Activity patterns and resource use by sheep and red deer grazing across a grass/heather boundary

Diet selection and activities of sheep and red deer grazing alone and together across a grass (primarily Lolium perenne/Agrostis capillaris ) and heather ( Calluna vulgaris ) boundary, between July and October, are described.
Throughout the experiment, both animal species had a much higher proportion of grass than heather in their diet. Time spent grazing on each type of vegetation was similarly greater on grass, and the bite rates of the animals were also more rapid on the grass than on the heather.
Red deer spent more time grazing per 24 hours than did sheep, with sheep grazing less during darkness hours than red deer. Total grazing time of both species did not decrease with decreasing daylength from July to October.
There were no consistent differences in the diurnal patterns of activity of sheep and red deer, with both species showing 2-3 large grazing peaks per day.
There were no consistent effects of mixing sheep and deer on their grazing behaviour and diet composition.     

Golden Eagles Aquila chrysaetos: land use and food in northeast Scotland

This paper contrasts changes in breeding numbers and breeding success of Golden Eagles Aquila chrysaetos on four areas with different land uses. On land primarily used for deer stalking but also for grouse shooting, and supporting abundant prey, the number of eagle pairs was steady in 1944-80. On deer land with fewer prey, the number of pairs declined greatly in the 1960s, when deer carrion became scarcer following increased shooting of red deer. On grouse moors incorporating deer stalking, the number of pairs declined in the 1950s due to persecution by gamekeepers, and then largely recovered as this lessened. On grouse moors with little or no stalking, the number of pairs fell after 1946 and remained low due to persecution, which has continued since then. Eagles on grouse moors bred poorly due to persecution. On deer land they were seldom persecuted by estate staff, and bred well. On an area of deer land, the mean annual number of young reared per undisturbed clutch in summer was related to the estimated weight of prey in the spring of the same year. The eagles have remained fairly pesticide-free, and bred well in 1963-65 when more contaminated birds in west Scotland bred poorly.     

Hen harriers and red grouse: economic aspects of red grouse shooting and the implications for moorland conservation

1.  Thirgood & Redpath (2008) propose ways in which red grouse : hen harrier conflicts could be resolved. It has also been suggested that grouse management could accept lower bag sizes (number of birds shot) thus reducing the need for intensive management of predators and habitats. This would allow hen harriers to co-exist more easily on grouse moors.
2.  We compare the bags, costs and incomes from these less intensive forms of grouse shooting with the more intensive driven shooting.
3.  Allowing high density grouse moors to decline to low density ones will result in greater loss of income than the corresponding saving of costs. This can result in moor owners abandoning grouse management and thus gamekeepers losing their employment.
4.  Losing gamekeepers from the uplands would jeopardize the protection of heather moorland and Special Protection Areas for birds, large areas of which are keepered and which currently support high numbers of breeding waders.
5.   Synthesis and applications . We agree with the study by Thirgood & Redpath that consideration of social and economic factors will be needed to resolve conflict but a reduction in management effort from driven to walked-up shooting is not the answer. A more satisfactory approach to the harrier : grouse conflict could be to try to reduce harrier predation by means of diversionary feeding and to address the problem of the rapid build-up in harrier numbers by exploring the use of a ceiling on harrier densities.     

The utilization of heather moorland by territorial Red Grouse Lagopus lagopus scoticus

Although the breeding density of Red Grouse Lagopus lagopus scoticus is related to the density of burned patches of heather Calluna vulgaris , territories appear to offer surplus food, and other factors, especially the risk of predation, may be important. In order to determine how Red Grouse make use of heather moorland, an individually marked population was studied on a moor in northeast Scotland from 1992 to 1994. Territorial birds preferred stands of tall heather during the daytime in autumn and winter, but in spring they increased their use of heterogeneous heather/grass mixtures. Furthermore, in spring, females showed a very strong affinity for edges between heather stands of different ages, where they had easy access to nutritious young heather and good cover. When near the vegetation edge, territorial birds spent most time on the side of the edge providing greater cover. Broods occupied ranges in areas with rather more old heather and with more edge than was available over the entire study site. Within their ranges though, broods clearly preferred heterogeneous areas during the first month after hatching, and showed a further preference for vegetation edge. Their behaviour changed during the second month after hatching, when they utilized tall heather to a greater extent, but edge remained important.     

Spatial synchrony in red grouse population dynamics

Red grouse Lagopus lagopus scoticus populations exhibit unstable dynamics that are often characterised by regular periodic fluctuations in abundance. Time-series' of grouse harvesting records collected from 287 management units (moors) across Scotland, England and Wales were analysed to investigate the broad scale patterns of synchrony in these fluctuations. Estimation of the spatial autocorrelation of grouse population dynamics across moors indicates relatively high levels of synchrony between populations on adjacent moors, but that this synchrony declines sharply with increasing inter-moor distance. At distances of greater than 100  km, grouse population time-series exhibit only weakly positive cross-correlation coefficients. Twenty-eight geographical, environmental and other candidate variables were examined to construct a general linear model to explain variation in local synchrony. Grouse moor productivity (average size of shooting bag), distance from the Atlantic coast moving in a north-easterly direction, April and June temperatures, and June rainfall significantly increased the explanatory power of this model. An understanding of the processes underlying synchrony in red grouse population dynamics is a prerequisite to anticipating the effects of large-scale environmental change on regional patterns of grouse distribution and abundance.     

Factors influencing red deer occurrence at the southern edge of their range: A Mediterranean ecosystem

The red deer is in the Iberian Peninsula at the southwestern edge of its European range and although widespread, red deer ecology in Portugal remains poorly understood. By using pellet group counts, we investigate how habitat structure, vegetation composition and human disturbance affect red deer occurrence. Red deer distribution was positively associated with areas with high density of heather, Leguminosae plants and patches with high cover of shrubs, ground cover and tree cover. Red deer occupied areas further away from roads and from villages. Red deer distribution was negatively associated with agricultural fields and areas with high canopy cover.In the perspective of the current climatic changes, continue research on red deer in these so-called edge populations represents an opportunity to assess the ecological responses within an evolutionary perspective and to provide important conservation suggestions for other countries located on the edge of its distribution range. The present results have implications for the conservation of red deer, emphasizing the need for wide range ecological studies. Red deer variation seems to be related to local factors rather than proximity to the edge of its range.     

Enhanced control of sheep ticks in upland sheep flocks: repercussions for red grouse co-hosts

Sheep ticks Ixodes ricinus (Acari: Ixodidae) and tick-borne diseases cause major economic losses in both upland sheep farming and moorland shoots of red grouse Lagopus lagopus scoticus. Sheep were treated with acaricide four times between March and October and double-vaccinated against louping ill virus (LIV), instead of the conventional regime of two acaricide treatments and no vaccinations, on two moors in northern England. Enhanced treatment started at Westerdale Moor in 1995 and at Danby Moor in 2000; the latter had previously represented a spatial control site. From 1992 to 2003, grouse chick condition, tick burdens, reproductive success, shooting bags and LIV seroprevalence were measured. A total of 1297 grouse chicks from 398 broods were examined for ticks. Enhanced acaricide treatment reduced tick burdens by 90%, and LIV seroprevalence decreased in relation to the number of years since treatment began. Breeding success and post-breeding densities of grouse in the current sample area remained unrelated to acaricide treatment, tick burdens or LIV seroprevalence, but 25% and 60% more grouse were shot on Westerdale and Danby, respectively, after treatment enhancement than before. By improving shooting bags, tick management schemes help to maintain the economic viability of grouse moors, which, in turn, provide upland landscape and wildlife benefits.     

Can acaricide‐impregnated leg bands fitted to female red grouse reduce sheep tick parasitization of chicks and increase chick survival?

In parts of northern England, North Wales and the Scottish Highlands, increasing numbers of sheep ticks Ixodes ricinus (Ixodida: Ixodidae), and the louping ill virus they can carry, are considered to be important factors that reduce red grouse Lagopus lagopus scotica productivity. The present study tested this hypothesis by fitting adult female grouse with leg bands impregnated with the acaricide cypermethrin to experimentally control ticks on their chicks on two managed grouse moors in northeast Scotland. The chicks of females fitted with acaricide leg bands showed reduced tick infestations and improved survival in one of the two study years, relative to chicks of control females. Acaricide leg bands constitute a potential management technique that may be adopted by grouse moor managers in circumstances of high tick infestations on grouse chicks.     

The effects of acaricide treatment of sheep on red grouse Lagopus lagopus scotica tick burdens and productivity in a multi‐host system

Ixodes ricinus (Ixodida: Ixodidae) ticks are of economic and pathogenic importance across Europe. Within the uplands of the U.K., management to reduce ticks is undertaken to benefit red grouse Lagopus lagopus scotica (Galliformes: Phasianidae). Management strategies focus on the acaricide treatment of domestic sheep Ovis aries (Artiodactyla: Bovidae), but the effectiveness of this is less certain in the presence of wild hosts, particularly red deer Cervus elaphus (Artiodactyla: Cervidae) and mountain hare Lepus timidus (Lagomorpha: Leporidae). This study examines the effects of sheep management on grouse tick burdens and productivity using sites with a range of wild host densities. Sites at which applications of acaricide were more frequent had lower tick burdens; this relationship was similar on sites with a range of deer densities. However, no direct link was detected between acaricide treatment interval and grouse productivity. Sites with higher deer densities had higher grouse tick burdens and lower productivity [mean ± standard error (SE) young : adult ratio: 1.2 ± 0.2] compared with sites with lower deer densities (mean ± SE young : adult ratio: 1.8 ± 0.1). Sites with higher grouse brood sizes and higher proportions of hens with broods were also those with higher mountain hare abundance indices. This study highlights the importance of the frequent treatment of sheep with acaricide to reduce tick burdens on grouse, even in the presence of wild hosts.     

Role of small mammals in the persistence of Louping-ill virus: field survey and tick co-feeding studies

Louping-ill (LI) is a tick-borne viral disease of red grouse, Lagopus lagopus scoticus Lath. (Tetraonidae: Galliformes), and sheep, Ovis aries L. (Bovidae: Artiodactyla), that causes economic loss to upland farms and sporting estates. Unvaccinated sheep, grouse and mountain hares, Lepus timidus L. (Leporidae: Lagomorpha), are known to transmit LI virus, whereas red deer, Cenrus elaphus L. (Cervidae: Artiodactyla), and rabbits, Oryctolagus cuniculus L. (Leporidae: Lagomorpha), do not. However, the role of small mammals is unknown. Here, we determine the role of small mammals, in particular field voles, Microtus agrestis L. (Muridae: Rodentia), in the persistence of LI virus on upland farms and sporting estates in Scotland, using field sampling and non-viraemic transmission trials. Small mammals were not abundant on the upland sites studied, few ticks were found per animal and none of the caught animals tested seropositive to LI virus. Laboratory trials provided no evidence that small mammals (field voles, bank voles, Clethrionomys glareolus L. (Muridae: Rodentia), and wood mice, Apodemus sylvaticus L. (Muridae: Rodentia), can transmit LI virus between cofeeding ticks and, in the field, LI virus was prevalent only in areas with known LI virus competent hosts (grouse, mountain hares or unvaccinated sheep) and absent elsewhere. In contrast to the case of tick-borne encephalitis (TBE) virus in Europe, it is concluded that small mammals seem to be relatively unimportant in LI virus persistence.     

Rising burden of immature sheep ticks (Ixodes ricinus) on red grouse (Lagopus lagopus scoticus) chicks in the Scottish uplands

The sheep tick Ixodes ricinus (L.) (Acari: Ixodidae) is an ectoparasite of major economic and pathogenic importance in Scotland. Its distribution in the Scottish uplands is assumed to be governed by the abundance and distribution of its definitive hosts (deer and sheep) and climatic variables such as temperature and rainfall. As the numbers of its major host in Scotland, red deer, have increased dramatically and climatic conditions have become more favourable, the level of parasitism could have been expected to rise. We use data gathered from tick counts on over 4000 red grouse chicks Lagopus lagopus scoticus Latham (Galliformes: Tetraonidae) in various experiments over the past 19 years to ascertain whether the intensity and prevalence of parasitism has been increasing. From 1985 to 2003 the average tick burden of a parasitized red grouse chick has grown from 2.60 +/- 1.12 ticks per chick to 12.71 +/- 1.44. Over this period the percentage of chicks of a given brood parasitized has also increased from 4 +/- 2% to 92 +/- 3%. The possible implications of this increase in parasitism for red grouse production are discussed.     

Long-term density-dependent changes in habitat selection in red deer (Cervus elaphus)

Understanding how habitat selection changes with population density is a key concept in population regulation, community composition and managing impacts on biodiversity and ecosystem services. At low density, it is expected that individuals select habitats in terms of their preference, but as population density increases, the availability of resources per individual declines on preferred habitats, leading to competition which forces some individuals to exploit less preferred habitats. Using spatial information of Scottish red deer (Cervus elaphus) winter counts, carried out in 110 areas across Scotland between 1961 and 2004 (a total of 1,206,495 deer observations), we showed how winter habitat niche breadth in red deer has widened with increasing population density. Heather moorland and montane habitats were most and least preferred for deer, respectively. Increasing density favoured the selection of grassland, to the detriment of the selection of heather moorland. The selection of heather and grassland decreased when temperature increased, while the selection of montane and peatland habitats increased. These findings are important for understanding how habitat use, density and population are likely to be affected by weather, and allow us to predict habitat impacts by large mammal herbivory and climate.     

Effects of human disturbance on the diet composition of wild red deer (Cervus elaphus)

Disturbance from human recreational activities may affect the nutrition of free-ranging herbivores due to trade-offs between feeding in preferred habitats and perceived predation risk. To test this hypothesis, we estimated diet composition for red deer in the Scottish highlands in spring, when recreational activity tends to be high, and in winter when it is lower. We analysed faecal samples from three habitat types (grassland, heather moorland and woodland) collected at sites close to a busy track (disturbed) and at a distance from it (less disturbed). The diet consisted of 39% grasses, sedges, herbs and rushes (GSHR) and 58% Calluna vulgaris and Erica spp. (heather) in spring, compared with 14% grasses and 77% heather in winter, with small quantities of Vaccinium spp. (berry) and Pinus sylvestris (tree) in both seasons. In spring, faeces from disturbed grass and woodland sites indicated a diet with less GSHR and more heather and tree than faeces from less-disturbed sites, which could be due to an increased need for vigilance in exposed grassland and the need to seek cover. In contrast, faeces from all disturbed sites in winter indicated a diet with more GSHR and less heather than faeces from less-disturbed sites. This could be due to a seasonal decline in recreation and increase in hunting activity reversing the disturbance levels at the different sites, since hunting is not normally carried out in areas used by the public for recreation. We conclude that there may be nutritional benefits to deer of reducing disturbance near open grassland.     

Behaviour, population changes and dispersal of mountain hares (Lepus timidus) in Scotland

Mountain hares in north-east Scotland spent the day in forms in long heather and moved downhill in the evening to feed in hill pastures. Here they grazed intensively, often in groups of 4–6 individuals. In June and July adult females, then pregnant or lactating, grazed in daylight on the pastures. Leverets spent less time grazing, and more in play and exploration. They spent the day in cover near the feeding areas, and if disturbed during grazing crouched or went into cover, while adults fled. During 1982 and 1983 males predominated in groups of grazing hares, but in 1984 both sexes were equally represented. These changes in sex ratio were reflected in the increased proportion of females trapped for marking between February and late July during 1982–86. There was a dominance order related to weight among male hares and dominant hares approached more females. There was no firm evidence of mate-guarding. Males approached females regardless of their oestrous state and were usually rebuffed with varying degrees of intensity including striking and chasing. There were no interactions between females. Adults of both sexes chased leverets for short distances but leverets joined groups of feeding adults. Neither leverets nor first-winter hares showed evidence of dispersal. Mountain hares avoided sheep and cattle and there were fewer hares after the arrival of sheep in May.     

Recovery of a dwarf birch (Betula nana) population following reduction in grazing by red deer (Cervus elaphus)

Summary

In Britain Betula nana is a nationally scarce plant, its distribution being limited by climate, heather burning and grazing by deer and sheep. The recovery of a population of B. nana was monitored for several years inside a Scots pine regeneration exclosure in N.E. Scotland. Browsing pressure was reduced in the exclosure but a few red deer (amounting to 3–4 km^?2) were deliberately retained. B. nana stem density increased after red-deer density was reduced. Over the same period there was a reduction in browsing inside the exclosure. B. nana stem density was negatively related to heather height, deer presence and tree canopy cover. Comparisons of growth were made with plants outside the exclosure between 1998 and 2000, the mean annual increment of stems, allowing for browsing loss, being respectively 1.25 and 0.45 cm for inside and outside the exclosure. Most stem basal diameters inside the exclosure were smaller than outside. Reducing red deer density to 3–4 km^?2 on similar upland sites with peaty soils could lead to regeneration of B. nana.     

Isolation of habitat patches limits colonisation by moorland Hemiptera

There has been a significant loss in the UK, and elsewhere in Europe, of moorland dominated by heather Calluna vulgaris. One response to this has been the initiation of moorland vegetation restoration projects. Heather moorland has distinct assemblages of invertebrates and the ability of these to colonise newly created habitat patches has not previously been experimentally tested. We established a dispersal and colonisation experiment by transplanting C. vulgaris-dominated turfs within grassland at different distances (up to 40 m) from heather moorland vegetation. Hemiptera were cleared from these turfs by the use of an insecticide and were sampled 1 year later to investigate re-colonisation rates. Hemiptera assemblages on transplanted turfs were most dissimilar to those of heather moor at the greatest distances of these turfs from heather moor. Colonisation rates of heathland-indicator Hemiptera declined exponentially with distance. The number of individual heathland-indicator Hemiptera was higher on turfs 5 and 10 m from heather moor than on turfs transplanted back into the heather moor, possibly due to a crowding effect. Our findings indicate that moorland Hemiptera assemblages may be limited by dispersal ability. We recommend that moorland restoration schemes should be prioritised on ground as close as possible to existing heather moors.     

Highest densities of mountain hares (Lepus timidus) associated with ecologically restored bog but not grouse moorland management

Over the last 20 years, ecological restoration of degraded habitats has become common in conservation practice. Mountain hares (Lepus timidus scoticus) were surveyed during 2017–2021 using 830 km of line transects in the Peak District National Park, England. Historically degraded bog areas were previously reported having low hare numbers. Following bog restoration, we found hare densities of 32.6 individuals km⁻², notably higher than neighboring degraded (unrestored) bog with 24.4 hares km⁻². Hare density on restored peatland was 2.7 times higher than on bogs managed for grouse shooting at 12.2 hares km⁻² and 3.3 times higher than on heather moorland managed for grouse shooting at 10.0 hares km⁻². Yearly estimates varied most on habitats managed for grouse, perhaps indicative of the impact of habitat management, for example, heather burning and/or possible hare culling to control potential tick‐borne louping ill virus in gamebirds. Acid grassland used for sheep farming had a similar density to grouse moorland at 11.8 hares km⁻². Unmanaged dwarf shrub heath had the lowest density at 4.8 hares km⁻². Hare populations are characterized by significant yearly fluctuations, those in the study area increasing by 60% between 2017 and 2018 before declining by ca. 15% by 2020 and remaining stable to 2021. During an earlier survey in 2002, total abundance throughout the Peak District National Park was estimated at 3361 (95% CI: 2431–4612) hares. The present study estimated 3562 (2291–5624) hares suggesting a stable population over the last two decades despite fluctuations likely influenced by weather and anthropogenic factors. Mountain hares in the Peak District favored bog habitats and were associated with restored peatland habitat. Wildlife management should be cognizant of hare density variation between habitats, which may have implications for local extinction risk.     

Trends in ixodid tick abundance and distribution in Great Britain

Abstract.  The popular, but rarely documented, view in Britain is that ticks have increased in distribution and abundance over recent years. To assess this, we gathered evidence for changes in tick distribution and abundance by distributing a survey questionnaire throughout Britain and by analysing trends in the prevalence of tick infestation on red grouse chicks Lagopus lagopus scoticus Latham (Galliformes: Tetranoidae), gathered over 19 years at three Scottish sites, and on deer (Cetartiodactyla: Cervidae) culled over 11 years on 26 Ministry of Defence (MoD) estates. Based on the survey, the current known distribution of Ixodes ricinus Linnaeus (Acari: Ixodidae) has expanded by 17% in comparison with the previously known distribution. The survey indicated that people perceive there to be more ticks today than in the past at 73% of locations throughout Britain. Reported increases in tick numbers coincided spatially with perceived increases in deer numbers. At locations where both tick and deer numbers were reported to have increased, these perceived changes occurred at similar times, raising the possibility of a causal link. At other locations, tick numbers were perceived to have increased despite reported declines in deer numbers. The perceptions revealed by the survey were corroborated by quantitative data from red grouse chicks and culled deer. Tick infestation prevalence increased over time on all grouse moors and 77% of MoD estates and decreased at six locations.     

The influence of different aspects of grouse moorland management on nontarget bird assemblages

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Management of heather for game and livestock

MOSS, R., 1989. Management of heather for game and livestock. Heather provides food and cover for moorland herbivores. Interactions between soils, drainage, climate, muirburn and grazing affect the species composition, structure and growth of moorland swards. Together, these aspects of a sward determine which and how many herbivores it can support. Grazing and burning maintain much heather ground which would otherwise revert to scrub or woodland. Heavy grazing by ungulates can turn a heathery sward into a graminaceous one, so reducing numbers of grouse and mountain hares. On good soils, invading graminoids are usually nutritious grasses and the carrying capacity for ungulates can increase. On poor soils, grazing and frequent burning may lead to swards dominated by poor quality grasses and sedges so that the carrying capacity for ungulates declines. One way of reinstating heather dominance is to remove livestock; where there are no heather plants left, other techniques may be necessary.