Christoph Leuschner Heinz Ellenberg Ecology of Central European Forests Vegetation Ecology of Central Europe Volume I Ecology of Central European Forests Christoph Leuschner • Heinz Ellenberg Ecology of Central European Forests Vegetation Ecology of Central Europe, Volume I Revised and Extended Version of the 6th German Edition Translated by Laura Sutcliffe Christoph Leuschner Heinz Ellenberg (deceased) Plant Ecology University of Göttingen University of Göttingen Göttingen, Germany Göttingen, Germany Translation of the revised and extended German language edition: Vegetation Mitteleuropas mit den Alpen, by Heinz Ellenberg/Christoph Leuschner, © 2010 by Eugen Ulmer KG, Stuttgart, Germany. All Rights Reserved. ISBN 978-3-319-43040-9 ISBN 978-3-319-43042-3 (eBook) DOI 10.1007/978-3-319-43042-3 Library of Congress Control Number: 2017943125 © Springer International Publishing Switzerland 2017 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, 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. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Cover illustration: Beech primeval forest Havešová in Slovakia Printed on acid-free paper This Springer imprint is published by Springer Nature The registered company is Springer International Publishing AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Dedicated to the memory of Heinz Ellenberg (1913–1997), eminent ecologist, teacher and friend Foreword With the publication in 1963 of the first edition of his classical textbook Vegetation Mitteleuropas mit den Alpen (Vegetation Ecology of Central Europe), Heinz Ellenberg produced a unique compendium of the diverse vegetation types of Central Europe and their ecology. This region in the heart of Europe with more than 7000 years of continuous human settlement may well, together with Britain, be the best studied region on earth with respect to its vegetation ecology, visible in a myr- iad of publications on the floristics and plant communities, and the ecology of spe- cies, communities and ecosystems. Covering an area of approximately one million km2 in the northern temperate zone, Central Europe harbours a rich variety of land- scapes from the Baltic and North Sea coasts in the north to the Alps in the south and from eastern France to eastern Poland. This diverse and well-studied region can provide profound insights into the complex interactions between environment, veg- etation and man. Ellenberg’s concept was so successful because he based his analysis on a thor- ough description and sound classification of the plant communities, as many eco- logical statements lose value if they cannot be related to a particular community or vegetation type. It is on this foundation that causal relationships between environ- ment, species composition, community dynamics and ecosystem functioning are explored for the main vegetation types. This truly interdisciplinary approach also includes the historical dimension of the vegetation, its recent change under the impact of human land use pressure and climate change, and current conservation issues. In the more than 50 years since the book’s first appearance, a tremendous amount of relevant research has been carried out, so that the task of providing a comprehen- sive overview of the plant, vegetation and ecosystem ecology of the Central European landscape mosaic has become even more challenging. However, the need for an interdisciplinary synthesis of facts and concepts in plant, vegetation and eco- system ecology is also increasing. Most current environmental problems such as climate change, land use intensification, eutrophication or acid rain involve multi- factorial causation, yet scientific training and research are increasingly narrowed into specialist fields. A broad synthesis of the existing ecological knowledge is vii viii Foreword therefore vital to promote sound understanding of the ecosystem as a whole. This was the motivation for me to take on the task of producing an English version of the book using the 6th German edition as a basis (Ellenberg/Leuschner, 2010: Vegetation Mitteleuropas mit den Alpen, Ulmer Verlag). For the English book, most of the text was revised, a wealth of new facts added and more than 400 recent references incor- porated (now over 5500 cited publications in total). For convenience, the book was split into two volumes (Vol. I, Ecology of Central European Forests; Vol. II, Ecology of Central European Non-Forest Vegetation), whereby Volume I also contains a general introduction into the physical geography, phytogeography and land use his- tory of Central Europe. The book has greatly profited from the careful work and editing of Laura Sutcliffe who, as a native English speaker trained in ecology in Germany, had the task of translating the text. Many of the frameworks, classifications and other terminologies in the German-speaking ecological literature have developed independently of its Anglo-Saxon counterpart, and we spent many hours discussing how best to translate these. During the process of writing, many colleagues, fellow workers, students and technical assistants have supported me by supplying information, pointing at mis- takes, correcting text passages, producing art work and helping to compile the refer- ences lists, especially Markus Hauck, Werner Härdtle, Dietrich Hertel, Michael Runge, Norbert Hölzel, Irmgard Blindow, Helge Walentowski, Yasmin Abou Rajab, Bernhard Schuldt, Jonas Glatthorn, Stefan Kaufmann and Ina C. Meier. I am very grateful to Bernd Raufeisen who produced all the figures and Astrid Röben who helped with the reference list. I would like to thank Valeria Rinaudo and Ineke Ravesloot from the Springer Editorial group and the production team around Prasad Gurunadham and S. Madhuriba for the dedication and support they gave this project. I hope that the two volumes of this book provide the reader with a useful and thought-provoking synthesis of the dynamics and functioning of Central European ecosystems with its characteristic vegetation types, habitats and landscapes. Clearly, such a book represents a subjective selection of topics and can cover only a fraction of the relevant publications, which is a severe shortcoming, but I hope that it serves to direct the reader to the relevant further reading. I am always grateful for correc- tions and any supplementary information. I hope above all that this book will con- tinue to inspire current and future vegetation ecologists and provide a solid information platform for action to protect and value these landscapes. Göttingen, Germany Christoph Leuschner March 2016 Contents of Volume I Part I The Natural Environment and Its History 1 Environmental and Historical Influences on the Vegetation of Central Europe.................................................................................... 3 1.1 The Climate and Phytogeography of Central Europe .................... 3 1.2 An Overview of the Geology and Soils of Central Europe ............ 10 1.3 Historical Influences on the Vegetation of Central Europe ............ 13 2 Life Forms and Growth Types of Central European Plant Species ............................................................................................ 23 2.1 Life Forms ...................................................................................... 23 2.2 Endogenous Rhythms..................................................................... 26 2.3 Plant Anatomy and Morphology .................................................... 27 Part II The Role of Man 3 The Central European Vegetation as the Result of Millennia of Human Activity ............................................................. 31 3.1 Phases of Forest Clearance............................................................. 31 3.2 The Effects on the Vegetation of Low-Intensity Grazing and Woodland Use ............................................................ 40 3.2.1 The Opening Up and Destruction of the Forest ............... 40 3.2.2 The Spread of Pasture Weeds ........................................... 45 3.2.3 Soil Degradation Through Low-Intensity Grazing .......... 47 3.3 From Coppiced Woodlands to Modern Forestry ............................ 52 3.3.1 Coppicing With and Without Standards ........................... 52 3.3.2 High Forest Management ................................................. 58 3.4 The Development of Arable Cultivation and Arable Weeds .......... 60 3.4.1 Pre-industrial Agriculture ................................................. 60 3.4.2 The Effects of Technological Advances on Crop Fields and Low-Intensity Pastures .............................................. 64 ix x Contents of Volume I 3.5 The Development of Meadows, Intensive Pastures and Other Grassland ....................................................................... 65 3.5.1 Straw and Fodder Meadows ............................................. 65 3.5.2 Continuous and Rotational Grazing ................................. 67 3.5.3 Agricultural Biocide Use, Energy Use and Crop Yield.... 69 3.6 Changes in Landscape Hydrology ................................................. 71 3.6.1 Modifications of River Valley Landscapes ...................... 71 3.6.2 The North Sea Dykes and Their Consequences ............... 74 3.6.3 The Destruction of Mires, and Attempts to Restore Them ............................................................... 77 3.6.4 Increasing Exposure of the Vegetation to Drought .......... 79 3.7 Chemical Pollution of the Environment and Its Impact on the Vegetation ............................................................................ 79 3.7.1 Long- and Short-Range Effects of Chemical Pollutants .. 79 3.7.2 Nutrient Enrichment of Soils and Water Bodies .............. 80 3.7.3 Acid Deposition ............................................................... 90 3.7.4 Sulphur Dioxide and Ozone Emissions............................ 92 3.7.5 Emissions of Heavy Metals and Other Substances .......... 98 3.8 Changes in Game Densities and Their Effect on the Vegetation ... 104 3.9 Introduction of Non-native Plant Species ...................................... 105 3.10 Recent Species Losses and Impoverishment of Plant Communities ..................................................................... 106 3.11 The Effects of Recent Climate Change on the Vegetation ............. 108 Part III General Ecology of Central European Forests 4 Abiotic Conditions, Flora, Ecosystem Functions and Recent Human Influence ................................................................ 119 4.1 The Flora of Central European Forests .......................................... 119 4.2 The Geographic Distribution of Forest Vegetation ........................ 119 4.2.1 Zonal, Extrazonal and Azonal Forest Vegetation ............. 119 4.2.2 The Potential Natural Vegetation of Central Europe ........ 123 4.2.3 Altitudinal Belts of Forest Vegetation .............................. 124 4.2.4 Water and Temperature Limitations of Forest Growth ..... 125 4.3 Environmental Conditions and Forest Habitat Classification ........ 127 4.3.1 The Climate of the Forest Interior .................................... 127 4.3.2 Soil Water Regime ........................................................... 134 4.3.3 Soil Chemical Properties .................................................. 141 4.4 Comparative Ecology of Central European Tree Species .............. 150 4.4.1 Important Characteristics of Crown Structure ................. 151 4.4.2 Traits Related to Productivity and Stress Tolerance ........ 151 4.4.3 Nitrogen Acquisition ........................................................ 165 4.4.4 Stress Tolerance ................................................................ 166 4.4.5 Litter Quality and Tree Species Effects on the Soil ......... 179 4.4.6 Competitive Abilities of the Tree Species ........................ 182 Contents of Volume I xi 4.4.7 The Effects of Elevation on Tree Growth ........................ 188 4.4.8 The Influence of Climate on Elevational Changes in Tree Species Composition ............................. 191 4.4.9 Forest Cover in Central Europe and the Current Coverage of Major Tree Species ...................................... 194 4.5 Forest Floor Plants and Shrubs of the Forest Interior: Ecological Niches and Ecological Grouping ................................. 196 4.5.1 Niches of Forest Shrubs ................................................... 196 4.5.2 Ecology of Forest Floor Plants ......................................... 207 4.5.3 The Ecological Grouping of Herbaceous Plants in Central European Broadleaved Forests ............. 243 4.6 Population Ecology of Forest Floor Plants .................................... 245 4.6.1 Phenology ......................................................................... 245 4.6.2 Life Cycles ....................................................................... 249 4.7 Productivity and Cycling of Water and Nutrients .......................... 252 4.7.1 The Biomass and Productivity of the Tree Layer............. 252 4.7.2 The Biomass and Productivity of the Herb Layer ............ 266 4.7.3 Ecosystem Carbon Cycling .............................................. 270 4.7.4 Water Cycling ................................................................... 272 4.7.5 Nutrient Cycling ............................................................... 283 4.8 V egetation Dynamics ..................................................................... 298 4.8.1 Tree Layer Dynamics ....................................................... 298 4.8.2 Fluctuations and Succession in the Herb Layer ............... 298 4.9 R ecent Human Influence ................................................................ 300 4.9.1 Forest Damage in the Past and the Present ...................... 300 4.9.2 Anthropogenic Changes in Forest Soil Conditions .......... 301 4.9.3 Recent Tree Damage and Its Potential Causes ................. 310 4.9.4 Anthropogenic Changes in the Herb Layer and in the Cryptogam and Fungal Flora of Forests .......... 324 4.9.5 Conservation and Restoration of Forests ......................... 337 Part IV Forest and Shrub Formations 5 Beech and Mixed Beech Forests ............................................................ 351 5.1 The Classification of Hardwood Broadleaved Forests ................... 351 5.2 The Classification of Beech Forests in Central and Western Europe ....................................................................... 356 5.3 Beech Forests on Rendzina and Pararendzina ............................... 361 5.3.1 Mesic Limestone Beech Forests (Hordelymo-Fagetum)...................................................... 361 5.3.2 Mull Beech Forests Rich in Wild Garlic .......................... 366 5.3.3 Sedge Beech Forests on Dry Slopes (Carici-Fagetum) .............................................................. 368 5.3.4 Beech Forests Without a Herb Layer (Fagetum nudum) ............................................................. 372