conserving european mountain habitats: what can …conserving european mountain habitats: what can...

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Conserving European mountain habitats: what can their past tell us about their future? R. A. ~od~shon' & E. G. A. 01sson2 I Institute of Geography and Earth Sciences, University of Wales, UK 2 Department of Biology, Norwegian University of Technology, Trondheim, Norway Abstract This paper looks at the strategies needed for the conservation and sustainable use of some of Europe's most fragile ecosystems: mountain habitats. Despite their appearance as wildscape, many traditional mountain ecosystems are semi-natural habitats whose character has been shaped by past forms of land use management. In some cases, this symbiosis has evolved over many centuries, even millennia, giving a deceptive natural stability to the ecosystems involved. However, as mountain areas across Europe have been drawn into more specialised forms of farm production, the progressive abandonment of traditional husbandry practices, including range grazing by livestock, has led to a reduction and fragmentation of those ecosystems and to the loss of key species within them. Using two sample mountain areas, the Scottish Highlands and Jotunheimen of Norway, this paper focuses on how the presence of particular ecosystems and species needs to be seen within a specific rather than generalised framework of traditional land use management. It will reconstruct this framework, highlighting the differences between the two sample areas and showing how their traditional resource management has contributed to the ecological dynamics of traditional mountain habitats. Drawing on detailed field data at both a community and species level, it will highlight the effect that changes to this traditional resource management have had on mountain habitats. Transactions on Ecology and the Environment vol 63, © 2003 WIT Press, www.witpress.com, ISSN 1743-3541

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Page 1: Conserving European mountain habitats: what can …Conserving European mountain habitats: what can their past tell us about their future?R. A. ~od~shon' & E. G. A. 01sson2 I Institute

Conserving European mountain habitats: what can their past tell us about their future?

R. A. ~ o d ~ s h o n ' & E. G. A. 01sson2 I Institute of Geography and Earth Sciences, University of Wales, UK 2 Department of Biology, Norwegian University of Technology, Trondheim, Norway

Abstract

This paper looks at the strategies needed for the conservation and sustainable use of some of Europe's most fragile ecosystems: mountain habitats. Despite their appearance as wildscape, many traditional mountain ecosystems are semi-natural habitats whose character has been shaped by past forms of land use management. In some cases, this symbiosis has evolved over many centuries, even millennia, giving a deceptive natural stability to the ecosystems involved. However, as mountain areas across Europe have been drawn into more specialised forms of farm production, the progressive abandonment of traditional husbandry practices, including range grazing by livestock, has led to a reduction and fragmentation of those ecosystems and to the loss of key species within them. Using two sample mountain areas, the Scottish Highlands and Jotunheimen of Norway, this paper focuses on how the presence of particular ecosystems and species needs to be seen within a specific rather than generalised framework of traditional land use management. It will reconstruct this framework, highlighting the differences between the two sample areas and showing how their traditional resource management has contributed to the ecological dynamics of traditional mountain habitats. Drawing on detailed field data at both a community and species level, it will highlight the effect that changes to this traditional resource management have had on mountain habitats.

Transactions on Ecology and the Environment vol 63, © 2003 WIT Press, www.witpress.com, ISSN 1743-3541

Page 2: Conserving European mountain habitats: what can …Conserving European mountain habitats: what can their past tell us about their future?R. A. ~od~shon' & E. G. A. 01sson2 I Institute

1 Introduction

Mountain habitats comprise some of Europe's most fragile yet threatened habitats. Despite their appearance as wildscape, many are semi-natural habitats whose character has been shaped by past forms of land use management. In some cases, this symbiosis has evolved over many centuries, even millennia, giving a deceptive natural stability to the ecosystems involved. As mountain areas across Europe have been drawn into specialised market-driven forms of production over the past century or so, the progressive abandonment of traditional husbandry practices, from the management of hay meadows to open range grazing by livestock, has brought about a reduction and fragmentation of those dependent on them. As ecosystems lying along the upper frontier of farming, expected changes in CAP as well as climate change are likely to accelerate change. This paper will explore some of these issues by comparing how changes in land use management have impacted on habitats in two mountain areas set in north-west Europe: the mountains of mid-Norway and those of the Scottish Highlands. It will first, define their traditional systems of land use and how they contributed to the formation of particular habitats. Second, it will look at how change, long-term and short-term, have reduced and fragmented them to the point at which whole habitats or individual species are threatened. Third, it: will conclude by arguing that to conserve what remains of such habitats, we need to go beyond their mere designation as protected sites and restore the specific management conditions and disturbance regimes that fostered them ab initio but should do so within a framework of sustainable farm output.

2 Scottish Highlands and mountains of mid-Norway: physical setting

As mountain areas, mid Norway and the Scottish Highlands share a broadly similar resource set. Physically, both comprise limited areas of valley ground that have long provided the primary focus for settlement and resource exploitation. Above this valley-ground, lay large areas of mountain land whose output was low and seasonally confined. Beyond these general similarities, the character of their resource set differed in its detail. Physically, core areas in mid- Norway have a greater proportion of high ground (over 300 m a.s.l.), and a greater proportion of ground rated as sub-marginal, as opposed to simply marginal, for farming when compared to the Scottish Highlands (Table 1). Whilst the mainland core and eastern areas of the Scottish Highlands could be described as having a more continental-type climate when compared with the more oceanic or maritime weather patterns of the western coastal areas, it is less continental in type than the climate of areas like the Jotunheimen (Table 1).

Prior to any human modification, there were differences in vegetation due to climate and topography. Overall, the core areas of mid-Norway like the Jotunheimen have a greater proportion dominated by alpine and sub-alpine habitats compared to the Scottish Highlands. The Jotunheimen habitats are

Transactions on Ecology and the Environment vol 63, © 2003 WIT Press, www.witpress.com, ISSN 1743-3541

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Table 1: Edaphic factors in Scottish Highlands, UK, and Jotunheimen, Norway. ' = Number of days per year with a mean temperature over 6OC.

Mean annual precipitation, in mm Mean annual temperature, OC

Vegetation period, length in days1 Climate regime Climatic tree line m a d

Dominating vegetation in sub-alpine region

Percent of ground above 300 m a.s.1. Latitude

Scottish Highlands, UK

Eastern Highlands, 800-1000 Western Highlands, coast, 1800-2000

Eastern Highlands, 6 Western Highlands, coast, 8

150-80

OceaniclMaritime Eastern Highlands 650-700 Northwest Highlands 2-300

Pine forest (Pinus sylvestris) with birch (Betula pubescens, B. pendula) and oak (Quercus sessile). Where forest cleared, dry heath with Calluna dominant or grassland, inc. Nardus. Where waterlogged, wet heath, blanket bogs, with Erica tetralix, Molinia caerulea, Scirpus cespitosus and Sphagnum spp.

60-80

E. Jotunheimen, Norway 400-500

Continental 900-1100

Dry pine (Pinus sylvestris) and mountain birch (Betula pendula) woodlands with Cladonia lichens, dry dwarf shrub heathlands with Calluna and Emperhrurn dominant

100

dominated by dry dwarf shrub and lichen communities (Table 1). The sub-alpine communities are dry Scot's pine (Pinus sylvestris) and mountain birch woodlands and pine and mountain birch [l]. In the Scottish Highlands, vegetation can be divided into sub-montane and montane habitats, the former being those areas where, prior to human impacts, vegetation cover was that of forest and the latter being those areas above the tree limit. The forest cover comprised Scots pine (Pinus sylvestris) and birch (Betula spp.), with oak (Quercus spp.) being present in the west [2]. Whilst the upper tree limit in the eastern Highlands was around 650-700 m a d . , it falls to below 300 m a d . in the north-west owing to the greater rainfall and windspeed of the area [3]. This natural forest survives today only in a localised, fragmented form. When cleared, it was replaced by dry heaths, mainly Calluna-Trichoporum in the eastern Highlands and Calluna-Eriophorum in the western Highlands, but where overgrazed, by grassland. Everywhere, but especially towards the north-west, wet Atlantic heaths, blanket bogs, with Erica tetralix, Molinia caerulea and Scirpus cespitosus dominant, are also present. As in the Jotunheimen, dwarf shrubs, moss heath and lichen dominate vegetation cover above the tree line [4].

Transactions on Ecology and the Environment vol 63, © 2003 WIT Press, www.witpress.com, ISSN 1743-3541

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3 Traditional land uses and habitat formation

3.1 Background

Farming communities in both areas evolved similar forms of resource use, with core settlements on the lower valley ground supplemented by seasonal sites on the higher ground from which they could exploit the short summer surge in biomass output from their higher mountain pastures. The character of these summer settlements, and the intensity of their exploitation strategies, varied over time. In both areas, the early phases of the Little Ice Age (1300-1500 AD) involved a withdrawal of settlement and land use from the higher ground, as climatic degradation along with the collapse of populations following the Black Death brought about a contraction of settlement onto the lower ground. Evidence for both regions suggests recovery of settlement was under way by the 17th century. In Norway the 17" century is interpreted as the period for the onset of development of the intensive mountain summer farming [5] , although livestock grazing in mountains started long before this time, in Bronze age, as interpreted from paleoecological studies [6 ] . In the Scottish Highlands too, signs of re-expansion are evident by the 17th century [7].

3.2 Mid Norway

In the Jotunheimen, high ground resources were exploited through the movement of stock to seasonally occupied summer farms, the number of such farms and, therefore the scale of high ground usage increasing from the 17th century onwards. In some cases, these summer farms were set as much as 50-70 km from the permanent winter farms [8]. The stock kept and routinely moved between winter and summer farms were mixed: horses, cattle, sheep and goats. In addition to mixed stocking, three other aspects of husbandry practice had potential significance for habitat change. First, high ground pastures were exploited via open range grazing, with no attempt to disaggregate stock or grazing areas. Second, summer farms were established as fixed sites, each one traditionally attached to an individual farmer with buildings for humans, hay and stock and a tied enclosure that was maintained and mown as a hay meadow (No. setervoll), stock only being allowed access after the hay crop had been taken. In mountains areas like that of Gauldalsvidda, where possibilities existed for harvested substitute fodder from wetlands, these enclosed meadows were small, averaging around 0.5 ha. In the higher and more continental areas like Jotunheimen where the climate did not favour development of wetlands, the enclosed meadows were notably larger, averaging 4 ha [g]. Third, the size of meadow enclosure has a bearing on how farms used the open pasture of their mountain ground. In the higher and drier mountain areas, where heathland dominated by Ericacae species prevailed, there were few or no possibilities for mowing hay from hill ground, so that the enclosed meadow land represents the only source of hay. However, hay was not the only source of fodder used in such areas, farms invested much labour in gathering lichen, mainly Cladonia and

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Cetraria, as fodder. By contrast, in the lower mountain areas, such as Gauldalsvidda, farmers invested much labour in mowing hay from the fens and mesic grasslands that existed on the land that lay around their summer farms.

These husbandry practices affected habitat development in a number of ways. First, the level of grazing, along with the high level of demand for wood for fuel and construction, suppressed tree growth. It has been estimated the tree line has been anthropogenically-suppressed by 200 m [10]. Added to this, the climatic degradation of the Little Ice Age would also have depressed the overall tree limit, recovery only taking place with the climatic amelioration of the late 19th and early 20th centuries. Second, plant ecological analyses of unimproved meadows and semi-natural pastures show higher levels of species richness when compared to modern improved leys and pastures. It was found that semi-natural sub-alpine pastures, formerly managed as meadows, had about twice the number of plant species, 28 per 0.25m2, compared to fertilized leys, 15 per 0.25m2 [ l l]. Third, the human use of the sub-alpine woodlands including livestock grazing, reduced the areas covered by woodlands and created semi-natural sub-alpine grasslands and heathlands. This implies that many alpine plant species have extended their distribution to sub-alpine regions and form a significant element in those plant communities [12]. Some of those species are now considered to be rare species. One of the very few endemic plants in Scandinavia, Primula scandinavica, belongs to this group. Some plant species display indications of population differentiation when growing in the alpine and the subalpine habitats, respectively. It is hypothesized therefore, that the shaping of those semi-natural sub-alpine habitats might contribute to an increase of biological diversity [13].

3.3 Scottish Highlands

Traditional systems of resource management in the Scottish Highlands were also based around the exploitation of the low ground valley areas, supplemented by the exploitation of high ground resources via seasonally occupied sites called shielings. Whilst the region experienced an expansion from the 16th century onwards, it was an expansion driven by the need to maximise the output of arable rather than pasture [14]. For this reason, its use of high ground resource differed in detail from that of areas like the Jotunheimen.

Traditional farming townships in the region maintained a range of stock similar to that found on farms in mid Norway: horses, cattle, sheep and goats. However, their management was compromised by the prioritised needs of arable in two critical ways. First, there was neglect of winter feed for stock and the habitats needed to sustain such feed. The expansion of arable over the 16'-18' centuries was invariably at the expense of land suitable for low lying meadow. Mid lgth century data shows that even in the central Highlands, where townships were dependent on hay for winter fodder owing to the need to winter stock indoors, the average extent of meadow per townships in three sample areas covering 123 townships was barely 2-6 ha. Reduced to the amount of meadow per holding, it represented an average of 0.58 ha per holding (North Lochtayside), 0.74 ha (Strathyre) and 1.73 ha (South Lochtayside) per

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holding 1151. Townships did not combat this deficiency of low ground meadow by establishing enclosed hay meadows around their summer shielings nor is there evidence that they cut hay on any scale from mountain pasture during summer. The home meadow around the main winter settlements was the total available. By expanding arable and reducing the role played by hay, communities limited the flow of nutrients from grass to arable via stock manure, raising their dependence on other manures, such as turf cut from hill pasture 1161.

Second, inadequate levels of winter feed meant that most touns did not carry enough stock through winter to exploit the summer grazing potential of their mountain pastures. Analysed in terms of Standard Livestock Units (SLU), with allowance for the smaller size of pre-19th century stock, stocking levels ranged between 0.0125 and 0.0251ha SLU, compared with 0.5-l/ha SLU for modern day stocking levels [17]. This low density of stocking was combined with a strategy that limited the time stock spent grazing the pasture around shielings sites. In the central Highlands, stock spent no more than six weeks on mountain pasture before being brought back to graze on pasture around the main settlement by the end of Julylearly August and to be folded - at night - on that part of the outfield land scheduled to be cultivated the following spring. However, farmers made heavy use of high ground resources in other ways: timber for building purposes and fuel; peat for manure and fuel; vast quantities of turf for the building of field walls, house walls and roof cover; and heather or bracken for thatch and bedding.

We can link these various resource strategies to a number of ecological impacts. Recent thinkmg has challenged the view that the central Highlands were covered with a great Caledonian forest, with extensive stands of Scots pine and birch, down to the eighteenth century. Pollen data suggests that a great deal of Highland forest clearance had taken place by 1300 AD [IS]. One estimate puts the amount of forest cover left in 1500 as low as 10-15%, so it was already a denuded landscape by then, with cover declining further to 5% by 1750 [19]. This ongoing clearance, 1500-1750, probably stemmed from the clear-felling of timber for export out of the region. However, local needs played a part, at least, in degrading open forest areas further and in preventing regeneration, including the demand for fuel; the use made of young growth for wattle-based housing and partitions; the heavy grazing of low-lying wood by stock searching for winter fodder and shelter; and the grazing out of regenerative growth on hill ground.

Once cleared of forest, hill ground was colonised by shrub heath dominated by Calluna. Pollen data suggests a gradual increase in heather cover over the 2- 300 years prior to the mid-19' century [20], a conclusion consistent with the further reduction in forest cover over this period. As a sera1 climax, Calluna- heath was stabilised via a management regime based on grazing and burning. Though levels of grazing in the region were comparatively light, 16"-mid 18 '~ centuries, the grazing impact of stock like cattle and goats was sufficient to sustain Calluna-heath by suppressing the regeneration of pine, birch and willow. Regular burning also played a vital part. Though it offered some feed advantage for game birds like red grouse, local court records make it clear that heather was burnt by tenants to provide early spring feed for sheep [21].

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What is striking about the Scottish Highlands is the smalI area managed as meadow by the 18" century. Townships maintained only modest areas of enclosed meadow, often on wetter ground close to their arable. Nor is there evidence to suggest that they established areas of enclosed meadow beside their summer shieling sites, or that hay was mown on an extensive scale from areas of open or range pasture. Tall herb grasslands, the type of habitat that forms the basis of mountain meadows elsewhere in Europe, are scarce in the Highlands today, being confined to inaccessible sites beyond reach of stock. Yet even allowing for the impact of heavy grazing on such habitats over the past 200 years, it is unlikely that such meadows were particularly extensive in the region under traditional land use regimes, 16 '~ -18~~ centuries. The species rich sub- montane pasture that survives today comprises a mix of low herbs and grass, with both alpine and sub-alpine species present. Work on alpine and sub-alpine grasslands suggests that areas both mown for hay and grazed had higher species richness than pastures that were simply grazed [22]. This would suggest that whilst light grazing fostered species rich pasture in the Scottish Highlands, especially with stock regularly moving between such pasture and sources of potential propagules, such pasture was not as species rich as it might have been had communities been more active in mowing hill pasture for hay. Further, areas around shieling sites were enriched through the nightly concentration of stock and their manure, the vegetation becoming grassier because higher levels of nutrient loading and much heavier grazing favoured grasses over herbs.

4 The impact of farming change

4.1 Mid Norway

The intensity of the use of the mountains via the summer farming developed and increased up to mid-19' century when a peak in the number of summer farms in Norway was reached [23]. From the 1920' and onwards the modernisation of agrarian practices also affected mountain summer farming and the most labour demanding practices like collection of lichen for winter fodder disappeared successively. Many mountain summer farms sites were connected with roads for motor vehicles and dairies were established. Thus the need for the full processing of milk into cheese, butter and other products at the summer farms gradually decreased. This in turn led to decreased need for wood fuel harvest and thus allowed regrowth of birch in the subalpine region. However, after the World War 11 a number of agricultural changes affected the mountains in Norway. Those changes can be summarized in 4 steps: 1) Abandonment of haymaking in semi-natural hay-meadows in favour of

cultivation of leys. This change included both the transformation of semi- natural grasslands in the mountains into leys by ploughing and re-seeding as well as transforming semi-natural haymeadows into pastures. This process was supported by governmental subsidies to stimulate increased yields in fodder production for livestock and thus in livestock productivity. The ecological results were a reduction in biological diversity on ecological

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community level and on species level. The landscape diversity also decreased by reduction of habitat variability.

2) Fertilization of semi-natural meadows and pastures to increase biological production. The ecological result was decreased biological diversity and the disappearance of plant species vulnerable to increased nitrogen and phosphorus levels [24].

3) Changes in the composition of herds. Horses disappeared along with the mechanisation of agriculture although horses never numerically significant in mountain areas. Goats, on the other hand, constituted the largest number of the livestock components in Sjodalen, eastern Jotunheimen in 1997 [25]. Goats are browsers, viz, they prefer to graze on woody species like shrubs and trees, and they have a significant grazing impact on the landscape. There was an evident decline in the number of goats in the mountains herds from 1920s and today goats exist solely in a few farms as monospecific herds as milk producers [26]. They have geographically limited impact on the landscape [27]. During the 1980s there has been a continuous increase in number of sheep and today the number of sheep in Norwegian mountains is twice as large as in the beginning of the 20" century [28].

4) Changes in grazing regimes - due to changes in livestock composition. Sheep prefer to graze in the alpine region of the mountains. Soon after being released onto the mountains after snow-melting in early June, they wander to the alpine sites and spend the summer there. The subalpine zone that was earlier grazed by goats and cattle has little or no grazing pressure today in many Norwegian mountains [29]. The result is that the birch forest is invading the sub-alpine semi-natural heathlands and grasslands here and the tree line is advancing up the mountains slopes [30].

In mid-Norway, we can document fundamental changes in landscape related to the fore-mentioned adjustments in agriculture, livestock composition and thus in the mountain resource use. Over the period 1964-1989, the open heathlands in Sjodalen, Jotunheimen, decreased with 70% and pine-birch woodland increased with a similar rate [31]. For the mountains north-west of Jotunheimen, Gauldalsvidda, changes were also recorded but due to differences in environment and less dramatic changes in livestock herds, the forest succession on former semi-natural areas in the subalpine region has been less dramatic. For both mountains though, the species-rich semi-natural grasslands used for hay-making in the enclosures at the summer farm sites, were generally transformed into leys.

4.2 Scottish Highlands

Change in the Scottish Highlands has been deep-rooted. Quite apart from the shifts evident before the 1 8 ~ ~ century, more fundamental institutional change took place from the mid-18" century, with arable-based townships being progressively cleared of their subsistence farmers and transformed into commercial sheep farms. This transformation shifted the balance of stock heavily towards sheep and, at the same time, increased the density of stocking. Data for over 30 sheep farms across the southern-central Highlands analysed

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before and after the clearances in terms of SLU puts the scale of this increase in the scale of stocking about 2x. If we add the shift into larger breeds like the Cheviot and Blackface, then the scale of increase was nearer 4x, producing stocking densities of 0.25-0.5 LUIha.

Less easily quantified was the impact of changes in grazing management. Quite apart from the disappearance of goats and reduced numbers of cattle and horses, post-Clearance farming was based on: the dis-aggregation of sheep into specialised flocks or hirsels based on age and sex; the enclosure of low lying and high ground pastures reducing the area of open range grazing; the introduction of rotational grazing on hill pasture, the grazing of stock on hill and mountain pasture for much longer; the use of arable for fodder crops and the re-seeding of pastures; extensive land drainage, reducing on-farm ecological variability.

These changes increased the ecological footprint of grazing substantially. In the area that saw the earliest clearances, the southern-central Highlands, reports of a decline in Calluna heath were being published by the 1790s. This contraction can be linked to the spread of sheep and intensification of local grazing pressure. A further surge in sheep numbers, driven by higher prices and a switch out of cattle, occurred in the 1860s and 70s. The increased grazing pressure reduced the extent of Calluna-heath further, a reduction now evident in pollen diagrams [32]. The spread of grasses at the expense of Calluna completed the succession of forest-Calluna-heath-grarninoids [33], a succession that &fines a key theme in the recent ecological history of the Scottish Highlands.

The fall in the price of sheep and wool after 1880 led to a depression in Highland sheep farming. On some estates, deer replaced sheep in an effort to maintain profitability whilst, on others, the management of heather moor for red grouse became more prominent in the estate economy. Likewise, some landowners began to use farm land to establish plantations of larch and spruce, a strategy taken further by the setting up of the Forestry Commission in 1919.

The post World War I1 period has seen a renewed the debate over sheep farming and its environmental effects. Despite growth in the national Scottish flock between 1945-2000 (6.7M - 9.2M), analyses suggest that overall sheep numbers in Highland areas have not changed fundamentally since 1945. This apparent steady state though, hides internal adjustments. The loss of considerable areas of both hill and valley ground grazings to forestry and to deer has meant that the same number of sheep as in 1945 are now being kept on less land. Numbers, therefore, have only been maintained through local increases in stocking. In Huntly, on the eastern edge of the Highlands, the number of breeding ewes increased by 38%, 1975-1990, even though forest cover also increased by 5 1 % [34]. Increasingly, hill farming has depended on a deepening and broadening range of subsidies. Some, like the Hill Livestock Compensation Allowance, have encouraged overstocking because they are headage based. New replacement subsidies now being developed under the Less Favoured Support Schemes (2001+) have addressed this problem by switching to area based payments. Alongside them are a growing range of payments designed to produce environmental benefits more directly. Agreements over the management of farm land lying with Sites of Special Scientific Interest can involve payments. With

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146 Ecosvstems and Sustainable Development IV

517,322 ha of land now incorporated into Sites of Special Scientific Interest in the Highland region alone, these payments form a sizeable investment in habitat management. Environment-friendly farming with reduced stocking and more sensitive grazing regimes has also been achieved through the designation of Environmentally Sensitive Areas (1987+), like the Breadalbane ESA, an ESA of over 180,000 ha in the southern Highlands. Further agri-environment support, decoupling subsidies from production, is being developed through a new all- embracing scheme, the Rural Stewardship Scheme (1999+).

5 Conclusion

Despite the compelling role played by climate and topography in their development, most European mountain habitats are semi-natural habitats whose natural succession has been modified or deflected in some way by the fact they form part of human resource systems, albeit habitats along the upper edge of those resource systems. By virtue of their position, mountain habitats have a long history of flux, both natural flux (i.e. climate change, acid deposition) and anthropogenically-induced flux (i.e. expansion and contraction of population, shift from common to private resource systems, shift from subsistence to commercial systems). If we add to this flux the insight of much recent work pointing to the sensitivity of mountain habitats and plant species to even slight changes in land management or grazing practice, then it is clear that mountain habitats have never been wholly stable ecosystems. Prior to the nineteenth century, this flux is likely to have worked positively, at least for some areas. Through a succession of phases when alpine species extended their range and phases when subalpine species extended theirs, some mountain areas acquired habitats that mixed both. Likewise, human strategies of resource use, by creating regular patterns of stock movement across a range of habitats and regular cycles of disturbance through hay mowing, etc., helped develop such species enrichment further. But if the net effect of flux before the nineteenth century was species enrichment, the net effect of more recent change has been to favour species impoverishment. By its very nature, commercial or market driven farming leads to the simplification of habitats, with fewer types of stock, the re- seeding of pastures, the draining of others, heavier stocking and the contraction of open range and shared grazings all having their effect. If the residual biodiversity of threatened mountain habitats is to be stabilised and re-expanded, we need land management systems that restores a diversity of husbandry practice in a diversity of situations, from mixed stocking and hay meadowing to the regular seasonal movement of stock across habitats that can serve as sources of potential of propagules. Arguably, such a diversity is best achieved not by so- called modulation, trimming a fraction of production support away from hill farm output and devoting it to agri-environment schemes, but by re-assigning all farm support in key areas to agri-environment ends so that the economics of restoring traditional land management practices are taken out of the marketplace.

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Ecosvstems and Sustainable Development IV 147

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Aas, B. and Faarlund, T. The present and the Holocene subalpine birch belt in Norwuy. Special issue. EFS Project. European Palaeoclimate and Man. 13. European Science Foundation. Gustav Fischer Verlag: Stuttgart, 1996. Birks, J.H.B., 'Long-term ecological change in the British Uplands' (Chapter 3) M.B.Usher and D.B.A. Thompson (eds.), Ecological Change in the Uplands, British Ecological Society: Oxford, special publication no.7, pp. 37-56, 1988; MacKenzie, N.A. The native woodlands. History, decline and present status. The Ecology, Land Use and Conservation of the Cairngorms, ed. C. Gimingham, Packard Publishing Ltd: Chichester, pp. 107-119,2002. Ratcliffe, D.A., and Thompson, D.B.A. The British Uplands: their ecological significance (chapter 2). Ecological Change in the Uplands, eds. M.B. Usher and D.B.A. Thompson. British Ecological Society, special publication no.7: Oxford, pp. 9-36, 1988. Rodwell, J.S. (ed.). British plant communities. Volume 2. Mires and

heaths. Cambridge University Press: Cambridge, 1992. Reinton, L. Saeterbruket I Noreg I. Saetertypur og driftsformer. Institutt for Sammenlignende Kulturforskning, H. Aschehoug and CO. (W. Nygaard): Oslo, 1955. Moe, D., Indrelid, S & Fasteland, A. The Halne area, Hardangervidda. Use of a high mountain area during 5000 years - an interdisciplinary case study (chapter 30). The Cultural Landscape - Past, Present and Future. eds. Birks, H.H., Birks, H.J.B., Kaland, P.E. and Moe, D. Cambridge University Press: Cambridge, pp. 429-444, 1988; Kaland, P.E. Pollenanalytiske undersokelser utenfor boplassen i Kotedalen. In: Kotedalen - en boplass gjemnnom 5000 br. Bind 2. Natuwitenskapelige undersokelser. eds. Hjelle, K.L., Hufthammer, A.K., Kaland, P.E. Olsen, A.B. & Soltvedt, E.C., Universitetet i Bergen: Bergen, pp. 65-89, 1992. For example, Mather, A.S. Pre-1745 land use and conservation in a Highland glen. Scottish Geographical Magazine, 86, pp. 159-69, 1970. Olsson, E.G.A., Austrheim, G. and Grenne, S.N. Landscape change patterns in mountains, land use and environmental diversity, Mid-Norway, 1960-1993. Landscape Ecology, 15, pp. 155-70,2000. Ibid, pp.369-79. Aas, B. and Faarlund, T. The present and the Holocene subalpine birch belt in Norway. Special issue. EFS Project. European Palaeoclimate and Man. 13. European Science Foundation. Gustav Fischer Verlag: Stuttgart, 1996. Austrheim, G, Olsson, E.G.A, Grontvedt, E. Land-use impact on plant communities in semi-natural sub-alpine grasslands of Budalen, central Norway. Biological Conservation 87, pp. 369-379, 1999. Ibid. DD. 369-79.

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[l31 Olsson, E.G.A., & Myklebost, H. Does grazing increase biological diversity in mountains? Population differences in Gentiana nivalis L. in alpine and subalpine habitats in Norwegian mountains. Submitted to Biological Conservation.

[l41 Dodgshon, R.A. From Chiefs to Landlords, Edinburgh University Press: Edinburgh, 1998.

[ l 51 Based on Mcarthur, M.M. Survey of Lochtayside 1769. Scottish History Society: Edinburgh, 3rd series, xxvii, 1936; Statistics of the Annexed Estates 1755-56. HMSO: Edinburgh, 1973.

[l61 Dodgshon, R.A. & Olsson, E.G. Productivity and nutrient use in eighteenth-century Scottish Highland townships. Geografiska Annaler 70B, pp. 39-51, 1988.

117) Based on Scottish Record, Edinburgh, GD112/9143 and GD1 12/16/12/3/l. [l81 Macklin, M.G., Bonsall, C., Davies, F.M. and Robinson, M.R. Human-

environmental interactions during the Holocene: new data and interpretations from the Oban area. The Holocene, 10, pp. 109-21, 2000.

[l91 Smout, T.C. Nature Contested. Environmental History in Scotland and Northern England Since 1600. Edinburgh University Press: Edinburgh, 2000.

[20] Stevenson, A.C. and Thompson, D.B.A. Long-term 'Calluna decline' in upland Britain and Ireland: paleoecological evidence. The Holocene, 3, pp. 70-6, 1993; Stevenson, A.C. and Birks, H.J.B. Heaths and moorland: long- term ecological changes, and interactions with climate and people (chapter 21). Heaths and Moorland: Cultural Landscapes, eds.D.B.A. Thompson, A.J. Hester and M.B. Usher (eds.), HMSO: Edinburgh, pp. 224-39, 1995.

[21] Typical early references to heather burning are provided by Scottish Record Office, GD501159, March 22nd, 1622 and GD50113611, April 17*, 1634.

[22] Fischer, M. and Wipf, S. Effect of low-intensity grazing on the species-rich vegetation of traditionally mown subalpine meadows. Biological Conservation, 104, pp. 1-1 1, 2002.

[23] Olsson, E.G.A. The Scandinavian mountains reflecting human subsistence use of the past -what are the options for future sustainability?, submitted to Mountain Research and Development.

[24] Berlin, G. Semi-natural meadows in southern Sweden - changes over time and the relationship between nitrogen supply and management. PhD thesis in Plant Ecology, Department of Ecology, University of Lund: Lund. 1998.

[25] Agricultural Statistics Norway. 1997. Agricultural statistics. Oslo. [26] Olsson. E.G.A. The Scandinavian mountains reflecting human subsistence

use of the past - what are the options for future sustainability? Submitted to Mountain Research and Development.

[27] Endresen, M. Effekter av husdyrsbeite p6 vegetasjonen I subalpine enger i Griningsdalen, Jotunheimen. Injluence of livestock grazing on subalpine mountain landscape and vegetation. Hovedfagsoppgave. Botanisk institutt. NTNU: Trondheim, 2001.

[28] Agricultural Statistics Norway. Agricultural statistics: Oslo, 1997.

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[29] Olsson, E.G.A., Austrheim, G. and Grenne, S.N. Landscape change patterns in mountains, land use and environmental diversity, Mid-Norway, 1960-1993. Landscape Ecology, 15, pp. 155-70,2000.

[30] Aas, B. and Faarlund, T. The present and the Holocene subalpine birch belt in Norway. Special issue. EFS Project. European Palaeoclimate and Man. 13. European Science Foundation. Gustav Fischer Verlag: Stuttgart, 1996.

[31] Olsson, E.G.A., Austrheim, G. and Grenne, S.N. Landscape change patterns in mountains, land use and environmental diversity, Mid-Norway, 1960-1993. Landscape Ecology, 15, pp. 155-70,2000.

[32] Stevenson, A.C. and Thompson, D.B.A. Long-term 'Calluna decline' in upland Britain and Ireland: paleoecological evidence. The Holocene, 3, pp. 70-6, 1993.

[33] Sydes, C. and Miller, G.R. Range management and nature conservation in upland Britain (chapter 25). Heaths and Moorland: Cultural Landscapes, eds.D.B.A. Thompson, A.J. Hester and M.B. Usher, HMSO: Edinburgh, pp. 323-37, 1995.

[34] Mather, AS. The inter-relationship between afforestation and agriculture in Scotland. Scottish Geographical Magazine, 112, 1996.

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