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Re-defining native woodland N. BROWN Oxford Forestry Institute, South Parks Road, Oxford, OX1 3RB, England Summary The term 'native' is widely used to refer to a small number of British tree species and naturally regenerating woodland. This paper proposes that the concept is rooted in an out-dated understanding of vegetation dynamics. Consequently, its definition is arbitrary and ecologically flawed, and should be replaced by terms that make clear the different but important aspects of biogeographical origin, local distinctiveness and conservation value. Introduction Growing concern in Britain over the decline of the original woodland cover to less than 1.5 per cent of the land surface has provoked a recent revision of Forestry Commission policy. The new policy seeks to promote the environmental and social benefits of forests. One of its principal objectives has been to protect and expand the area of native woodland. The policy of expansion of the area of native woodland has been put into practice in two ways. First, changes to the Woodland Grant Scheme (Forestry Commission, 1993) have promoted the planting of native species. Second, the Commission has made widely available detailed information on their ecology and silviculture. As a consequence of these measures there has been a dramatic increase in planting of native species for conservation and amenity purposes. While this is to be welcomed the policy needs to be examined critically to ensure that it is achieving sensible results. Many foresters believe that the promotion of native species has gone beyond sound conservation to the point of arbitrary discrimination against exotic species and may have <} Institute of Chartered Foresters, 1997 reinforced the unhelpful distinction between production forests and forests for conservation and amenity. In this review the author examines the ways that the term 'native' has been used by foresters and ecologists and suggests alternatives which may help overcome the ambiguity and apparent arbitrariness of the term. Conventionally, the word native implies belonging to a natural or original state which has not been created artificially. The word native has been applied to both species and woodland. When applied to a species in Britain, it implies that it is a natural post-glacial colonist of these islands and has not been introduced by humans. When applied to woodland the definition varies with the region of Britain. In England and Wales the term native woodland has been used to refer to communities of naturally occurring species regenerating by natural means, even where the woodland is not of natural origin. However, Stevens and Carlisle (1959) defined the native pine woods of Scotland as the naturally regenerated remnants of the post-glacial pine forest. They proposed that Forestry. Vol. 70, No. 3. 1997 192 FORESTRY woodland was only genuinely native if it was descended from one generation to another by natural means. Following this lead the term is now commonly used in Scotland to refer to woodland that is descended directly, without human interference, from those trees which spread over the British Isles after the last glaciation. In England and Wales the terms natural or semi-natural are more commonly used for this purpose. Native species The identification of native species is based on the assumption that it is possible to disentangle the effects of human interference from biogeography. It is now frequently accepted that native tree species are those which established themselves in Britain after the most recent glacial period, approximately 11 000 years ago, and before significant human civilization. It may be assumed that only a very few tree species continued to occupy these isles throughout the last glacial period. As the botanical slate was all but wiped clean the question of which of the many types of forest that have occupied these lands in the past is used as the base-line for assessing what is natural can be avoided. Genera such as Tsuga and Pterocarya, though present in Britain in previous interglacial periods can now be conveniently classified as exotics. In Britain, climatic history has enabled us to avoid the difficult issue of how far we should go back into biogeographical history to find our native woodland. However, we still face the thorny issue of how close we dare approach the modern day before we decide that human influence is too pervasive and woodland composition no longer natural. Conventionally, civilized human populations are not considered to be natural. Therefore the advent of human civilization in Britain during the Mesolithic has often been used to define the point at which the vegetation ceased to be natural and any further species immigrations became ineligible for native status. It is, however, a far from simple biogeographic task to detect which tree species were natural colonists of the British Isles and which were introduced by humans. It is likely that many species may have colonized and been introduced to Britain on many occasions. While in a forestry context it is common to refer to species as native to Britain, the country does not function as an ecological unit. Species that are technically defined as native to Britain may be incapable of naturally colonizing many areas. More importantly there may be considerable intraspecific local adaptation and consequently important genetic variation in local populations of a widespread species (Soutar and Spencer, 1991; Forrest, 1992). The use of this term at a country scale reflects a bizarrely nationalistic view of biogeography and has neither ecological nor practical value. There may, however, be considerable value in being able to be able to identify whether a particular species is native to a given site within the country. For example, one of the requirements for grants for new woodland under the Woodland Grant Scheme is that 'where nature conservation is an important aim, this will normally mean that you should use the species which are, or might have been, native to the site' (Forestry Authority, 1993). It is far from an easy task to determine whether or not a species belongs naturally at any particular site. The natural range of a species can no longer be easily delineated in Britain. One approach might be to use the original-natural distribution of species (sensu Peterken, 1996) or its putative distribution before humans had a significant influence. While it has been possible to reconstruct the preMesolithic biogeography of many tree species there is considerable uncertainty over the extent to which the biogeography of many of those species which did colonize Britain before the Mesolithic is without human influence. There is speculation that humans may have played a significant role in altering the speed or magnitude of the post-glacial tree invasions and the subsequent dispersal across the country. For example, Rackham (1986) has proposed that Mesolithic communities of humans may have been responsible for the great abundance of hazel found during this period. Indeed, the extinction of many large herbivores, hastened by human hunting, may have been a necessary precondition for the post-glacial re-establishment of woodland. It is clear that while our putative RE-DEFINING NATIVE WOODLAND native species may have colonized one or more parts of the British Isles naturally, their subsequent biogeography and history have been profoundly influenced by human activity. Hence, post-glacial pre-Mesolithic vegetation distribution may not be the original-natural pattern it is purported to be. There may not be a natural pattern at all. It is likely that the present-natural distribution of most species today (that which would prevail if humans had not become a significant influence) would be very different from a putative original-natural distribution (Peterken, 1996). The distribution of vegetation is very sensitive to climate, especially where a species is growing at the limit of its environmental tolerance (Cannell et al., 1989). For example, the decline of lime (Tilia cordata) over the last 5000 years is likely to have been induced by a slight but progressive cooling of the UK climate. The range of any plant species is constantly shifting and it will temporarily overlap with parts of any number of countries as the vagaries of a dynamic environment allow. With the exception of the Caledonian race of Pinus sylvestris and possibly Betula pubescens Ehrh., B. pendula Roth., Populus tremula L., and Sorbus aucuparia L. none of our so-called native trees has occupied Britain for longer than ten thousand years. They are all recent invaders from Europe. Indeed, many are pioneer species with wide ecological tolerance and remarkable geographic ranges. The present day distribution of aspen (Populus tremula L.) for example, stretches from Norway to North Africa (Worrell, 1995). These species are ecological nomads and do not have a fixed geographical range but invade wherever chance creates an opportunity and environmental conditions are suitable. Ingrouille (1995) suggests that accidental events and influences which are not necessarily predictable may have been most influential in determining the pattern of our original-natural vegetation. Plant species did not re-invade Britain as predictable wave fronts but colonized from any number of unpredictable foci where they had fortuitously established. The modern fragmentation of British woodland cover and increase in tree planting have obscured the present-natural tree species distributions. It is very difficult to predict what their 193 natural biogeography would now be. Similarly, it is impossible to do more than speculate as to which of the European species of tree which failed to colonize any part of Britain naturally before the Mesolithic would subsequently have arrived without human assistance. Nor can we predict accurately what the geography of their subsequent invasion might have been. Native woodland The concept of native woodland attaches great value to naturalness. The most rigorous definition of native woodland implies that it is possible to identify components of woodland structure and composition which have been inherited from the original-natural forest cover. A more modern usage of the term still places great value on natural structure and composition and the process of natural regeneration. The relationship between the original-natural, or prehistoric wildwood and the fragments of semi-natural woodland which still exist today is a complex one. In the first half of the twentieth century ecologists believed that, unless disrupted and altered by humans, ecosystems would invariably develop along well defined pathways to a stable, self-sustaining climax community. The natural structure and composition of this community was believed to be determined by the environment alone (Weaver and Clements, 1938). The climax community in which Clementsian succession terminates was viewed as permanent and stable since it was in total harmony with a stable habitat. The concepts of native and climax plant communities share considerable conceptual ground. Recent Forestry Authority recommendations for creating new native woodlands in Britain (Rodwell and Patterson, 1994) encourage foresters to create 'the kind of climax vegetation that would develop . . . if succession was allowed to take its full course'. A similar view is reflected in Peterken's (1996) description of potential-naturalness as 'the state which would develop if people's influence were completely removed and the resulting succession were finished in a single instant. This hypothetical state expresses our understanding of the current potential of the native species on a site under the prevailing climate.' The climax 194 FORESTRY community is seen as natural and its structure and composition are determined by the environment. It is now believed that environmental determinism and the climax community are unrealistic models. Disturbance is an inherent part of all ecosystems. Where the life-span of major organisms in a community exceeds the frequency of disturbance then community composition will bear little relation to the potential environmental climax of a site. This is certainly the case for woodland where the structure and composition of vegetation reflects a continuous process of response to and recovery from disturbance. Since trees are typically very long lived organisms, disturbance events of a magnitude sufficient to influence their survival and reproduction will occur with a frequency well within their life spans. A rare weather event, such as the hurricane which hit southern Britain in 1987, may occur only once in many decades but may have an effect on tree survival that has ramifications for many tree generations. Peterken and Mountford (1996) report how the 1976 drought has had a profound influence on the structure and successional trajectory of Lady Park Wood in Gwent over the last 16 years. As a consequence the composition of natural tree communities may be extremely dynamic from one generation to the next. History is the primary force sculpturing woodland composition. An ecosystem is highly dynamic and cannot be preserved in the same way that our cultural heritage may. Chance patterns of dispersal and survival will determine the pool of species available. Chance weather events, disease, changes in predator pressure and human use will all alter the way woodland regenerates. For example, Jones (1952) attributes the even age of many British beech woods to be the consequence of prolific natural regeneration in very rare warm years. The relative competitive status of different species of tree is constantly shifting as disturbance and a fluctuating forest environment give advantage first to one species and then to another. Present patterns are the product of unique historic events that were neither predictable nor likely to be repeated. Harvey (1995) concludes that its is neither desirable nor possible to manage nature in order to maintain the status quo. Changes in species composition should be accepted as a natural trend. The 'post-glacial pre-Mesolithic' definition of native woodland attempts to fossilize nature. It fails to recognize that the composition of any plant community is continuously changing. Invasions and extinctions are not only natural but are the very essence of woodland ecology. Recommendations such as those of Rodwell and Patterson (1994) that foresters should aspire to re-create native climax woodland may simply re-inforce an ecological myth. Clearly the structure and composition of present-day native woodland is not a relic of a past natural climax community. But it is also unlikely to be a reliable indication of what wildwood might have been like today in the absence of human influence (present-natural sensu Peterken, 1996). Our so-called native woodland is, by its nature, a community in transition. Many of the 35 or so species which were naturally able to colonize this island are invasive species. They have an ecology that is typical of the early arrivals after disturbance. The socalled native British tree flora is an early successional one and it is highly improbable that in the absence of human disturbance it would have remained as a stable community up until the present day. Late successional species would almost certainly have followed these early colonists had forest fragmentation not brought their advance to a premature end. Beech (Fagus sylvatica), hornbeam (Carpinus betulus), lime (Tilia cordata and T. platyphyllos) and yew (Taxus baccata) are the few truly shade tolerant, late successional trees in the native British flora. Beech is believed by some to have naturally invaded parts of southern Britain and is only regarded as native south of a line from the Wash to South Wales (Rodwell and Patterson, 1994). Hornbeam is a hardy species of wide ecological range (Savill, 1991) but is deemed to be native to the south-east of Britain alone. The putative native ranges of these species are highly unlikely to be their present-natural distributions. As Sprugel (1991) has emphasized, the past provides an equivocal guide to what the present might have been without humans. The fact that woodland may not have been subject to direct management or exploitation does not mean that it is free from important human influence. Humans have had profound RE-DEFINING NATIVE WOODLAND indirect effects on the structure and composition of even protected natural areas. The current protected status of much native woodland in Britain may itself mean that trees are no longer exposed to the same disturbance regime that gave rise to their present structure and composition (Hobbs and Huenneke, 1992). The natural levels of wild herbivores in British woodland are unknown. Populations of deer continue to increase to levels where natural regeneration of many native species is impossible. Other herbivores which would have inflicted different types of pressure on tree populations such as beaver (Caster fiber Linn.) and wild boar (Sus scrofa Linn.) have gone extinct (Kirby et al., 1994). Many important processes of human induced environmental change now act at a landscape scale and even protected forest areas are subject to their influence. Changes in the composition of the atmosphere due to pollution are already known to have had a profound effect on natural plant communities (Woodin and Farmer, 1993). It is possible that if climate change continues our native woodland may become unsuited to the prevailing climatic conditions (Billington and Pelham, 1991). The UK Department of the Environment has concluded that the likely increase in mean annual temperatures of 0.5°C over the next decade will bring about noticeable changes in plant and animal distributions. There will be significant movement of species northwards at a rate of at least 50-80 km per decade (DoE, 1996). The number of species in the UK is likely to increase due to an increased rate of immigration. In the face of these ubiquitous processes of human induced change it is untenable to use the concept of native to imply that any present day British woodlands have a structure or composition that is largely a function of natural processes. The traditional view of the climax ecosystem is that once human disruption ceases, nature will take over and succession will slowly return community composition to its original-natural state. This model has been replaced by a view that the trajectory of succession is determined by the starting conditions and that even within a single environment different starting points will lead to very different outcomes. Succession is therefore a chaotic rather than a deterministic 195 process. Past management of British woodland has not been obliterated by the passage of time but has played a central role in determining the path of future natural succession. Spencer (1996) has demonstrated that in the New Forest past exploitation has had an impact on composition that has persisted for many centuries due to major changes in stand dynamics and ecology. During the seventeenth and eighteenth centuries the forest was heavily exploited for oak leading to a major shift to dominance by beech. Beech trees are now reaching the end of their natural life span and are prone to catastrophic disruption from drought and windblow. This in turn has profound consequences for the present day forest disturbance regime and the nature of tree regeneration. Forest composition has not returned to an environmentally determined climax condition. The ramifications of past events continue to determine the future. Nature: a powerful myth The concept of native species depends on the assumption that it is possible to determine whether or not a species belongs naturally at a particular site. Native woodland has been defined as a community of naturally occurring species regenerating by natural means. Naturalness is central to the value attached to native species and woodland. This review has sought to demonstrate that naturalness is an abitrary quality. Peterken (1996) suggests that we should regard naturalness as a continuous variable such that a forest might be classified as being more or less natural. This review has striven to demonstrate that natural process and human influence should not be separated. Humans have had a diverse range of impacts on many ecosystem processes many of which it is no longer possible to quantify. There is a suspicion that a classification of naturalness such as that proposed by Peterken (1996) will rank only conspicuous effects as unnatural. All British woodland reflects the intimate interplay of both natural and human influences. Not only is it naive to believe that the two forces may be disentangled but such an approach denies the very real biological and historical interest in understanding their relation- 196 FORESTRY ship. While conservation biologists clearly recognize the pervasiveness of human influence in British woodland there is a powerful culture of.valuing what is perceived as natural over and above human influenced systems. Cronon (1995) claims that attempts to exclude human influence from the concept of nature is a product of the very social history that it seeks to deny. He suggests that a consumer society which is divorced from the country as a place for production has created a myth that true nature can only occur when humans are not there to despoil it. In this case that myth is the Clementsian climax forest. This vision of nature appeals because it provides a standard against which the failings of a modern society can be judged and is a central tenet of the environmental movement. A romantic urban-based wilderness fantasy has no place for human beings to make a living. It is dangerous because it leaves no place for discovering what a sustainable human place in nature might be. Furthermore it sets an arbitrarily high standard against which cultural and productive landscapes are regarded as inferior. As both Peterken (1996) and Rackham (1986) have pointed out much of the value for nature conservation of ancient woodland may lie in a detailed historical understanding of human impacts. Much of our woodland may be locally distinctive in both structure and composition as a consequence of human use. The grove of wind-torn sycamores sheltering a Pennine farmhouse are a perfect example. These cultural landscapes may well be unique to Britain but their importance is reduced by a forest policy in which native woodlands are prioritized for conservation and amenity values. A need for a new definition The concept of native woodland as it is presently used in British forestry confuses the separate issues of local adaptation, local distinctiveness and species biogeography. The recent development of classifications of native woodlands (Dunlop, 1994; Forestry Commission, 1994) which measure nativeness on the basis of whether woods are natural or planted and composed of appropriate species of local genotype reflects the fact that the term is insufficiently precise to distinguish important woodland types. The term native is misleading as it creates arbitrary and ecologically unsound distinctions. The biogeography and cultural history of a species should be of academic and silvicultural interest not a reason for discrimination. We should adopt a more sophisticated approach to our use and conservation of our tree flora, promoting and conserving woodland for more rationally defined values. For example, there is obvious conservation value in protecting the ancient Caledonian pine forests. However, that value is not a consequence of the fact that the species qualifies for the definition 'native' but because of their' awesome beauty, their habitat value, and the unique genetic structure of the Scottish population. Ecotype The ecological term 'ecotype' is used to describe a population adapted to local environmental circumstances. It may be composed of trees that are genetically distinct or that have been managed in a way that has created a unique or important structure. Thus it is entirely possible for an introduced species to develop a local ecotype. While it is likely that much of the beech woodland in the Chilterns has been planted, the local ecotype has a distinctive white timber and is integral to the landscape. Locally distinctive There is clearly considerable value in being able to identify woodland which is locally distinctive. Such woodland is composed of trees that are an intimate part of the local landscape, regardless of whether or not they are native. Sycamore trees for example, are an integral part of a bleak Pennine farm landscape, as are Scots pines in Breckland. The campaign group Common Ground has promoted the role of trees, particularly apple trees, in bestowing important local character, patterns of planting may give distinctive character, and create an essential component of the cultural landscape. RE-DEFINING NATIVE WOODLAND Natural colonist There is both conservation value and scientific interest in distinguishing the natural colonists in any ecosystem from introduced species. The issue of scale is important in describing the colonization process. Many tree species have been introduced to this country but have subsequently colonized large areas naturally. On the other hand all of the post-glacial, pre-Mesolithic natural colonists of the British Isles have been introduced to a significant part of their present day range. Conclusions There would seem to be no rational, objective criterion by which native woodland in Britain can be distinguished from non-native woodland. British woodland comprises a continuum of naturalness, from the ancient descendants of the prehistoric wildwood to modern, intensive plantations. It is composed of species that range from those recently introduced from other parts of the world to those that have occupied parts of this island since prehistoric times. The decision about where, along these continua the line is drawn between native and non-native is arbitrary and ecologically unsound. It has been open to a wide variety of interpretations for different purposes. The use of the term native has confused the important issues of biogeographic origin, local distinctiveness and conservation value. It is time that a more precise forestry made clear the important difference. British forestry needs to shake off a concept that is jingoistic and rooted in an outdated understanding of vegetation dynamics. Acknowledgements I am grateful to Keith Kirby, Peter Savill, Steve Jennings, Mary Stockdale and Dan Bebber for comments. This paper is partially based on ideas presented at Institute of Chartered Foresters Discussion Meeting on Native and Non-native in British Forestry, Warwick, 31 March-2 April 1995. References Billington, H.L. and Pelham, J. 1991 Genetic variation in the date of budburst in Scottish birch pop- 197 ulations: implications for climate change. Fund. Eco. 5(3), 403-409. Cannell, M.G.R., Grace, J. and Booth, A. 1989 Possible impacts of climatic warming on trees and forests in the United Kingdom: a review. Forestry 62(4), 337-364. Cronon, W. 1995 The trouble with wilderness; or, getting back to the wrong nature. In Cronon, W. (ed.) Uncommon Ground: Towards Re-itwenting Nature. W.W. Norton, New York. 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Savill, P.S. 1991 The Silviculture of Trees Used in British Forestry. CAB International, Wallingford. 198 FORESTRY Soutar, R.G. and Spencer, J.W. 1991 The conservative of genetic variation in Britain's native trees. Forestry 64(1), 1-12. Spencer, J. 1996 Forest exploitation in the New Forest; the biological consequences of a thousand years of forest use and ahuse in lowland England. Paper presented at a conference on Sustaining Ecosystems and People in Temperature and Boreal Forests. Victoria, British Columbia, September 1996. Sprugel, D.G. 1991 Disturbance, equilibrium and environmental variability: what is 'natural' vegetation in a changing environment? Biol. Conserv. 58(1), 1-18. Stevens, H.M. and Carlisle, A. 1959 The Native Pinewoods of Scotland. Oliver & Boyd, Edinburgh. Weaver, J.E. and Clements, F.E. 1938 Plant Ecology. McGraw-Hill, New York. 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