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.     

Parental differences in brood provisioning by Hen Harriers Circus cyaneus

Capsule Females varied their provisioning patterns according to brood age and brood size, whereas males did not.

Aims To quantify how parents balance the needs of their offspring for food and protection.

Methods We studied 13 nests from hides and spent on average 101 hours per nest monitoring prey types, provisioning rate and the time spent at the nest by both sexes in relation to brood size and brood age.

Results Males always provided more food than females. Males brought similar amounts of prey items irrespective of brood size and nestling age, whereas females brought more prey and bigger items to larger and older broods. Females spent less time brooding larger broods, particularly early on.

Conclusions Hen Harrier parents share the provisioning burden, with each parent delivering prey as a function of brood care requirements, hunting capability and the behaviour of the other parent.     

Activity patterns of breeding Hen Harriers Circus cyaneus assessed using nest cameras

Nest camera footage from 13 Hen Harrier Circus cyaneus nests was analysed to document patterns of adult attendance, incubation, brooding and prey delivery rates. Nest attendance was high throughout the incubation stage and began to decrease when chicks were five days old. Chick provisioning increased gradually after hatching and peaked when the chicks were five days old. Daily activity rates were highest during the middle of the day, from 08:00 to 19:00 hours.     

Habitat selection and foraging success of Hen Harriers Circus cyaneus in west Scotland

The foraging behaviour of Hen Harriers breeding in west Scotland was studied in a variety of afforested and moorland habitats. Time budget information from sample areas, stratified by habitat type and controlled for nest distance, was used to investigate habitat selection. Preferences were then related to measures of foraging performance in each habitat. Hen Harriers foraged preferentially over young first rotation coniferous forests, and selected heathland and grassland habitats ahead of closed canopy woodland. Harrier preference for the various habitats was positively correlated with the frequency with which they struck at, and captured, prey. Prey strike success was much lower than that reported for moorland managed for Red Grouse Lagopus lagopus scoticus. The results explain many of the reported changes in local Hen Harrier populations. The extent of young first rotation forestry is currently in decline. It is predicted that this will lead to a reduction in numbers of Hen Harriers breeding in west Scotland, and an increase in the proportion of the British harrier population occupying moorland managed for Red Grouse.     

Is female investment in defence correlated with breeding success in Hen Harriers Circus cyaneus

CapsuleThe intensity of parental defence, irrespective of the value of offspring, may be one of the proximate causes of breeding success, reflecting the quality of breeders.

Aims To test whether female Hen Harrier investment in defence towards a human predator influences breeding success.

Methods Generalized linear mixed models were used, with nest content, date of visit, the interaction (nest content*date), breeding success (number of fledglings/clutch size) and presence or absence of male as explanatory variables. ‘Female’ was included as a random variable in the model. Alarm call rate by a female Hen Harrier during the first three minutes after my arrival at a nest was used as a dependent variable (as an indicator of parental investment in defence towards a human predator).

Results Females were present and alarmed in 100% of visits whereas males were present in 37% of visits. The individual variations in female investment in defence, after controlling for the principal determinants of nest defence, were significantly correlated with breeding success of each pair.

Conclusion The relationship between female investment in nest defence and reproductive success can be explained through differences in female quality. Good quality females, as measured by the level of investment in nest defence, probably also invested more in other breeding aspects not controlled in this study obtaining higher reproductive success.     

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.     

Food niches of sympatric Marsh Harriers Circus aeruginosas and Hen Harriers C. cyaneus on the Dutch coast in winter

Analysis of pellets collected from adjacent communal winter roosts of Marsh Harriers Circus aeruginosus and Hen Harriers C. cyaneus on an extensive saltmarsh in the southwest Netherlands showed highly significant differences between the diets of the two species. Marsh Harrier diet showed no change throughout January, February and March. They specialized on ducks, which were about half of their prey numerically and more important by weight. Although Marsh Harriers sampled other prey, this remained at a low level and showed no seasonal response to the availability of young lagomorphs. Hen Harriers occupied the niche of a generalist predator, having a broader diet and responding to the presumed changes in availability of prey with diet shifts. In November, about half of their prey items were passerines but these declined in importance in December, and small mammals rose. The proportions of both small mammals and birds fell in February and March, due to increasing dependence on young lagomorphs. Diet overlap between the two species was greatest in January and declined in February and March.     

Factors affecting the vulnerability of farmland birds to predation by Hen Harriers Circus cyaneus in winter

Capsule At a communal roost in Cambridgeshire, selection of common birds of arable farmland as prey is interpreted in terms of differences in prey species’ behaviour and size.     

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.     

Sex ratio, survival and territorial behaviour of polygynous Hen Harriers Circus c. cyaneus in Orkney

Polygyny is widespread among Orkney's Hen Harriers Circus cyaneus . From 1975–81, it was associated with a sex ratio estimated from resightings of colour-marked birds up to six years of age to be 29 females:ten males. The sex ratio of fledglings changed significantly from a greater proportion of females in the 1950s and early 1960s to a greater proportion of males subsequently. The mean estimated 'survival' rates (birds colour-marked in Orkney which were seen there in later years) of males and of females 0–2 years old were 14 and 29%. The mean estimated annual survival rates of males and females from 2–6 years old were 72 and 90%. It is suggested that the uneven sex ratio resulted in more frequent intra-sexual encounters and displays by females. The results of temporary removals of two females (immediately replaced) and of two males (not replaced) in spring indicates that there was a shortage of males in the population.     

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.     

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.     

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.     

Anthropogenic pressures within the breeding range of the Hen Harrier Circus cyaneus in Ireland

ABSTRACT

Capsule

Patterns in the frequency and co-occurrence of anthropogenic pressures associated with suitable breeding habitat for Hen Harriers Circus cyaneus demonstrates the need for specific, focussed management and policy options aimed at mitigating impacts on this threatened population.     

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.     

Status of the Hen Harrier Circus cyaneus in the UK and the Isle of Man in 1998

A survey of breeding Hen Harriers in the UK and the Isle of Man carried out in 1998 consisted of two parts: a complete census of 10-km squares at the core of the species'known range, and a random sample of 10-km squares across the rest of its known recent range. From this, the UK and Isle of Man breeding population was estimated at 570 territorial pairs: 436 in Scotland, 19 in England, 28 in Wales, 38 in Northern Ireland and 49 on the Isle of Man. Although there was no overall change in the size of the UK and Isle of Man population between 1988–94 and 1998, numbers declined in Orkney and increased in Northern Ireland. In Scotland, numbers increased on grouse moors and decreased in young plantation between 1988–89 and 1998, so that 55% of the population is now found on grouse moor. A similar situation exists in England, but in Northern Ireland 45% of the population was found breeding in open areas within mature conifer plantations in 1998.     

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.     

Diet composition influences annual breeding success of Montagu's Harriers Circus pygargus feeding on diverse prey

Capsule Three similarly sized passerine species with various breeding strategies behaved differently in the presence of models of mammalian and avian predators.

Aims We tested whether solitary breeders behave in the presence of a predator in order to avoid disclosing their nest site in: hole-nesting Pied Flycatcher Ficedula hypoleuca, shrub-nesting Blackcap Sylvia atricapilla and ground-nesting Meadow Pipit Anthus pratensis.

Methods The behaviour of parents in response to two types of predator model (visually oriented Corvid birds and olfactory oriented Mustelid mammals) placed sequentially at 40, 5 and 1 m distance from the nest was recorded from a shelter.

Results The hypothesis was supported in open-nesting species (Meadow Pipit and Blackcap), as parents did not approach the nest in the presence of a predator. In the hole-nesting Pied Flycatcher the parents disclosed the nest site in most cases (by entering the nest). The intensity of alarm calling increased with decreasing distance of a predator from the nest in all species except Meadow Pipit in the presence of Raven Corvus corax models. The intensity of attacking changed only in Meadow Pipit with decreasing distance of Stoat Mustela erminea from the nest.

Conclusion The results showed that anti-predator behaviour was species-specific, depending on type of predator, habitat and nest inaccessibility.