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Forest and Woodland Ecology Forestand Woodland Ecology Instituteof TerrestrialEcology NaturalEnvironment Research Council á aim64 á Natural Environment Research Council INSTITUTE OF TERRESTRIAL ECOLOGY FOREST AND WOODLAND ECOLOGY An account of research being done in ITE ITE Symposium No. 8 Edited by F.T. LAST ITE, Bush Estate, Penicuik, Midlothian, EH26 OQB and A.S. GARDINER Merlewood Research Station, Grange-over-Sands, Cumbria, LA11 6JU Printed in Great Britain by NERC Reprographic Section © Copyright 1981 Published in 1981 by Institute of Terrestrial Ecology 68 Hills Road Cambridge CB2 1LA ISBN 0 904282 51 1 The frontispiece was drawn by Dr. Julia Wilson, ITE, Edinburgh (Bush). COVER PHOTOGRAPHS (photographers' names in brackets) At the top: flower of Triplochiton scleroxylon, 'obeche' (K.A. Longman). Moving clockwise: Cervuselaphus, mature male red deer (B.W. Staines); Wychwood, Oxfordshire, type 4 in National Woodlands Classification (R.G.H. Bunce); excavated structural roots of Picea sitchensis, Sitka spruce, 16 years after planting in Dumfriesshire, Scotland (E.D. Ford); profile of a podzol (J. Miles); Sitka spruce plantation in Dumfriesshire, Scotland (E.D. Ford); Accipiter nisus, female sparrowhawk at nest with fledglings (R.S. Smith). Centre top: stomata of Scots pine, Pinus sylvestris, in unpolluted (left) and polluted (right) environments (A. Crossley). Centre bottom: mycorrhizal roots of Betula pendula, silver birch (S. Donaldson). The Institute of Terrestrial Ecology (ITE) was established in 1973, from the former Nature Conservancy's research stations and staff, joined later by the Institute of Tree Biology and the Culture Centre of Algae and Protozoa. ITE contributes to, and draws upon, the collective know- ledge of the fourteen sister institutes which make up the Natural Environment Research Council, spanning all environmental sciences. The Institute studies the factors determining the structure, composition and processes of land and freshwater systems, and of individual plant and animal species. It is developing a sounder scientific basis for predicting and modelling environmental trends arising from natural or man-made change. The results of this research are available to those responsible for the protection, management and wise use of our natural resources. Nearly half of the ITE's work is research commissioned by customers, such as the Nature Conservancy Council who require information for wildlife conservation, the Depart- ment of the Environment, the DepartFnent of Energy and the European Economic Community. The remainder is fundamental research supported by NERC. ITE's expertise is widely used by international organisations in oVerseas projects and programmes of research. 1 CONTENTS Introduction 5 Nature of tree and woodland resources 7 1. British woodlands in an European context. 7 R.G.H. Bunce, ITE Merlewood. 2. Scottish deciduous woodlands: a cause for concern? 12 T.W. Parr, ITE Monks Wood. 3. Trees in town and country. 16 J.E.G. Good, ITE Bangor and R.C. Munro, ITE Edinburgh (Bush). 4. Hedgerows as a resource. 20 M.D. Hooper, ITE Monks Wood. Forest and woodland dynamics 24 24 5. Woodland regeneration with examples drawn from investigations of oak woodland. M.W. Shaw, ITE Bangor. 29 6. Changes in the growth and structure of a young plantation forest. E.D. Ford, ITE Edinburgh (Bush). 7. Monitoring in woodlands. 32 J.M. Sykes, ITE Merlewood. 41 8. The recovery of ground vegetation in coppicewood: the significance of buried seed. A.H.F. Brown, ITE Merlewood. 45 9. The conservation of pasture-woodlands. P.T. Harding, ITE Monks Wood. Tree physiology and the influence of the physical environment 49 49 10. Site factors influencing the growth of stands of Scots pine in Great Britain. E.J. White, ITE Edinburgh (Bush). 54 11. The role and use of water in Sitka spruce plantations. R. Milne, ITE Edinburgh (Bush). 58 12. Root growth and its relation to the water economy of a Sitka spruce plantation. J.D. Deans, ITE Edinburgh (Bush). Trees and the deposition of atmospheric pollutants 62 62 13. Dry deposition of airborne pollutants on forests. D. Fowler, ITE Edinburgh (Bush). 66 14. Wet deposition and the movement of pollutants through forests. I.A. Nicholson, ITE Banchory (Brathens). 72 15. An improved method of estimating the contribution of crown leachates to the chemical composition of rain collected beneath trees. K.H. Lakhani, ITE Monks Wood. 2 Trees, their nutrition and effects on soils 75 16. Growth of sycamore and birch in relation to soil chemical properties. 75 A.F. Harrison, ITE Merlewood and D.R. Helliwell, Waterloo Mill, Wooton-under Edge, Glos., G L12 7JN. 17. Nutritional variants of birch. 78 J. Pelham and P.A. Mason, ITE Edinburgh (Bush). 18. Toadstools and trees. 82 P.A. Mason, ITE Edinburgh (Bush). 19. Effects of trees on soils. 85 J. Miles, ITE Banchory (Brathens). Within-species variation and its genetical control 89 20. Betula pubescens—the affinities of types ih the Scottish highlands with those of continental 89 Europe: a study of leaf morphology or the application of morphometrics. A.S. Gardiner, ITE Merlewood. 21. Analysis of the variation within Sitka spruce, lodgepole pine and loblolly pine. 94 M.G.R. Cannel! and C. Cahalan, ITE Edinburgh (Bush). 22. Within-species variants of trees for planting derelict land. 98 J. Wilson, ITE Edinburgh (Bush). Flowering and vegetative propogation 101 23. Possibilities of controlled reproduction in trees. 101 K.A. Longman, ITE Edinburgh (Bush). 24. Physiological approaches to the conservation and improvement of Triplochiton scleroxylon— 105 a West African timber tree. R.R.B. Leakey, ITE Edinburgh (Bush). The fauna, including pests, of woodlands and forests 110 25. Insects on exotic broadleaved trees of the Fagaceae, namely Quercus borealis and species 110 of Nothofagus. R.C. Welch, ITE Monks Wood. 26. The subcortical fauna of oak: Scolytid beetles as potential vectors of oak wilt disease. 116 M.G. Yates, ITE Furzebrook. 27. Trees and the phytophagous insects data bank. 118 L.K. Ward, ITE Monks Wood and D.F. Spalding, ITE Cambridge (Hills Rd.). 28. Population studies of woodland butterflies. 120 E. Pollard, ITE Monks Wood. 29. Factors affecting the breeding of sparrowhawks and the occurrence of their song-bird prey 125 in woodlands. I. Newton, ITE Monks Wood and D. Moss, ITE Bangor. 30. Grey squirrel damage and management. 132 R.E. Kenward, ITE Monks Wood. 31. Abundance of field voles (Microtus agrestis) in conifer plantations. 135 W.N. Charles, ITE Edintiurgh (Craighall Rd.). 3 32. Deer and their woodland habitats. 138 B.W. Staines and D. Welch, ITE Banchory (Brathens). Viruses of trees 143 33. Viruses of trees. 143 II. Cooper and M.L. Edwards, NERC Institute of Virology, 5 South Parks Road, Oxford, OX1 3UB. Lists of common names and latin binomials. 146 158 Addresses of ITE stations. á 5 INTRODUCTION Are we sufficiently alert to the fact that trees affeCt their environments (in addition to being Despite the extensive programme of planting affected by them) and those of associated plants started in the 1920s, Great Britain remains one of and animals? Are the natural assemblages of the least afforested of the developed countries. plants in the uplands, where most of our coni- Its stock of trees, mainly coniferous (Figure 1 ferous plantings are being made, at risk? What of and Plate 27), which occupies about 7% of its the assemblages in former coppicewoods? Can land area, is unlikely to provide more than 10% the effects of afforestation on populations of of the country's requirement of timber and songbirds and their predators be predicted? timber products. By disrupting the former pattern of land uses, shouldn't we have expected that damage by deer Since the 1920s, attitudes have changed. The and squirrels would be intensified? conservation, amenity and landscape interests of tree planters in the 18th and 19th centuries have come to the fore, but with a greater aware- ness of the indirect effects of afforestation, eg its effects on the quantity of water collected from 13•2% Oak Scots Pine 20% 5•3% Beech 2•5% Ash Corsican Pine 3% 2% Sycamore Lodgepole Pine 4% 1% Birch 4%Other Hardwoods - 2-5% Other Conifers Sitka Spruce 20% 3-5% Douglas Fir 7%Hybrid & Japanese Larch 4% European Larch Norway Spruce 8% Fig. 1 Proportion by area, of different tree species la woogland and afforested areas of Great Britain. (Locke, 1970). different catchments, the effects of thinning and Foresters too frequently forget that they are felling on soil erosion and water quality, and dealing with the woody equivalents of ancient hence the development of fish and aquatic plants, einkorn wheats. They are only now beginning to etc. It cannot be denied that concern has been appreciate the full-potential that exists in their triggered by the scale of extant and projected chosen species which, compared with most planting programmes with sometimes dramatic modern agricultural crops, are extremely variable. changes in the mosaic of land uses, including the Fundamental studies of the mechanisms and conservation of semi-natural woodlands. But do strategies of tree growth are now beginning we fully appreciate the diversity of our natural to signal the way in which improvements may woodlands? Do we know how to conserve them, be sought. But in seeking these improvements recognising that all ecosystems are naturally shouldn't more attention be paid, than heretofore, subject to 'make or break'—they are dynamic? In to the exploitation of differences between species? considering the management of forests and It is hardly necessary to recount the damage done woodlands, it is unquestionably desirable to by the aggressive strains of Ceratocystis ulmi,the adopt an integrated approach which takes into Dutch elm pathogen, to our vast stock of English account their environments (atmospheric and elms which are thought to have been vegetatively edaphic), associated shrubs and herbs, harmful propagated (naturally, as in hedgerows, and and beneficial microbes, pests and other animals artificially) from relatively few introduced (invertebrate and vertebrate), in addition to saplings.
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