Woodland biodiversity, palaeo-human ecology and some implications for conservation management

Journal of Biogeography, 26, 1, 33–43

Woodland biodiversity, palaeo-human ecologyand some implications for conservationmanagementRichard Tipping∗, James Buchanan, Althea Davies and Eileen Tisdall Department of

Environmental Science, University of Stirling, Stirling FK 9 4 LA, U.K.

AbstractThis paper appraises the role of prehistoric human societies in modifying the natural

woodland biodiversity of several regions throughout Scotland. It draws attention to the

likelihood that, contrary to popular belief, biodiversity was enhanced by interactions

between agricultural communities and woodlands. The possible purposefulness of such

interactions is discussed.

Mindful of the present concern to re-establish ‘native’ woodlands in many parts of northern

Britain, we then review three approaches to the identification of native tree types in Scotland,

and conclude that insufficient attention in reafforestation schemes is being paid to (a) the

palaeoecological record as a record of native woodland or (b) the likely former high

taxonomic diversity of woodlands. Both these failings will lead to the creation of new

woodlands lacking in species diversity.

Keywords‘Native’ woodland, palaeoecological record, Scotland, woodland biodiversity.

INTRODUCTION 6000 years ago are, for some purposes, inappropriate and

are based on false assumptions. Scotland is also the principalWe wish to address two major issues in this paper. The first

recipient of grants for the ‘restoration’ and recreation ofof these is the assumption that human societies are, and have

‘native’ woodlands.always been, guilty of reductions in plant species diversity. We

We use palynological data to make assertions concerningwill present three case studies which suggest that tree species

past biodiversity. As Odgaard (1999) indicates, this is fardiversity has, in the past, been enhanced by human activity.

from straightforward. Biodiversity as a concept is also farWe will briefly consider the mechanisms by which this occurred,

from well-defined (Gaston, 1996). Here we adopt Harper &and whether this enhancement could have been deliberately

Hawksworth’s (1994) definition of ‘organismal diversity’ asmanipulated.

the scale of our observations. Although we discuss woodlandOur second purpose is to highlight what we see as deficiencies

diversity, we cannot assume that the appearance of treein the current selection of trees being planted in the drive to

taxa represents the colonization by associated ground-layerre-establish ‘native’ woodlands. We identify inadequacies in

communities; current views on tree migration stress itsthis procedure that will lead, unless revised, to woodlands

individualistic nature (Birks, 1989; Delcourt & Delcourt,lacking their original diversity, which is not the intended policy

1991). This is the reason for our not operating at the levelbehind these woodland schemes.

of ‘ecological diversity’ (Harper & Hawksworth, 1994).This paper concentrates exclusively on data from Scotland.

Pollen data are often not good indicators of species,This is because much palynological (pollen-analytical) data

through imprecision in our taxonomy (Moore et al., 1991).are available from this coldest and wettest region of the

We concentrate our attention on tree taxa, as for more thanBritish Isles, and it is possible to reconstruct to some degree

most life-form groups in north-west Europe, these are betterthe original distribution of woodland communities (McVean

served by pollen taxonomy (Figs 3,4,5). Given that arboreal& Ratcliffe, 1962; Birks et al., 1975; Birks, 1977, 1988, 1996;

pollen is generally abundantly produced and dispersed, thereBennett, 1989; Tipping, 1994). Later, we shall suggest that

are inevitable uncertainties in determining either the firstthese country-wide maps of the vegetation of the country

appearance in a region of a taxon (Bennett, 1986) or its

local presence. Here we adopt the recommendations of

Huntley & Birks (1983) in defining the proportions of pollen

of a tree taxon likely to represent its local presence.∗Corresponding author.

1999 Blackwell Science Ltd

34 Richard Tipping et al.

Figure 1 The broad pattern of woodland

communities and unforested areas in Scotland

at 6000 cal. BP derived from pollen data

(modified from Tipping, 1994).

THE NATURAL WOODLANDS OF SCOTLAND Isles (Birks & Madsen, 1979; Wilkins, 1984; Bennett et al.,

1990; Birks, 1991; Bennett et al., 1992; Bunting, 1994; Tipping,

1994), and the extent of naturally unforested areas above tree-Figure 1 is the most recent compilation of palynologicallines (Birks, 1988; Bennett, 1989; Tipping, 1994). Nevertheless,data relating to Scottish plant communities before the firstthe major woodland types recognized have not changedmajor impacts on this vegetation by Neolithic agriculturalsignificantly since McVean & Ratcliffe’s (1962) initialcommunities (Tipping, 1994). The time represented in thisreconstructions.diagram is around 6000 cal. years Before Present (BP). This

This does not imply consistency in our understanding oftime-period was determined by previous reconstructions (Birkspast natural woodlands. Much of Scotland does not have aet al., 1975; Bennett, 1989) which considered this to representhigh-quality array of palynological data (Tipping, 1994; Birks,the period of maximum woodland development (below).1996). Large spatial gaps in our knowledge exist: for example,Fig. 1 was compiled to allow comparison with these earliernorth-east Scotland is barely represented by any modern-interpretations. Major differences between reconstructions still

standard reconstructions. The details in the spatial patterningexist in, for instance, the differing interpretations of woodland

cover and species representation in the Western and Northern of woodlands able to be depicted in Fig. 1 are more an artefact

Blackwell Science Ltd 1999, Journal of Biogeography, 26, 33–43

Woodland biodiversity 35

Figure 2 The locations of the three case-

studies and other localities mentiones in the

text.

of the frequency of investigations than the natural variability reconstructions are spatially very imprecise, lack any sort of

subtlety with regard to altitudinal, climatic or edaphicof the woodlands. For example, the woodlands of both Skye

and Mull are able to be sensitively depicted with relation to gradients, and are simplistic.

In addition, there is an assumption in such reconstructionsmountainous areas through the detailed work of Vasari &

Vasari (1968) and Birks & Williams (1983) on Skye & of Lowe that the period 6000 cal. years ago represents the fullest

expression of woodland complexity and diversity. This idea& Walker (1986; Walker & Lowe, 1985, 1987) on Mull. Other

areas are far less favoured by research. In addition, there is no may be applicable to much of southern Britain, but is almost

certainly not appropriate for Scotland. The principal migrationconsideration given to variations in the pollen recruitment

characteristics of pollen sites; Pennington et al’s (Pennington routes for the major tree taxa colonizing the British Isles are

from the south-east and east, from the continent (Birks, 1989).et al., 1972) large lake records cannot be compared directly

with peat records from the same region (Anderson, 1995), Pinus is one of the very few trees which does not accord with

this pattern (Bennett, 1995). The major tree types had clearlyalthough they are given equal weight in Fig. 1. These weaknesses

imply that this and earlier reconstructions of the natural become established in southern Britain before 6000 cal. BP, and

a degree of stability in woodlands may have been met (Bennett,woodlands of Scotland cannot be used in themselves as the

basis for future ‘native tree’ planting programmes. These 1983, 1986), with no major colonizations of new taxa, and



Figure 3 Selected pollen taxa for Yetholm

Loch over the time-period 7000–4000 cal. BP.

disturbances operating within effectively closed systems. This Pinus expanding and contracting more-or-less synchronously

in diverse regions throughout Scotland (Pears, 1968; Bennett,is not the case in Scotland, where Holocene climatic changes,

for example, can be expected to have led to greater disturbances 1984, 1995; Dubois & Ferguson, 1985; Bridge et al., 1990), and

still not stable within the later Holocene, after 4000 cal. BPand consequent taxonomic re-shufflings. Thus we can see



(Gear & Huntley, 1991). Equally, the establishment of Alnus l.p.a. zone B, probably dominated by hazel (Corylus) and birch

(Betula), with alder (Alnus) and oak (Quercus), but not elmis highly complex within Scotland, and has very different times

of expansion between sites (Birks, 1989; Chambers & Elliott, (Ulmus). Human interference is not identified until l.p.a. zones

C and D, when cereal-type pollen (Gramineae > 37l m) and1989; Bennett & Birks, 1990; Tallantire, 1992). Human agency

may have been an important component in maintaining pastoral indicator herbs (Behre, 1981) are recorded. Alnus

pollen percentages expand, and other tree taxa do not decline,woodland instability, and providing niches for increasing

biodiversity. but Ulmus pollen percentages increase to generate a continuous

and prominent curve. The interpretation is that human activity

created sufficient gaps in the existing woodland to allow theCASE STUDIES

eventual colonization of elm in this upland landscape.

The Carn Dubh pollen sequence (Fig. 5) is from a valleyThis section presents three examples where human impact on

woodlands in Scotland appears to have encouraged a greater peat lying at ≈300 m OD in the calcareous-rich Dalradian

schists of the Eastern Grampians (Ferreira, 1959). These uplandsorganismal diversity than existed prior to that impact. The

sites (Fig. 2) are (a) Yetholm Loch in south-east Scotland were dominated by hazel (Corylus), but elm (Ulmus) was also

abundant locally, probably because of base-rich soils. Quercus(Tipping, 1992, 1996) (b) Sourhope, near Yetholm Loch but

deep within the recesses of the Cheviot Hills (Tipping, 1996), (oak) was less significant, and Betula (birch) and pine (Pinus)

may not have grown around the peat basin. It is unlikely thatand (c) Carn Dubh in the East Grampian uplands east of

Pitlochry (Tipping, 1995; Rideout, 1996). The examples are alder (Alnus) grew locally either, until after the initial human

impacts on this landscape, from c. 5800 cal. BP. At this time ataken from time periods after the first settlement in these areas

of farming communities. This is to emphasize the inherent clear elm decline is observed (Fig. 5), and the landscape is

opened, indicated by a rise in wild grasses (Gramineae < 8lminstability of woodlands in Scotland after 6000 cal. BP, and to

more confidently establish human impact as causal in the anl-D). At this time also, Alnus percentages expand, and it is

argued (Tipping, 1995) that this marks the local establishmentchanges in tree types – it is easier to identify agricultural

activities from pollen analyses than the more subtle of a large population of alder trees previously present but in

only low numbers. The association with human activity ismodifications of hunter-gatherer communities (Maguire, 1983).

Yetholm Loch is a continuous lowland (150 m OD) Holocene emphasized at this site by the remarkably ‘late’ radiocarbon

date for the local expansion of alder (compare with Birks,sediment and pollen record dated by seven internally consistent

radiocarbon dates (Tipping, unpublished). Fig. 3 is a pollen 1989), well into the Neolithic period.

The patterns in woodland changes suggested from thesediagram of selected taxa plotted against time BP, between 7000

and 4000 cal. BP. This sequence is discussed in detail in Tipping three case-studies are clear. It is argued that human activity

facilitated the enrichment of existing natural woodlands with(1996); here we concentrate on changes in tree taxa occurring

at the characteristic elm (Ulmus) decline, in local pollen new tree taxa. This is not a new argument. The greater

abundance of light-demanding trees such as Fraxinus (ash),assemblage (l.p.a.) zone E between 5800 and 5500 cal. BP. A

probable anthropogenic origin for the elm decline has been and that of the majority of the tall shrubs, after anthropogenic

woodland clearance, have been noted for many years (Godwin,proposed (Tipping, 1996), and although a decline in several

tree pollen types is seen, reversing this trend is the appearance 1956). Smith (1984) argued a very similar case to that at Carn

Dubh for the Alnus rise at Newferry in northern Ireland, andof Tilia (lime) pollen. Percentages of this genus are low, but

Tilia is a very low pollen producer (Andersen, 1970; Greig, the idea of disturbance to existing woods has been argued for

the same feature across Britain by Bennett & Birks (1990). Our1981; Huntley & Birks, 1983) and its occurrences at Yetholm

Loch are comparable to those identified at sites in the English discussion is different in suggesting that it is not only the

adventitious r-strategists that can benefit from increasedLake District by Pigott & Huntley (1980), where local presence

is attested. The interpretation (Tipping, 1996, 1997a) is that woodland disturbance, but also more conservative k-strategists

like Tilia and Ulmus.Tilia was able to colonize areas of base-rich soils only following

the decline of Ulmus, which probably originally grew The key factor in these examples is that the woodland was

rendered more diverse following or during human interferencepreferentially on these more nutrient rich soils. Conventionally,

Tilia is believed to have reached a climatically controlled limit than before. The mechanisms taking place at an ecological

scale are difficult to discern. Perhaps the simplest interpretationnear the line of Hadrian’s Wall by 6000 cal. BP (Pigott &

Huntley, 1980; Birks, 1989), but here it is suggested that a is that pre-existing woodlands were, until human disturbance,

capable of resisting invasion by late-arriving or less competitivemigration route existed into southern Scotland, encouraged by

milder climate on the east coast (Moyle, 1980). However, taxa. Climatic constraints may also have changed through time,

as they appear to have done to facilitate the later Holoceneestablishment of sufficient Tilia trees to appear in the pollen

record was only possible following anthropogenic reductions competitive spread of Pinus in northernmost Scotland (Gear

& Huntley, 1991). However, it seems that it was human activityin the competitive pre-existing Ulmus communities.

Sourhope is a valley peat, ≈15 km south of Yetholm Loch, that was the dominant factor, through providing gaps in the

existing woodland large enough or long enough to facilitate150 m higher and with a nutrient-poor soil. Fig. 4 depicts a

selected pollen stratigraphy from 7000 to 4000 cal. BP (Tipping, the colonization of new species from nearby but small seed

sources (Huntley, 1996).1996). The site in a deep valley between the 600 m high Cheviot

Hills supported a much less diverse natural woodland, seen in Woodland clearance by grazing, either intentional or



Figure 4 Selected pollen taxa for Sourhope

over the time-period 7000–4000 cal. BP.



Figure 5 Selected pollen taxa for Carn Dubh

over the time-period 7000–4000 cal. BP.



unintentional (Buckland & Edwards, 1984) would appear to the prevailing ‘crisis’ which is reassuring in that action can be

taken to restore the equilibrium.present difficulties for the establishment of new taxa, as it

would for existing tree types, and the likeliest opportunity for In Scotland, Darling’s arguments for the periods of principal

woodland decline are likely to be incorrect (Tipping, 1993a,b,colonization would occur at periods of reduced or nonexistent

grazing pressures, within woodland regeneration phases. It is 1994). Nevertheless, the current enthusiasm for ‘reforesting

Scotland’, backed by substantial moneys from the Government’sthis sort of context that Smith (1984) identified as critical for

the expansion of alder, at a time when it is competitively equal Millennium Fund, has its roots in Darling’s (1955) view of the

recent landscape impoverishment. This is not to criticise theto other tree species. Nevertheless, at these periods existing

species, particularly the aggressive adventitious colonizers like purpose of these major woodland schemes; there are many

valid arguments, economic and aesthetic, for the establishmentbirch and hazel, would have advantages over trees like elm

and lime. Possibly it was clearance for domestic settlement, of new woodlands (Peterken, 1996). Here we draw attention to

the practical definitions of ‘native’ woodlands being established,with grazing animals kept away from garden plots and houses,

that would have led to conditions most advantageous to the given the implications of increased past biodiversity from pollen

analyses presented above.establishment of new tree types.

Smith (1970) argued for the anthropogenic introduction of Buchanan (1996) identified three potential approaches in

determining the tree species composition of ‘native’ woodlands;hazel to the British Isles in the early Holocene ‘whether by

accident or design’ (p. 82). Current interpretations of the hazel (a) the use of palaeoecology to define past woodland species

(b) the identification of extant tree taxa in an area, and (c) therise tend to refute its association with human activity (Edwards,

1990). However, we should not disregard the potential that application of NVC classifications (Rodwell, 1991; Rodwell &

Patterson, 1994) to define the potential woodland composi-human societies have for the conscious manipulation of

woodlands in ways that are more subtle and complex than tion of an area. These are comparable to Peterken’s (1996)

‘composition’ options, the original-natural, present-natural andsimple tree-removal. Woodland management can be highly

sophisticated (Rackham, 1980). This is an argument presented future-natural options. Current estimates of the selection

processes undertaken by woodland conservationists tend tofor West African forests by Fairhead & Leach (1996), armed

with an extensive series of ethnographic observations on the stress the present-natural woodland; in other words, re-planting

is based on what is growing at the moment. However, since thesevaried ways by which human groups deliberately encourage

particular tree taxa. In north-west Europe we are largely woodland fragments and relicts are by no means established as

being derived from native stands (guidelines only require thedivorced from such traditions, but records abound of the

preferential manipulation of tree species for utilitarian woodland to have been present on maps in c. AD 1750; Walker

& Kirkby, 1987), and may have been managed, and are oftenpurposes, magic and medicine (Fraser, 1932; Darwin, 1996).

Boyd (1988) is one of the few workers to suggest such links, preserved in unnatural locations sheltered from grazing animals,

this option represents the poorest of the three approacheswith regard to macro-plant finds from a late Iron Age

archaeological site in southern Scotland, where the range of (Buchanan, 1996). It can be shown to seriously underestimate

the natural diversity of woodlands (Cairngorms Partnership,tree and tall shrub macro-fossils collected from ditch fills was

at variance with that depicted from pollen analyses (Tipping, 1996).

Buchanan (1996) found the palaeoecological approach more1997b). We should not underestimate the intimacy of the

relation between woods and human groups. encouraging in the identification of woodland diversity in his

field area of Beinn Eighe, in north-west Scotland (Fig. 2). The

numbers of pollen sites in this small region (Durno & McVean,NEW ‘NATIVE’ WOODLANDS AND THE

1959; Birks, 1972; Kerslake, 1982; Taylor unpublished) allowedMAINTENANCE OF ORGANISMAL

the detailed description of woodland habitats through timeBIODIVERSITY

and in space. Many more tree taxa than are currently present

could be regarded as once local and therefore native. However,For many years there has been a commonly held consensus

that human impacts on the woodlands of Scotland have been with regard to re-planting, the very dynamism and inherent

change depicted by pollen analyses present difficulties of choiceentirely damaging. The first part of this paper has hopefully

redressed this impression to some extent. In Scotland the for the woodland conservationist. Each of these ‘past-natural’

woodlands represents an event at a unique point in time andinfluence of Frank Fraser Darling in framing our perceptions

cannot be over-emphasized. His influential writings, most space, and the thoughtless replication of ancient woods, even

if this were possible from palynological data, could notnotably in the West Highland Survey of 1955, have enforced

the view that woodland destruction was largely through accommodate subsequent edaphic, climatic or biotic changes.

‘Future-natural’ options are argued (Rodwell & Patterson,the economic depredations of rapacious landowners and

commercial interests in the last two hundred years. His 1994) to provide predictive models of unhindered vegetation

succession based on what shrub and herb taxa are currentlyarguments are now reproduced in emotive terms by ‘green’

authors such as James Hunter (1994). Smout (1993) has with present. This process makes the assumption that what ground-

layer plants and shrubs remain from a once-extensive woodlandmore constraint traced the influence on ‘green consciousness’

of Darling. The idea that woodland losses are very recent is can reflect what trees were there and so should be there. This

may seem a useful approach (e.g. Pyatt, 1997), but at best itseen elsewhere in the world, and Fairhead & Leach (1996) argue

that, to many conservationists, this provides an immediacy to reflects only the most recent and possibly degenerate woodland,



Anderson, D. (1995) An abrupt mid-Holocene decline of Pinus sylvestrisand this approach shares with the ‘present-natural’ option thein Glen Torridon, north west Scotland: implications for pa-criticisms that this recent woodland need not be natural, needlaeoclimatic change. University of Oxford, School of Geographynot be fully floristically diverse, and need not be adapted toResearch Papers no. 52.future change.

Behre, K.-E. (1981) The interpretation of anthropogenic indicators inThese are difficult but potentially expensive dilemmas for

pollen diagrams Pollen Spores, 23, 225–245.the woodland ecologist (Peterken, 1996). It should be possible

Bennett, K.D. (1983) Postglacial populations of forest trees in Norfolk,to explore approaches to re-afforestation which are more U.K. Nature, 303, 164–167.confidently based on palaeoecological data, and which endorse Bennett, K.D. (1984) The Post-glacial history of Pinus sylvestris in thethe ‘past-natural’ option of Peterken. This is being developed British Isles Quat. Sci. Rev, 3, 133–155.on the National Trust for Scotland (NTS) property of West Bennett, K.D. (1986) Competitive interactions among forest tree popu-

Glen Affric in northern Scotland (Fig. 2) (Davies et al., 1997). lations in Norfolk, England, during the last 10 000 years. New Phytol,

103, 603–620.The basis for this work is the great likelihood that pastBennett, K.D. (1989) A provisional map of forest types for the Britishwoodlands were far more diverse than present estimates of

Isles 5000 years ago. J. Quat. Sci, 4, 141–144.existing woods or NVC communities, would admit.Bennett, K.D. (1995) Postglacial dynamics of pine (Pinus sylvestris)

and pinewoods in Scotland. Our pinewood heritage (ed. by J.R.CONCLUSIONSAldhous), pp. 22–39. SNH, Edinburgh.

This paper has looked back into the past to identify discrete Bennett, K.D. & Birks, H.J.B. (1990) Postglacial history of alder (Alnus

glutinosa (L.) Gaertn.) in the British Isles. J. Quat. Sci, 5, 123–134.but not unusual events where human activity, either intentionalBennett, K.D., Boreham, S., Sharp, M.J. & Switsur, V.R. (1992)or by coincidence, has increased the organismal diversity of

Holocene history of environment, vegetation, and human settlementScotland’s woodlands. The principal reason why such activitiesin Catta Ness, Lunnasting, Shetland J. Ecol, 80, 241–273.should have increased tree taxonomic diversity is through

Bennett, K.D., Fossitt, J.A., Sharp, M.J. & Switsur, V.R. (1990) Holo-woodland clearance for settlement. These anthropogenic

cene vegetational and environmental history at Loch Lang, Southwoodland gaps were of sufficient duration, size or character

Uist, Western Isles, Scotland. New Phytol, 114, 281–298.to provide niches for late-arriving or poorly competetitive taxa.

Birks, H.H. (1972) Studies in the vegetational history of Scotland. III.Past woodlands were thus more diverse than many woodland A radiocarbon-dated pollen diagram from Loch Maree, Ross and

ecologists have acknowledged from examination of the Cromarty. New Phytol, 71, 731–754.depauperate remnants we see today, and human activities were Birks, H.J.B. (1977) The Flandrian forest history of Scotland: a pre-

central to this diversity. This paper then looks forward to the liminary synthesis British Quaternary studies – recent advances (ed.

by F.W. Shotton), pp. 119–136 Clarendon Press, Oxford.major re-afforestation schemes currently being funded, andBirks, H.J.B. (1988) Long-term ecological change in the British uplands.prompts an urgent reconsideration of the approaches to tree-

Ecological change in the Uplands (ed. by M.B. Usher and D.B.A.planting currently being practiced. We may be in danger ofThompson), pp. 37–56. Blackwell, Oxford.planting ‘native’ woodlands that are a sad and impoverished

Birks, H.J.B. (1989) Holocene isochrone maps and patterns of tree-reflection of the past.spreading in the British Isles. J. Biogeogr, 16, 503–540.

Birks, H.J.B. (1991) Floristic and vegetational history of the OuterACKNOWLEDGMENTS

Hebrides. Flora of The Outer Hebrides (ed. by R.J. Pankhurst and

J.M. Mullin), pp. 32–37. London.We are very grateful to the Millennium Fund, through itsBirks, H.J.B. (1996) Great Britain – Scotland. Palaeoecological eventsprogramme ‘Millennium Forests for Scotland’, and to the

during the last 15000 Years (ed. by B.E. Berglund et al.), pp. 95–143.National Trust for Scotland, for their financial support andWiley, Chichester.encouragement in West Glen Affric, and for allowing us the

Birks, H.J.B., Deacon, J. & Peglar, S.M. (1975) Pollen maps for theopportunity for palaeoecological ‘experimentation’. We are

British Isles 5000 years ago. Proc. Roy. Soc. Lond, B, 189, 87–105.particularly pleased to acknowledge the support of Robin

Birks, H.J.B. & Madsen, B.J. (1979) Flandrian vegetational history atTurner (NTS Archaeologist) and James Fenton (NTS

Little Loch Roag, Isle of Lewis, Scotland. J. Ecol, 67, 825–842.Ecologist). Richard Tipping wishes to thank Historic Scotland, Birks, H.J.B. & Williams, W.W. (1983) Late-Quaternary vegetationaland Patrick Ashmore in particular, for support with the work history of the Inner Hebrides. Proceedings Roy. Soc. Edinb, 83B,at the three case-studies discussed, and to Gordon Cook of 269–282.the Scottish Universities Research and Reactor Centre for Boyd, W.E. (1988) Methodological problems in the analysis of fossil

non-artefactual wood assemblages from archaeological sites. J. Arch.radiocarbon dating. Roger Mercer (then of the ArchaeologySci, 15, 603–619.Department, Edinburgh University) and Jim Rideout (then of

Bridge, M.C., Haggart, B.A. & Lowe, J.J. (1990) The history andAOC Scotland) encouraged the work, and Andy Akhtar andpalaeoclimatic significance of subfossil remains of Pinus sylvestrisRichard Kynoch processed the samples. James Buchanan thanksin blanket peats from Scotland. J. Ecol, 78, 77–99.the Forth Valley Enterprise for financial support during his

Buchanan, J. (1996) Re-creating the Caledonian Forest. A comparisonM.Sc. at Stirling University.

of different approaches to determining tree mix in the expansion

and re-creation of native woodlands in Scotland. M.Sc Thesis, StirlingREFERENCES

University.

Buckland, P.C. & Edwards, K.J. (1984) The longevity of pastoralAndersen, S.Th. (1970) The relative pollen productivity and pollen

episodes of clearance activity in pollen diagrams: the role of post-representation of North European trees, and correction factors for

tree pollen spectra. Danm. Geol Unders. Ser II, 96, 1–99. occupation grazing. J. Biogeogr. 11, 243–249.



Bunting, M.J. (1994) Vegetation history of Orkney, Scotland: pollen Moore, P.D., Webb, J.A. & Collinson, M. (1991) Pollen analysis.

Blackwell Scientific, Oxford.records from two small basins in west Mainland. New Phytol, 128,

771–792. Moyle, D.W. (1980) Pollen analysis of peat deposits near Edlingham,

Northumberland. MSc Thesis, University of Durham.Cairngorms Partnership (1996) The Cairngorms assets. Cairngorms

Partnership: Grantown-on-Spey. Odgaard, B. (1999). The use of palynological diversity indices as

measures of vegetation diversity. J. Biogeogr, 26, 7–17.Chambers, F.M. & Elliott, L. (1989) Spread and expansion of Alnus

Mill. in the British Isles: timing, agencies and possible vectors. J. Pears, N.V. (1968) Post-glacial tree-lines of the Cairngorm Mountains,

Scotland Trans. Bot. Soc. Edinb, 40, 361–394.Biogeogr, 16, 541–550.

Darling, F.F. (1955) West Highland Survey. Oxford University Press, Pennington, W., Haworth, E.Y., Bonny, A.P. & Lishman, J.P. (1972)

Lake sediments in northern Scotland. Phil. Trans. Roy. Soc. Lond,Oxford.

Darwin, T. (1996) The Scots Herbal. The plant lore of Scotland. Mercat B, 264, 191–294.

Peterken, G.F. (1996) Natural woodland: ecology and conservation inPress, Edinburgh.

Davies, A., Tipping, R. & Tisdall, E. (1997) Palaeoenvironmental Northern Temperate Regions. Cambridge University Press, Cam-

bridge.perspectives on woodland sustainability. Scottish Woodland History

Discussion Group (ed. by T.C. Smout), pp. 16–20. SNH, Battleby. Pigott, C.D. & Huntley, J.P. (1980) Factors controlling the distribution

of Tilia cordata at the northern limits of its geographic range. II.Delcourt, H.R. & Delcourt, P.A. (1991) Quaternary ecology – a

palaeoecological perspective. Chapman & Hall, London. History in north-west England. New Phytol, 84, 145–164.

Pyatt, D.G. (1997) A site classification for Scottish native woodlands.Dubois, A.D. & Ferguson, D.K. (1985) The climatic history of pine in

the Cairngorms based on radiocarbon dates and tree-ring D/H ratios. Bot. J. Scot, 49, 455–468.

Rackham, O. (1980) Ancient woodland. Cambridge University Press,Rev. Palaeobot. Palynol, 54, 181–185.

Durno, S.E. & McVean, D.N. (1959) Forest history of the Beinn Eighe Cambridge.

Rideout, J. (1996) Carn Dubh, Moulin, Perthshire: survey and ex-Nature Reserve. New Phytol, 58, 228–236.

Edwards, K.J. (1990) Fire and the Scottish Mesolithic: evidence from cavation of an archaeological landscape 1987–90. Proc. Soc. Antiq.

Scotl, 125, 139–195.microscopic charcoal. Contributions to the Mesolithic in Europe

(ed. by P.Vermeesch and P.van Peer), pp. 71–79. Leuven. Rodwell, J.S. (1991) British plant communities. I. Woodlands and

scrub. Cambridge University Press, Cambridge.Fairhead, J. & Leach, M. (1996) Reframing forest history: a radical

reappraisal of the roles of people and climate in west African Rodwell, J.S. & Patterson, G.S. (1994) Creating new native woodlands.

Bulletin HMSO, London.vegetation change. Time-scales and environmental change (ed. by

T.S.Driver and G.P.Chapman), pp. 169–195. Routledge, London. Smith, A.G. (1970) The influence of Mesolithic and Neolithic man on

British vegetation: a discussion. Studies in the vegetational historyFerreira, R.E.C. (1959) Scottish mountain vegetation in relation to

geology. Trans Bot. Soc. Edinb, 37, 229–250. of the British Isles (ed. by D. Walker and R.G. West), pp. 81–96.

Cambridge University Press, Cambridge.Fraser, J.G. (1932) The Golden Bough: a study in magic and religion.

MacMillan & Co, London. Smith, A.G. (1984) Newferry and the Boreal-Atlantic Transition. New

Phytol, 98, 35–55.Gaston, K.J. (1996) What is biodiversity? Biodiversity. A biology of

numbers and difference (ed. by K.J. Gaston), pp. 1–12. Blackwell Smout, C. (1993) The Highlands and the roots of Green consciousness

1750–1990. Scott. Nat. Her. Occ. Paper no. 1. SNH, Edinburgh.Science, Oxford.

Gear, A.J. & Huntley, B. (1991) Rapid changes in the range limits of Tallantire, P.A. (1992) The alder [Alnus glutinosa (L.) Gaertn.] problem

in the British Isles: a third approach to its palaeohistory. New Phytol,Scots Pine 4000 years ago. Science, 251, 544–547.

Godwin, H.E. (1956) History of the British flora. Cambridge University 122, 717–731.

Tipping, R. (1992) The determination of cause in the generation ofPress, Cambridge.

Greig, J. (1981) Past and present lime woods of Europe. Archaeological major prehistoric valley fills in the Cheviot Hills, Anglo-Scottish

Border. Alluvial archaeology in Britain (ed. by S. Needham andaspects of woodland ecology (ed. by M. Bell and S. Limbrey), pp.

23–55. Brit. Arch. Report Int Ser. 146. BAR, Oxford. M.G. Macklin), pp. 111–121. Oxbow Press, Oxford.

Tipping, R. (1993a) The ‘History of the Scottish Forests’ revisited: PartHarper, J.L. & Hawksworth, D.L. (1994) Biodiversity: measurement

and estimation Phil. Trans. Roy. Soc. Lond. B, 345, 5–12. I. Reforesting Scotland, 8, 16–21.

Tipping, R. (1993b) The ‘History of the Scottish Forests’ revisited: PartHibbert, F.A. & Switsur, V.R. (1976) Radiocarbon dating of Flandrian

pollen zones in Wales and northern England. New Phytol, 77, II. Reforesting Scotland, 9, 18–21.

Tipping, R. (1994) The form and fate of Scottish woodlands. Pro-793–807.

Hunter, J. (1994) The other side of sorrow. Mercat Press, Edinburgh. ceedings Soc. Antiq. Scotl. 124, 1–54.

Tipping, R. (1995) Holocene landscape change at Carn Dubh Dubh,Huntley, B. (1996) Quaternary palaeoecology and ecology. Quat. Sci.

Rev, 15, 591–606. near Pitlochry, Perthshire. J. Quat. Sci, 10, 59–75.

Tipping, R. (1996) The neolithic landscapes of the Cheviot Hills andHuntley, B. & Birks, H.J.B. (1983) An atlas of past and present pollen

maps for Europe: 0–13000 years ago, Cambridge University Press, hinterland: palaeoenvironmental evidence. Neolithic studies in no-

man’s land (ed. by P. Frodsham), pp. 17–35. Northern ArchaeologyCambridge.

Kerslake, P.D. (1982) Vegetational history of wooded islands in Scottish Special Edition. NAG, Newcastle.

Tipping, R. (1997a) Vegetation history of southern Scotland. Bot. J.lochs, PhD Thesis, Cambridge University.

Lowe, J.J. & Walker, M.J.C. (1986) Flandrian environmental history Scot, 49, 151–162.

Tipping, R. (1997b) Medieval woodland history from the Scottishof the Isle of Mull, Scotland. II. Pollen analytical data from sites in

western and northern Mull New Phytol, 103, 417–436. Southern Uplands: fine spatial-scale pollen data from a small wood-

land hollow. Woodland history (ed. by C. Smout), pp. 52–75. ScottishMaguire, D.J. (1983) The identification of agricultural activity using

pollen analysis Integrating the subsistence economy (ed. by M. Cultural Press, Edinburgh.

Vasari, Y. & Vasari, A. (1968) Late- and post-glacial macrophyticJones), pp. 5–18. Brit. Arch. Report S181. BAR, Oxford.

McVean, D.N. & Ratcliffe, D.A. (1962) Plant communities of the vegetation in the lochs of northern Scotland. Acta Bot. Fenn, 80,

1–120.Scottish Highland. HMSO, Edinburgh.



Walker, G.J. & Kirkby, K.J. (1987) An historical approach to woodland BIOSKETCHESconservation in Scotland. Scott. For, 41, 87–98.

Walker, M.J.C. & Lowe, J.J. (1985) Flandrian environmental history

of the Isle of Mull. I. Pollen-stratigraphic evidence and radiocarbon Richard Tipping is Senior Lecturer in the Department ofdates from Glen More, south -central Mull. New Phytol, 99, 587–610. Environmental Science at the University of Stirling. He has

Walker, M.J.C. & Lowe, J.J. (1987) Flandrian environmental history published widely on palaeo-environmental analyses,of the Isle of Mull. III. A high-resolution pollen profile from Gribun, principally related to vegetation history, historicalwestern Mull. New Phytol. 106, 333–347. geomorphology and human impact in northern Britain.

Wilkins, D.A. (1984) The Flandrian woods of Lewis (Scotland). J. Ecol. James Buchanan is a researcher with the Scottish72, 251–258. Environmental Protection Agency, following completion of

a Master’s degree in Environmental Management at the

University of Stirling. Althea Davies and Eileen Tisdall are

postgraduate research students exploring different aspects

of Holocene landscape evolution in northern Scotland,

supported by Millennium Forests for Scotland and the

National Trust for Scotland. Their work focuses on natural

and anthropogenic change in west Glen Affric. Althea is

reconstructing the fine-spatial patterning of northern

Scottish native woods and the causes of woodland decline

and loss, through a network of pollen sites. Eileen is

analysing the timing, intensity and scale of climate change

through reconstructions of lake-level change and blanket

bog hydrology.


Documents

Woodland biodiversity, palaeo-human ecology and some implications for conservation management