger clutch size and higher breeding success than the Montagu's Harrier, and an earlier breeding phenology. It was thus less adversely affected by harvesting activities than Montagu's Harrier. Both species showed a reduced breeding success with increasing laying date. There was a large diet overlap between the two species, possibly leading to competitive interactions. Overall, the Montagu's Harrier should be considered as the more vulnerable of the two species, necessitating conservation measures, such as protection of nests from early harvesting activities. Nevertheless, to maintain both species in agricultural habitats, farming practices that preserve sufficient food should also be promoted.     

Hen Harrier Circus cyaneus population trends in relation to wind farms

Capsule: The data presented here demonstrate a considerable spatial overlap between wind farms and the breeding distribution of Hen Harriers in Ireland, but evidence for a negative impact of wind farms on their population is weak.

Aims: To assess the extent of the overlap between wind farms and breeding Hen Harriers and to investigate their potential impact on Hen Harrier population trends.

Methods: Data on Hen Harrier breeding distribution in 10?km?×?10?km survey squares from national surveys were used in conjunction with information on the location of wind farms to examine whether, and to what extent, changes in Hen Harrier distribution and abundance between 2000 and 2010 were related to wind energy development.

Results: Of the 69 survey squares holding Hen Harriers during the 2010 breeding season, 28% also overlapped with one or more wind farms. Data from 36 of the squares with breeding Hen Harriers during the 2000 survey revealed a marginally non-significant negative relationship between wind farm presence and change in the number of breeding pairs between 2000 and 2010.

Conclusions: A considerable overlap exists between Hen Harrier breeding distribution and the location of wind farms in Ireland, particularly in areas between 200 and 400?m above sea level. The presence of wind farms is negatively related to Hen Harrier population trends in squares surveyed in 2000 and 2010, but this relationship is not statistically significant, and may not be causal. This is the first study to assess the influence of wind energy development on Hen Harriers at such a large geographic and population scale.     

Hen Harrier Circus cyaneus foraging activity in relation to habitat and prey

Capsule In west Scotland, the birds foraged in accordance with the distribution of prey and vegetation associated with early-growth conifer forests.

Aims To investigate foraging Hen Harrier activity in relation to vegetation cover and structure and the abundance of potential prey.

Methods Forty 1 × 1-km sample observation areas were monitored at four sites in west Scotland.

Results Foraging time in the sample squares was dependent on the proximity of harrier nests. Harriers preferred areas with relatively high densities of Field Voles Microtus agrestis and Meadow Pipits Anthus pratensis, associated with tall ground vegetation and shrub layers. They also preferred areas planted with trees but preference declined with increasing tree foliage 2–5 m tall and bare ground.

Conclusion Conifer forests can provide favourable conditions for foraging harriers during the early growth stages. Further work is required to determine the value of other types of woodland.     

Status of the Hen Harrier Circus cyaneus in the UK and Isle of Man in 2004, and a comparison with the 1988/89 and 1998 surveys

Capsule The population of breeding Hen Harriers in the UK and Isle of Man increased between 1998 and 2004, although country and regional trends differed.

Aims To estimate the size of the UK and Isle of Man Hen Harrier breeding population in 2004, and to compare this with previous estimates made in 1988/89 and 1998.

Methods Surveys were carried out in core and a stratified random sample of 10-km squares throughout the known range, using the same methods as in the 1988/89 and 1998 national surveys.

Results There were an estimated 806 territorial pairs in the UK and Isle of Man in 2004, a significant 41% increase from the 1998 estimate of 570 pairs. Increases were found throughout, with the exception of south and east Scotland and England, where numbers decreased. Scotland held 79% of the UK and Isle of Man breeding population in 2004, and 10% of Scottish pairs were associated with non-moorland habitats, such as mature conifer plantations and scrub/brash.

Conclusions There was a substantial increase in the breeding population in most regions of the UK and Isle of Man between 1998 and 2004, possibly aided by increased use of non-moorland habitats. However, populations declined in upland areas of southern and eastern Scotland and northern England. Continuing illegal persecution arising from perceived conflicts between breeding Hen Harriers and driven grouse shooting may be a major cause of these regional declines.     

Interspecific and intraspecific differences in habitat use and their conservation implications for Palaearctic harriers on Sahelian wintering grounds

The floodplains of the West‐African Sahel region have experienced extensive habitat transformation during the past four decades, coinciding with an impoverishment of raptor populations. We investigated foraging patterns of Palaearctic migratory Eurasian Marsh Harriers Circus aeruginosus, Pallid Harriers C. macrourus and Montagu’s Harriers C. pygargus on a floodplain system in northern Cameroon to assess species, sex‐ and age‐related habitat preferences. Sex and age have rarely been incorporated into general studies of raptor habitat associations, despite clear evidence of intrasexual and age‐related differences in foraging strategies and diet composition, potentially carrying strong conservation implications. We found evidence of sexual differences in foraging preference related to land use, particularly in the most sexually dimorphic Pallid Harrier, and evidence that juveniles used different habitats to adults. This constitutes the first quantitative documentation of such differentiation by Palaearctic raptors on African wintering grounds, indicating that general patterns of habitat use in wintering raptors may obscure sex‐ and age‐specific preferences. Contrary to expectations, we found limited evidence for interspecific foraging segregation. Food partitioning by prey mass was related to harrier body mass and facilitated by a diverse availability of prey on human‐transformed floodplains. Anticipated further large‐scale conversion of floodplain habitat into predominantly desiccated grasslands raises concerns about the survival of wintering harriers.     

Ornithological Surveys of Nature Reserves, 1954

A survey of Hen Harrier winter roosts traced 202 sites in Britain to the end of 1985/86 winter, and 12 in Ireland. Communal roosting had been recorded at more than 90% of sites, and 43% were known to be used every winter. Of reported sites 39% were in E England, where coverage was undoubtedly more comprehensive. The majority of sites in England were reported to have been discovered since the mid 1970s. Hen Harriers almost invariably roosted amongst rank ground vegetation, in a variety of open habitats, and 28% of sites were on d rtl ground. Most sites were at low altitudes, but some had been found up to 427 m above sea level. Co-ordinated counts at roosts recorded maximum overall numbers at mid-winter in both 1983/84 and 1984/85, although the pattern of occupancy varied between regions. No estimate could be made of the sizes of the winter populations in Scotland, Ireland and Wales, but extrapolation of counts to estimate the numbers wintering in England indicated populations which peaked at about 400 birds in 1983/84 and 1984/85. Roosts in western Britain held a higher proportion of grey males than in the east. Ringing recoveries indicate that Hen Harriers wintering in England have mixed origins.     

Comparative breeding performance of Marsh Harriers Circus aeruginosus along a gradient of land‐use intensification and implications for population management

Assessing variation in breeding performance in relation to habitat characteristics may provide insights into predicting the consequences of land‐use change on species ecology and population dynamics. We compared four Marsh Harrier Circus aeruginosus populations subject to similar environmental conditions, but which differed in habitat composition, ranging from natural habitats to intensively cultivated areas. Using a 6‐year dataset, we characterized breeding habitat and diet in these four study sites, and analysed breeding performance in relation to this gradient of land‐use intensification. There was minimal variation in breeding performance between study years but consistent variation between study sites. Unexpectedly, Marsh Harriers breeding in intensively cultivated habitats had higher reproductive success than those breeding in more natural habitats, which, however, hosted higher breeding densities, so overall net population productivity (fledglings per unit area) was similar across sites. This resulted from combined effects of density‐dependence and different predation rates between study sites. The colonization of intensive farmland habitats may not necessarily impact negatively on population sustainability when breeding success and population density are traded against each other. However, our findings should not mask longer‐term conservation issues for populations breeding in these intensively managed areas, and further studies should assess potential long‐term negative effects of occupancy of human‐altered habitat.     

Habitat use by Hen Harriers Circus cyaneus on Orkney: implications of land-use change for this declining population

The use of habitats by hunting Hen Harriers Circus cyaneus just prior to the settlement period was investigated on Orkney, where numbers have declined by 70% over the last 20 years. Both males and females hunted over areas that were closer to subsequent breeding territories. Neither sex differed in the amount of time they hunted over areas dominated by either intensive pasture, moorland or rough grazing. However, male hunting was significantly related to the amount of unmanaged grass habitat with a litter layer. Female hunting was related negatively to vegetation height, and to the prevalence of both Heather Calluna vulgaris and managed grass; after controlling for these habitat features, female hunting also tended to be associated negatively with Common Starling Sturnus vulgaris abundance. Dramatic changes in land use on Orkney have occurred over the last 40 years, with increases in the amount of intensive pasture and decreases in the amount of rough grazing. These changes, coupled with a doubling in sheep Ovis aries densities over the last 20 years, are likely to have reduced the amount of unmanaged grass. These changes will have been detrimental to hunting male Harriers by reducing the amount of food they can supply to the females prior to egg laying and during the incubation period.