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Ecology of Red Deer a Research Review Relevant to Their Management in Scotland

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Ecology of Red Deer a Research Review Relevant to Their Management in Scotland Ecologyof RedDeer A researchreview relevant to theirmanagement in Scotland Instituteof TerrestrialEcology Natural EnvironmentResearch Council á á á á á Natural Environment Research Council Institute of Terrestrial Ecology Ecology of Red Deer A research review relevant to their management in Scotland Brian Mitchell, Brian W. Staines and David Welch Institute of Terrestrial Ecology Banchory iv Printed in England by Graphic Art (Cambridge) Ltd. ©Copyright 1977 Published in 1977 by Institute of Terrestrial Ecology 68 Hills Road Cambridge CB2 11LA ISBN 0 904282 090 Authors' address: Institute of Terrestrial Ecology Hill of Brathens Glassel, Banchory Kincardineshire AB3 4BY Telephone 033 02 3434. 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 knowledge of the fourteen sister institutes which make up the Natural Environment Research Council, spanning all the 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 ITE'Swork is research commissioned by customers, such as the Nature Conservancy Council who require information for wildlife conservation, the Forestry Commission and the Department of the Environment. The remainder is fundamental research supported by NERC. ITE's expertise is widely used by international organisations in overseas projects and programmes of research. Contents vii ACKNOWLEDGEMENTS 1 1. INTRODUCTION 2 2. BACKGROUND 2.1 Taxonomy, distribution and variability of 'red deer' 2 2 2.2 A brief biology of the animal 2.3 The status of red deer in Britain 3 .2.4 Importance to man 5 7 3. DISPERSION: FEEDING, SOCIAL AND OTHER BEHAVIOUR AFFECTING DISTRIBUTION 7 3.1 Factors affecting gross distribution 3.2 Factors affecting local distribution 9 3.2.1 Food and nutrition 9 9 3.2.1.1 Nutritional requirements 10 3.2.1.2 Grazing activity 11 3.2.1.3 Species eaten 3.2.1.4 Factors affecting plant selection 12 12 3.2.1.4.1 Extent of selection 14 3.2.1.4.2 Plant habit 14 3.2.1.4.3 Food quality 16 3.2.2 Weather 3.2.2.1 Effect of weather on energy metabolism 16 3.2.2.2 Shelter-seeking behaviour 16 17 3.2.3 Disturbance 3.2.3.1 Interactions with other grazing animals 18 3.2.4 Social behaviour 19 3.2.4.1 Social organisation 19 3.2.4.2 Group size 20 3.2.4.3 Group composition 20 3.2.4.4 Factors affecting the segretation of the sexes 21 3.2.5 Home range 21 3.2.5.1 Size 21 3.2.5.2 Tradition 22 3.2.5.3 Maintenance of home range 22 23 4. IMPACT ON THE HABITAT 4.1 Overview of studies on impact 23 4.1.1 Studies in foreign countries and their relevance to Britain 23 4.1.2 Studies in Britain 24 4.2 Effects on moorlands and grasslands 24 24 4.2.1 Composition changes 25 4.2.2 Succession to woodlands 4.2.3 Controlling impact in moorlands and grasslands 26 26 4.3 Effects on woodlands 4.3.1 Browsing 26 4.3.2 Bark-stripping 28 4.3.2.1 Occurrence 28 4.3.2.2 Consequences 29 4.3.2.3 Causes 30 4.3.3 Thrashing, frayirig and other effects 30 4.3.4 Controlling impact in woodland 30 4.4 Effects on soils 31 4.5 Effects of the habitat-management practices associated with red deer 32 33 4.6 Effects of habitat management and red deer on conservation vi With 11 tables, 4 figures and 24 photographs vfi Acknowledgements It is impractical to name individually all those people who helped us in the preparation of this review. Whilst we intend no discourtesy towards those not included, there are, we feel, some who deserve special mention. First, we are grateful to the ITE Man- agement Group, and in particular to Dr. M.W. Holdgate and Dr. J.P. Dempster for their special interest, guid- ance and criticism during the preparation of the drafts. Next, we must thank those people at home and abroad who were kind enough to respond to our requests for information on work in progress. Amongst these are the several students or ex-students who allowed us to quote their unpublished theses, and other colleagues who sent copies of their unpublished manuscripts. Finally, we thank Janet M. Crisp and Carolyn M. Cummins who helped greatly in the literature searching and proof reading, and all those involved in the typing, especially Elizabeth McDonald. Most photographs were taken by Brian Mitchell, others by Carolyn Cummins (Plate 7B), Tim Parish (Plate 1B; Plate 12, lower), Ian Paterson (Plates 1A, 6 and 7A), Brian Staines (cover, stag and browsed conifer), J.M. Sykes (Plate 8, lower), Adam Watson (Plate 11, upper) and Muriel Welch (cover, landscape). Landscape locations are as follows: cover— Assynt, Sutherland; frontispiece—Isle of Rhum; Plates 1 A and 7A— Glen Feshie, Inverness-shire; Plate 1B —Inverpolly, Wester Ross; Plate 6, lower—Glen Croe, Argyll. 2 1 1. Introduction Complex problems of land-use associated with free- deer are part. Hence deer anatomy is not considered, ranging populations of red deer (Germs elaphus L.) in and morphology and physiology are only partially many parts of the world have stimulated a great deal examined. In the fields of behaviour, performance and of ecological research. Our objective is to assess exist- population dynamics, as also in vegetational dynamics, ing knowledge relevant to the management of red deer we adopt a world-wide perspective in weighing and and their range in Scotland, with a view to highlighting sifting the results of research. Knowledge on other present research needs. large herbivores, particularly deer species, and on the A review is opportune for several reasons. The research grazing process, is also taken into account. studies conceived over a 20-year period by staff of the To achieve a wildlife-management capability it is former Nature Conservancy are now mostly completed, necessary to be able to predict the interactions of, and and future projects will be planned in changed circum- to have control over, the component parts of ecologi- stances. Two new bodies (Institute of Terrestrial Ecol- cal systems; we attempt to evaluate the research ogy and Nature Conservancy Council) have been creat- findings in these respects. Whilst recognising that wild- ed from -the Nature Conservancy, and the 'customer- life-management objectives generally depend on several contractor' principle has been introduced into govern- different sorts of motivating or limiting factors, we give ment research, giving greater weight to practical greater emphasis to the biological than the economic relevance, rather than scientific interest, as a justifica- or sociological considerations. tion of new expenditure. Also significant changes in land-use are occurring, or are likely to occur, in the Because of the importance of red deer to man, there Scottish uplands, with increased recreational pressures, is an extensive literature, with a great deal of sporting increased afforestation, changes in forestry practice, and popular material, only some of which is useful new attitudes towards deer in plantations, and in- in the present context. A bibliography by Kirsch Et creased demand for animal protein, this being reflected Greer (1968) gives over 1,200 references on wapiti or in the rising value of venison and the development of American elk, and 240 on other forms of red deer, deer farming. published up to December 1968; but the total of poten- The basic problem associated with red deer in the tially relevant material is very much greater than this. Scottish uplands concerns the balance between their We therefore mention only selected publications, and value as a resource, and their detrimental effects on have to some extent been obliged to cite their con- habitats and different land-use interests. As with most clusions irrespective of possible technical imperfections other resources, there is a desire to maximise output, and debatable inferences. Techniques of investigation but because deer can roam with relative freedom over are discussed where these are most limiting in either large tracts, of countryside, those benefiting from the research or management. Our scrutiny of the literature resource, in terms of sport, meat production or published since 1974 is less wide-ranging. aesthetic pleasure, do not necessarily experience the We have found it convenient to present first some adverse effects of their presence. Farm crops and forest basic information on the biology, importance and man- plantations can, however, be much damaged by grazing agement of red deer (BM), then the literature survey in and browsing, and even on land specially reserved for the fields of dispersion, feeding and behaviour (BWS), red deer there is a need to consider the well-being impact (DW), and population dynamics and perform- of the habitat. Also it is widely believed that the ance (BM), followed by a discussion on what is known performance of red deer in Scotland is relatively poor, and how the gaps should be filled, together with some and has been declining, because of habitat changes and comments on current management. Finally, the course over-stocking. and achievements of the Nature Conservancy research Our approach is to focus on those aspects of deer on red deer are outlined in an appendix.
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