Ecological Studies, Vol. 167 Analysis and Synthesis Edited by I.T. Baldwin, Jena, Germany M.M. Caldwell, Logan, USA G. Heldmaier, Marburg, Germany O.L. Lange, Würzburg, Germany H. A. Mooney, Stanford, USA E.-D. Schulze, Jena, Germany U. Sommer, Kiel, Germany Ecological Studies Volumes published since 1997 are listed at the end of this book. Springer-Verlag Berlin Heidelberg GmbH L.Nagy G. Grabherr Ch. Körner D.B.A. Thompson (Eds.) Alpine Biodiversity in Europe With 89 Figures, 17 in Color, and 61 Tables Springer Dr. Laszlo Nagy Prof. Dr. Christian Körner McConnell Associates Institute of Botany 41Eildon Street University of Basel Edinburgh, EH3 5JX Schönbeinstr. 6 Scotland 4056 Basel, Switzerland Prof. Dr. Georg Grabherr Prof. Dr. Desmond B.A. Thompson Department of Vegetation Ecology Advisory Services and Conservation Biology Scottish Natural Heritage Institute for Ecology 2 Anderson Place and Nature Conservation Edinburgh, EH6 5NP University of Vienna Scotland Althanstr. 14 1091 Vienna, Austria Cover illustration: Background: Mutterberger Seespitze, Stubai Alps, Austria (3302 m, L. Nagy). Dark green fritillary butterfly Argynnis aglaja, Swiss Alps (M. Raviglione); dotterel Charadrius morinellus, Scottish Highlands (S. Austin); Pyrenean frog Rana pyrenaica> Pyrenees (J. Serra Cobo); alpine daisy Leucanthemopsis alpinum> Mt. Schrankogel, Stubai Alps, Austria (L. Nagy); isard Rupicapra pyrenaica, Pyrenees (J.-P. Martinez-Rica); grasshopper Podismopsis poppiusi, South Urals (Y. Mikhailov) ISSN 0070-8356 Library of Congress Cataloging-in-Publication Data Alpine biodiversity in Europe / L. Nagy... [et al.]. p. cm. - (Ecological studies, ISSN 0070-8356; vol. 167) Includes bibliographical references and index. ISBN 978-3-642-62387-5 ISBN 978-3-642-18967-8 (eBook) DOI 10.1007/978-3-642-18967-8 1. Mountain ecology—Europe. 2. Biological diversity—Europe I. Nagy, L. (Laszlo), 1961-11. Series. QH135.A47 2003 57.5'4'094-dc21 203042503 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permit- ted only under the provisions of the German Copyright Law of September 9,1965, in its current version, and per- missions for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. http://www.springer.de © Springer-Verlag Berlin Heidelberg 2003 Originally published by Springer-Verlag Berlin Heidelberg New York in 2003 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Production: Friedmut Kröner, 69115 Heidelberg, Germany Cover design: design & production GmbH, Heidelberg Typesetting: Kröner, 69115 Heidelberg, Germany 31/3150 YK - 5 4 3 2 1 0 - Printed on acid free paper Preface The United Nations Conference on the Environment and Development (UNCED), held in Rio de Janeiro in 1992, spawned a multitude of pro- grammes aimed at assessing, managing and conserving the earth's biological diversity. One important issue addressed at the conference was the mountain environment. A specific feature of high mountains is the so-called alpine zone, i.e. the treeless regions at the uppermost reaches. Though covering only a very small proportion of the land surface, the alpine zone contains a rela- tively large number of plants, animals, fungi and microbes which are specifi- cally adapted to cold environments. This zone contributes fundamentally to the planet's biodiversity and provides many resources for mountain dwelling as well as lowland people. However, rapid and largely man-made changes are affecting mountain ecosystems, such as soil erosion, losses of habitat and genetic diversity, and climate change, all of which have to be addressed. As stated in the European Community Biodiversity Strategy, "the global scale of biodiversity reduction or losses and the interdependence of different species and ecosystems across national borders demands concerted international action". Managing biodiversity in a rational and sustainable way needs basic knowledge on its qualitative and quantitative aspects at local, regional and global scales. This is particularly true for mountains, which are distributed throughout the world and are indeed hot spots of biodiversity in absolute terms as well as relative to the surrounding lowlands. At the European scale, ALPNET, a European Science Foundation sponsored Alpine Biodiversity Network (1998-2000), was established to synthesise avail- able knowledge on alpine biodiversity. ALPNET was established following an ESF-ICALPE (International Centre for Alpine Environments) exploratory workshop at the Centro di Ecologia Alpina, Monte Bondone, Italy, in 1995. Pri- ority was given to the cataloguing and synthesis of the available knowledge on: (1) biodiversity of alpine environments, (2) spatial patterns along eleva- tion gradients, and (3) vegetation dynamics. As ALPNET was considered to be the first stage of integrating alpine biodiversity, field research was restricted to characterising the bioclimate of the European alpine areas through the use of soil temperature data loggers. This book is the culmination of ALPNET's VI Preface work. Three workshops (in Monte Bondone, Italy; Innsbruck, Austria; Car- gese, Corsica) between 1998-2000 contributed substantially to much of the material presented here, amounting to our current understanding of alpine biodiversity in the high mountain regions of Europe. The ALPNET model has since been adopted on a global scale to establish the Global Mountain Biodiversity Assessment programme of DIVERSITAS (http://www.unibas.ch/gmba).Another new initiative, with many participants fromALPNET,is GLORIA-Europe (http://www.gloria.ac.at.launchedon 1 Jan- uary 2001), the European component of a long-term global observation sys- tem for recording vegetation and temperature at alpine summits. Essentially, this book is about the taxonomic richness of plant and animal communities, their pattern and diversity in space, their temporal changes in species richness and community composition, and the underlying ecosystem processes above the treeline in the European high mountains (except the dry Mediterranean high mountains of Crete, the Canary Islands and the Azores). The book begins with an overview of Europe's alpine areas (Section I, Chap. I). This is followed by a synthesis ofbioclimate of the alpine zone across locations ranging from northern Scandinavia to the Sierra Nevada in a north- south direction and from the Sierra Nevada to the Jakupica Mountains in cen- tral Macedonia, FYR, from west to east. Section I concludes with a physio- graphic and ecological characterisation of the European high mountain regions (Chaps. 3.1-3.10). Section II (Chaps. 4-11) discusses plant taxonomic and community (assemblage) diversity patterns. The scales range from local through the sub-regional (mountain range) to the European continent. The patterns are, in the main, discussed in relation to physiographic factors: lati- tude, longitude, altitude, substratum and topography. Section III (Chaps. 12-17) presents a collection of accounts on high mountain invertebrate species and assemblages, ordered similarly to Section II on plants. The major factors discussed in relation to processes and changing patterns are global change (primarily climate warming) and land use (grazing). Section IV (Chaps. 18-23) focuses on vertebrate distribution, population ecology and ecosystem impacts, especially grazing. Several accounts in Section V (Chaps. 24-28) examine changes ranging from the timberline in the Urals to those observed at the upper limit of plant life in the Alps. Whilst these reports focus on climate amelioration as the main factor of changes (the reports are from areas little influenced by man), the role of herbivores and their manage- ment are at least of equal importance. The closing paper (Chap. 29) is a syn- thesis of alpine biodiversity in space and time. Unless stated otherwise, plant nomenclature in this volume follows Flora Europaea (Tutin et al.1964, 1968-1980). Preface VII Acknowledgements. We thank the ESF for funding the ALPNET network and contribut- ing to the cost of colour illustrations in this volume. The Bristol Foundation, Switzer- land, and Scottish Natural Heritage, Scotland, contributed support towards financing the editing work. References Tutin TG, Heywood VH, Burges NA, Valentine DW, Walters SM, Webb DA (eds) (1964) Flora europaea Vol I. Cambridge University Press, Cambridge Tuting TG, Heywood VH, Burges NA, Moore DM, Valentine DW, Walters SM, Webb DA (eds) (1968-1980) Flora Europaea Vols. 2-5. Cambridge University Press, Cambridge February 2003 The Editors Punta Minuta, Corsica, 2556 m (L. Nagy) Fundamental to the establishment of ALPNET was the unstinting enthusi- asm, driving commitment and knowledge of Jennifer Nagy (nee McConnell), who died shortly before the first workshop in 1998. Our bryologist, Patricia Geissler, was killed in a traffic accident in Geneva in 1999. We shall always remember Jennifer and Patricia. This book is dedicated to the memory of these two excellent scientists, friends and colleagues. They were very special people. Contents I Europe's Alpine Areas 1 An Outline of Europe's Alpine Areas ..... 3 G. GRABHERR, 1. NAGY and D.B.A. THOMPSON 1.1 The Mountains of Europe . . . . . . . . . . . . . . . . . . 5 1.2 An Ecological Classification of Europe's Alpine Areas . . . . 8 1.3 The Regional Accounts (Chaps. 3.1-3.10) . . . . . . . . . .. 11 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11 2 A Bioclimatic Characterisation of Europe's Alpine Areas 13 CH. KORNER, J. PAULSEN and S. PELAEZ-RIEDL 2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . 13 2.2 Site Selection and Methods . . . . . . . . . . . . . . . . . . . 13 2.2.1 What Was Measured and Where? 13 2.2.2 Sensors and Sensor Positioning ............... . 14 2.2.3 Data Analysis ......................... . 16 2.3 Results ............................. . 17 2.3.1 Between-Year and Between-Site Variation 17 2.3.2 Temperature Extremes .................... . 19 2.3.3 The Daily Course of Ground Temperature ......... . 20 2.3.4 The Seasonal Course of Ground Temperature ....... . 20 2.3.5 Seasonal Mean Temperatures ................ . 22 2.3.6 Growing Season Length . . . . . . . . . . . . . . . . . . . . . 22 2.3.7 Thermal Sums . . . . . . . . . . . . . . . . . . . . . . . . . . 23 2.4 Alpine Biota Share a Common Climate Across Europe 26 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 28