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

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1 Journal of Applied Ecology 2009, 46, doi: /j x FORUM Hen harriers and red grouse: economic aspects of red grouse shooting and the implications for moorland conservation Nick Sotherton* 1, Stephen Tapper 1 and Adam Smith 2 1 Game & Wildlife Conservation Trust, Fordingbridge, Hampshire, UK; and 2 Game & Wildlife Conservation Trust, Couston, Newtyle, Angus, UK Summary 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. Key-words: conflict, economics, grouse shooting, raptors, moorland, waders, predation Introduction Resolving human wildlife conflicts is of increasing importance in conservation, and the conflict between hen harriers Circus cyaneus (L.) and red grouse Lagopus lagopus scoticus (Latham) described by Thirgood & Redpath (2008) is one of the most intractable in Britain. Thirgood & Redpath (2008) correctly conclude that social and economic as well as ecological factors play a role in the problem and its resolution. Conservationists, policy makers and their advisors cannot ignore these non-biological issues, especially those relating to the economy of the land where the conflict plays out. Although not discussed in Thirgood & Redpath (2008) much of the conflict between red grouse and hen harriers arises from the need to produce high grouse densities to justify the *Correspondence author. nsotherton@gct.org.uk large investment made by moor owners in moorland management. Therefore, in the wider debate over conflict resolution it has been suggested that a less intensive management regime with consequent lower harvest rates would be an acceptable way forward to resolving this problem. In this study, we explore the social, economic and conservation benefits of the more intensive forms of red grouse management, and explain the economic differences between driven shooting and the less intensive walked-up shooting and the moorland management differences associated with each. These two types of shooting represent the options at either end of a spectrum of management intensity. Walked-up shooting involves smaller bags (numbers of grouse shot) and it is argued that this should more easily allow hen harriers to co-exist with upland game shooting. We consider the viability of this option using data from the Game & Wildlife Conservation Trust s bag recording scheme (Aebischer & Baines 2008). Ó 2009 The Authors. Journal compilation Ó 2009 British Ecological Society

2 956 N. Sotherton et al. The extent of grouse shooting There were estimated to be 459 grouse moors in the UK in the 1990s (Hudson 1992). In England they covered 4428 km 2 or 56% of the total upland cover (defined by the Department for Environment, Food and Rural Affairs as land above the designatedmoorlandline).inhighlandscotlandtherewere191 grouse moors, 105 grouse moors in the Southern Uplands of Scotland, 153 moors in northern England and 10 in Wales. Hudson (1992) estimated that these moors employed 480 keepers in Scotland and 223 in England. Grouse management will have decreased since 1992 with losses in Wales, Bowland and Cumbria, and Scotland, so the extent and influence of grouse management in these areas has decreased but the intensity of management has increased in the Pennines, North York Moors, Angus Glens and north-east Scotland as new owners tenants have moved in. An update of the extent of moorland management is urgently needed. Grouse shooting provides other rural employment in the off-season months in these Less Favoured Areas. In 2001, this was worth 144Æ8M of gross domestic product (GDP) in Scotland and was calculated to have supported 904 full-time equivalent jobs in hotels and other types of accommodation in Scotland (Dunlop 2001). Game shooting is estimated to be worth 1Æ6 billion to the UK economy. Approximately 12% of the total UK shooting provision consists of grouse shooting (Anon. 2006). Styles of grouse shooting: driven vs. walked-up On a day s driven-grouse shooting, some local, selfemployed people are hired to act as beaters, sweeping in an area of moorland and driving the birds over a line of between eight and 10 guns (shooters) usually standing behind stone or turf-built butts. Additional people can be employed as gun loaders and a further team with gun-dogs are employed as pickers-up to retrieve shot grouse. Catering is also usually provided. Depending on grouse densities there can be up to 50 driven days per season. An alternative method of shooting is the so-called walkedup shooting. This method either involves two or three guns shooting over pointing dogs or a more extended line of 4 7 guns with retrieving dogs. The aim is to walk across the moor flushing and shooting birds as they are encountered. Both styles of shooting are organized by the moors gamekeepers who are full-time employees engaged in managing the grouse all year, not just during the shooting season (from 12 August to 10 December). However, walked-up shooting employs considerably fewer people, usually only 10% of those involved in a driven day. Although management for grouse is essentially the same for both styles of shooting, driven shooting requires more birds, a larger work force on shoot days and more infrastructures (tracks and butts) than walked-up shooting. Grouse production on moors which regularly walk-up is usually lower than on the more intensively managed driven moors. At least one gamekeeper per 1500 ha of heather moor is required to regularly exceed the 60 grouse per 100 ha in August needed to make driven grouse shooting worthwhile (Hudson 1992; Hudson & Newborn 1995). We suspect that this estimate needs to be reviewed upwards because the accuracy of estimating grouse density has improved in recent years. The extent and economics of different shooting styles To compare the extent of these grouse shooting styles we used the Game & Wildlife Conservation Trust s National Gamebag Census (NGC). This is a voluntary scheme that currently collects bag statistics from over 600 UK estates annually (Tapper 1992). Up to and including the 2002 season, the NGC contains a sample of 157 properties in England, Scotland and Wales that shoot grouse. Of these, 62 have driven shooting only and 67 walked-up only; the remaining 28 run both. On each property the total bag of grouse shot, the numbers of gamekeepers employed and the area of moor managed is recorded. Moor owners record the number of days spent either shooting walked-up or driven. In our sample of moors returning bag data 56% of all shooting days are driven days. In Wales, only two properties shoot grouse and both walk them up. In England, 94% of shooting days are driven, whereas in East Scotland only 40% are driven, and in West and North Scotland only 27% of shooting days are driven. To assess the average income and expenditure of running shoot days we used average figures for 2008 provided by the Game & Wildlife Conservation Trust s upland advisory staff. To run an average-sized driven day we have assumed the following costs: 25 beaters 40 = 1000; three pickers-up 40 = 120; transport = 300; total = In this calculation we have ignored other incidental costs such as catering, loaders and the significant cost of running the moor with the employment of gamekeepers, their clothing, equipment, housing and transport (but see below). Capital costs (butts, lunch huts, tracks, etc.) are not included. For walked-up grouse shooting, costs are minimal for running the day but we have estimated 150 for two helpers with their dogs. Grouse shooting income is valued per brace (two birds) in the bag. In 2008, typical values were: Driven per brace; Walked-up per brace. We analysed the NGC sample using these figures collected up to We included only those moors which were able to show clearly the number of days walked-up and driven, as well as the total bag shot by style. We assumed the cost of a driven day to be 1420 and a walked-up day at 150. We valued driven grouse at 140 per brace and walked-up at 75. The net revenues given in Table 1 are therefore the bag for all days of each style multiplied by the relevant value per brace of grouse, minus expenditure per day times the number of days organized. Regional differences and incomes generated by grouse shooting Table 1 shows there are strong regional differences in the style of shooting. The average moor size in England and East

3 Harrier Grouse conflict 957 Table 1. Grouse shooting bags for 2002 from a sample of National Gamebag Census estates which provide information on numbers of walkedup and driven days. Number of gamekeepers refers to the mean number across all moors in the sample in that region. Total bag is the average of the totals across all moors in the sample in that region. Driven days is the average number of days across all the moors that provide driven shooting (n). Walked days is the average number of days across all the moors that provide walked-up shooting (n). Revenues are the average net incomes from grouse shooting for the two styles of shooting (see text for costs and expenses) not including the costs of employing gamekeepers and other moor management costs. Region No. moors Mean area (ha) Mean no. gamekeepers Mean total bag Driven days Walked days Driven revenue Walked revenue Wales Æ (2) NW England Æ (5) 4 (2) NE England Æ (22) E Scotland Æ (3) 5 (17) W Scotland Æ (3) 7 (18) Cumbria, Lancashire. 2 Northumberland, Durham, North Yorkshire. 3 Grampian, Tayside, Central, Lothian & Borders. 4 Highland, Strathclyde. Scotland is 2500 ha but it is double this in West Scotland. Wales has little or no shooting and what there is, is casual and walked-up. In our sample, six brace of grouse (range 10 2) was the average of the walked-up days. NW England is principally the western edge of the Pennines and the Bowland Fells. The average walked day in this area was 14Æ5(18Æ5 10) brace and a driven day averaged 48 (63 22) brace. In NE England, which isthebulkofthepenninesandthenorthyorkmoors,all the grouse shooting in our sample was driven and yielded an average of 72 (141Æ5 9) brace per day. In East Scotland (moors from the Grampian Mountains and Southern Uplands) the minority of moors that organized driven shoots averaged 70 (80 56Æ5) brace while the majority with walked-up shooting averaged just eight (12Æ5 2Æ5) brace. Bags for West Scotland averaged 27Æ5 (45 10) brace and 6Æ5 (26 2) brace respectively for driven and walked-up. There are striking differences in projected revenues from the different styles and regions. Driven shooting generates roughly 10 times the revenue of walked-up shooting, and moors in England are generating more than double the revenue of Scottish ones. English moors produced higher densities of grouse over a 5-year average than Scottish moors and this may be as a result of the intensity of keepering modified by the prevalence of grouse diseases and some inherent climatic and nutritional factors (Hudson 1992; Smith, Campbell & Redpath 2000). Scottish upland moors can also provide other sources of revenue, such as fishing and deer stalking, which provide additional revenue. Such revenue sources are less readily available in England where stalking red deer (Cervus elephus L). is not available. Were English moor owners to adopt a less intensive approach to managing their moors the data suggest that, even if they employed on average one less gamekeeper on each moor, the loss in revenue would be disproportionately greater (Table 1). Despite these revenues, nowhere are grouse moors likely to be profitable. None of the calculations include the cost of employing a gamekeeper (around per year, including housing, clothing, a vehicle and equipment). It is evident that only in NE England are grouse moors likely to produce enough revenue to cover these costs. In Scotland, none of the moors in our sample appeared to be generating enough income from grouse shooting to cover the cost of moorland management or of employing gamekeepers, and so grouse moor management remains a highly subsidised operation by the private sector. Possible financial and social reasons for smaller grouse bags In some instances, investment in management can be so successful that some English moors are able to produce large numbers of grouse, sometimes in excess of what can actually be harvested. Cattadori et al. (2003) found that large variations in harvesting yields were common at both high and low population densities. In these circumstances, slightly lower grouse densities would not result in lost income. While, theoretically, ecological and economic interests could be used to set grouse densities, restricting management and financial investment to limit growth of a red grouse population is, in practice, unlikely because much of the annual variation is not within management control (e.g. weather). The second reason that may support a move towards smaller grouse bags is a combination of social factors. The multiple satisfaction approach comes from studies of hunter attitudes outside the UK, which show that satisfaction is not only related to the density of the quarry species but to a combination of experiences that include recreation, companionship, environmental ambiance and time of the hunting season (Gigliotti 2000; Heberlein & Kuentzel 2002). Hunters reported significantly higher quality of the hunting experience when a combination of both hunting success and wildlife interactions (other species seen but not bagged) was fulfilled (Wynveen et al. 2005). Studies aiming to identify the relative roles of hunter satisfaction could lead to a new model of red grouse management incorporating both optimal red grouse density for economically viable shooting and conservation of predator

4 958 N. Sotherton et al. species. The willingness to pay for higher conservation value and lower grouse density might be as high for low conservation value and high grouse density but this remains to be tested. The ecology of intensively managed moors We suggest that altering the objectives for management could degrade the conservation value of British moorland. Such an effect would be most noticeable in England (Pennines and North York Moors) where a high proportion of moorland is intensively managed and where potential grouse productivity is greater than on other UK moors. One extra gamekeeper is employed per property on the average English moor (Table 1) compared with Scottish moors. This stimulates more systematic and less fragmented predator control, a more ubiquitous patchwork burning of heather and routine control of grouse diseases (Strongylosis and louping ill virus). Although grouse moors cover some 56% of the English uplands they are not distributed randomly between regions but are mostly concentrated along the Pennines and the North York Moors. In these regions, foxes Vulpes vulpes (L.) occur at very low densities and other predators, such as the carrion crow Corvus corone (L.), are killed very quickly after they appear on the moor (Hudson 1992). Therefore, these regions take on some of the characteristics of islands free of ground predators, and are becoming havens for ground nesting birds vulnerable to nest predation, such as red grouse, black grouse Tetrao tetrix (L.), golden plover Pluvialis apricaria (L.), curlew Numenius arquata (L.), lapwing Vanellus vanellus (L.) and dunlin Calidris alpina (L). Ninety per cent of black grouse leks in northern England are on the margins of grouse moors (Warren & Baines 2004). We believe it to be no accident that the North Pennines, the South Pennines and the North York Moors have all been designated as Special Protection Areas (SPA) mostly on the basis of their substantial numbers of breeding waders. Some 74% of these three SPA are managed as grouse moor. In the Peak District National Park 65% of the upland area is managed for grouse, grouse moors cover 70% of the Park s Special Areas of Conservation designation and 76% of its area designated as a SPA (Sotherton et al. 2009). In the North York Moors National Park 95% of the SPA is grouse moor (Sotherton et al. 2009). Priority upland biodiversity habitats dominated by dwarf shrubs such as Calluna vulgaris (L.), have been better retained on grouse moors than areas of moor not managed for grouse. In a study in Scotland over 40 years, grouse moors lost 24% of their heather cover whereas where grouse management was lost, heather cover had reduced by 41% (Robertson, Park & Barton 2001). In a survey of upland breeding birds across England and Scotland, Tharme et al. (2001) found that some ground nesting waders were more common on grouse moors than on other moorland and that taking account of differences in habitat management and geography, predator control remained as an important factor influencing breeding numbers of some of these species. Two- to fivefold increases in breeding density were reportedinspeciessuchascurlewandlapwing.inan8-year upland predation experiment 57% of golden plover, curlew and lapwing collectively reared chicks on plots where there was predator control compared with 18% of pairs on plots where there was not (Fletcher 2008). On a Scottish moor when grouse were managed for 8 years and then were not managed for a further 7 years, three species of waders, skylark Alauda arvensis (L.) and hen harrier were more abundant when the moor was managed for grouse whilst carrion crow increased in the later period when grouse management ceased and lapwing were virtually lost after keepering ceased (Baines et al. 2008). Hen harriers and grouse moors Breeding hen harriers are absent from the majority of English and Scottish grouse moors and gamekeepers prevent this species from colonising many suitable upland areas keeping it a rare breeding bird in England, if not in Scotland as a whole. However, we argue that a reduction in management intensity and shooting bag sizes is not the way forward. First, our data suggest that the drop in shooting revenue would be substantial. A large decrease in grouse bags would, in all probability, lead to a drop in investment in shooting properties with a negative impact on the local community. With an even greater level of support required from the grouse moor owners under this scenario, grouse shooting could collapse entirely in England and parts of north-east Scotland (as it has performed in Wales and much of the rest of Scotland). Secondly, we suggest that a reduction in the intensity of management of driven grouse moors would significantly reduce the favourable conservation status of heather moorlands and upland wading birds across the UK, thus jeopardising the status of large SPA. Alternative sources of income in the uplands are less attractive to the conservation interests there. Historically, headage payments maintained upland sheep flocks but tended to encourage overstocking which grouse moor management resisted (Robertson et al. 2001). Such payments have been removed posing increasing challenges for upland graziers and for moorland conservation (Hanley et al. 2008; RSE 2008; Condliffe 2009). It now appears that one of the few ways to retain the cultural and conservation benefits of sheep stocking without subsidy is to engage with the grouse moor owners to provide habitat and parasite control services. Alternative land uses are ecologically unacceptable, for example monoculture forestry or woodland expansion which reduces the extent of heather-dominated communities of international conservation value (Thompson et al. 1995). Other upland economic models such as carbon trading, or water quality supply management are largely unproven for multiple contiguous land holdings in the UK uplands (White & Wadsworth 1992; BFRS 2002; RSE 2008) and may even pose long-term risks to water and carbon storage (Davies et al. 2008) but these roles clearly need to be unravelled (Chapman et al. 2009; Dallimer et al. 2009). Wildlife tourism is often cited as an alternative source of income but again potential income figures are scarce and would operate mainly outside the shooting season leaving income sources scarce in the early winter months. However, for the North Pennines, Black (2009) showed that tourists would be more willing to visit such landscapes if more

5 Harrier Grouse conflict 959 favourable land cover types were accessible. Thus, the full extent of tourism in these areas is not yet explored and could contribute to the local economy. Black (2009) also showed that grouse moor owners were prepared to invest in conservation, especially if supported by the public sector. Such income could facilitate landowners being less dependent on grouse shooting, which is currently their largest source of income. Resolving the hen harrier : red grouse conflict should not undermine the management of English grouse moors for sound conservation reasons. These views have been supported by multiple signatories in Scotland (Anon. 2002) and in 2004 English Nature also came to this view (Duff 2004). This is not to say we think grouse moors should operate in the complete absence of hen harriers. There are potentially useful but currently unproven ways of mitigating harrier predation by providing harriers with additional diversionary food to reduce predation on grouse (Redpath, Thirgood & Leckie 2001). Models indicate that driven grouse shooting mightbesustainedinthepresenceofmoderatenumbersof hen harriers (Redpath & Thirgood 2003). Thirgood and Redpath accurately describe the pros and cons of supplementary feeding, and we agree that the impact of the technique on the grouse, the harriers and other generalist predators need to be fully quantified. To date, attempts to experimentally examine these issues have been unsuccessful. It has proved difficult to find large areas of grouse moor with sufficient red grouse and hen harriers to provide data for a statistical analysis powerful enough to be meaningful. However, the only attempt to sustain driven grouse shooting in the presence of an unregulated number of hen harriers failed because the local population of hen harriers increased from two pairs to 20 in <6 years, cutting the autumn numbers of grouse by 50% and eventually causing breeding numbers to decline (Thirgood et al. 2000). The consequent closure of the shoot reduced the commercial value of the land by c per year (Redpath & Thirgood 2003). This apparent tendency of hen harriers to form a breeding colony under the right conditions is probably the biggest issue that needs to be overcome in finding a formula for successfully managing grouse and conserving hen harriers on the same piece of ground. Thirgood & Redpath s (2008) suggestion of a ceiling in harrier numbers certainly deserves further investigation, indeed the idea originated at the Game & Wildlife Conservation Trust (Potts 1998), as does the encouragement of hen harriers on areas of upland Britain not managed for red grouse. References Aebischer, N.J. & Baines, D. (2008) Monitoring gamebird abundance and productivity in the UK: the Game & Wildlife Conservation Trust long-term datasets. Revista Catalana d Ornitologia, 24, Anon. (2002) Scotland s Moorland: Unique And Important. A Statement of Intent. Scottish Natural Heritage, Battleby. Anon. (2006) The Economic and Environmental Impact of Sporting Shooting. Public and Corporate Economic Consultants, Cambridge. Baines, D., Redpath, S., Richardson, M. & Thirgood, S. (2008) The direct and indirect effects of predation by Hen Harriers Circus cyaneus on trends in breeding birds on a Scottish grouse moor. The Ibis, 150(Suppl. 1), BFRS (2002) The Countryside Management Industry in the Scottish Borders: Sustainability and the Pursuit of New Jobs and Revenues (eds Scott Wilson Resource Consultants and Scottish Agricultural Colleges). The Borders Foundation for Rural Sustainability, Foulden. Black, J. (2009) Conservation values of the North Pennines. PhD thesis, Imperial College, London. Cattadori, I.M., Haydon, D.T., Thirgood, S.J. & Hudson, P.J. (2003) Are indirect measures of abundance a useful index of population density? The case of red grouse harvesting. Oikos, 100, Chapman, D.S., Termansen, M., Quinn, C.H., Jin, N., Bonn, A., Cornell, S.J., Fraser, E.D.G., Hubacek, K., Kunin, W.E. & Reed, S. (2009) Modelling the coupled dynamics of moorland management and upland vegetation. Journal of Applied Ecology, 46, Condliffe, I. (2009) Policy change in the uplands. Drivers of Environmental Change in the Uplands (eds A. Bonn, T. Allott, K. Hubacek & J. Stewart), pp Routledge, Abingdon, Oxon. Dallimer, M., Tinch, D., Acs, S., Hanley, N., Southall, H.R., Guston, K.J. & Armsworth, P.R. (2009) 100 years of change: examining agricultural trends, habitat change and stakeholder perceptions through the 20th Century. Journal of Applied Ecology, 46, Davies, G.M., Gray, A., Hamilton, A. & Legg, C.J. (2008) The future of fire management in the British uplands. International Journal of Biodiversity Science and Management, 4, Duff, K. (2004) Report on the Hen Harrier Recovery Project and Outline Proposals for the Future. GC P Paper for General Committee of Council, English Nature, Peterborough. Dunlop, S. (2001) An Economic Study of Grouse Moors: An Update (2001). Fraser of Allender Institute for Research, University of Strathclyde, Glasgow. Fletcher, K. (2008) Predation control and moorland birds. Annual Review of the Game & Wildlife Conservation Trust, 39, Gigliotti, L.M. (2000) A classification scheme to better understand satisfaction of black hill deer hunters: the role of harvest success. Human Dimensions of Wildlife, 5, Hanley, N., Davies, A., Angelopoulos, K., Hamilton, A., Ross, A., Tinch, D. & Watson, F. (2008) Economic determinants of biodiversity change over a 400- year period in the Scottish uplands. Journal of Applied Ecology, 45, Heberlein, T.A. & Kuentzel, W.F. (2002) Too many hunters or not enough deer? Human and biological determinants of hunter satisfaction and quality. Human Dimensions of Wildlife, 7, Hudson, P. (1992) Grouse in Space and Time: The Population Biology of a Managed Gamebird. Game Conservancy Ltd, Fordingbridge, Hampshire. Hudson, P. & Newborn, D. (1995) Red Grouse and Moorland Management. The Game Conservancy, Fordingbridge, Hampshire. Potts, G.R. (1998) Global dispersion of nesting hen harriers: implications for grouse moors in the UK. The Ibis, 140, Redpath, S. & Thirgood, S. (2003) The impact of the hen harrier (Circus cyaneus) predation on red grouse (Lagopus lagopus scoticus) populations: linking models with field data. Birds of Prey in a Changing Environment (eds D.B.A. Thompson, S.M. Redpath, A.H. Fielding, M. Maquiss & C.A. Galbraith), pp The Stationary Office, Edinburgh. Redpath, S.M., Thirgood, S.J. & Leckie, F.M. (2001) Does supplementary feeding reduce predation of red grouse by hen harriers? Journal of Applied Ecology, 38, Robertson, P.A., Park, K.J. & Barton, A.F. (2001) Loss of heather Calluna vulgaris moorland in the Scottish Uplands: the role of red grouse Lagopus lagopus scoticus management. Wildlife Biology, 7, RSE (2008) Committee of Inquiry into the Future of Scotland s Hills and Islands. (Chair) G. McCrone Royal Society of Edinburgh, Edinburgh. Smith, A.A., Campbell, S. & Redpath, S.M. (2000) The Influence of Moorland Management on Grouse and Their Predators. Department of the Environment, Transport and the Regions, Bristol. Sotherton, N., May, R., Ewald, J., Fletcher, K. & Newborn, D. (2009) Managing uplands for game and sporting interests: an industry perspective. Drivers of Environmental Change in Uplands (eds A. Bonn, T. Allott, K. Hubacek & J. Stewart), pp Routledge, Abingdon, Oxon. Tapper, S.C. (1992) Game Heritage: An Ecological Review from Shooting and Gamekeeping Records. Game Conservancy Ltd, Fordingbridge, Hampshire. Tharme, A.P., Green, R.E., Baines, D., Bainbridge, I.P. & O Brien, M. (2001) The effect of management for red grouse shooting on the population density of breeding birds on heather-dominated moorland. Journal of Applied Ecology, 38, Thirgood, S. & Redpath, S. (2008) Hen harriers and red grouse: science, politics and human wildlife conflict. Journal of Applied Ecology, 45,

6 960 N. Sotherton et al. Thirgood, S.J., Redpath, S.M., Haydon, T.J., Rothery, P., Newton, I. & Hudson, P.J. (2000) Habitat loss and raptor predation: disentangling long- and short-term causes of red grouse declines. Proceedings of the Royal Society of London B, 267, Thompson, D.B.A., MacDonald, A.J., Marsden, J.H. & Galbraith, C.A. (1995) Upland heather moorland in Great Britain: a review of international importance, vegetation change and some objectives for nature conservation. Biological Conservation, 71, Warren, P.K. & Baines, D. (2004) Black grouse in northern England: stemming the decline. British Birds, 97, White, B. & Wadsworth, R. (1992) A bioeconomic model of heather moorland management and conservation. Ecological Economics, 9, Wynveen, C.J., Cavin, D.A., Wright, B.A. & Hammitt, W.E. (2005) Determinants of a quality wild turkey hunting season. Environmental Management, 36, Received 19 February 2009; accepted 25 June 2009 Handling Editor: E.J. Milner-Gulland