Gabriele Broll Beate Keplin Mountain Ecosystems Studies in Treeline Ecology Gabriele Broll Beate Keplin (Eds.) Mountain Ecosystems Studies in Treeline Ecology With 96 Figures PROF. DR. GABRIELE BROLL DEPARTMENT OF GEO- AND AGROECOLOGY, ISPA UNIVERSITY OF VECHTA P.O. BOX 1553 49364 VECHTA GERMANY DR. BEATE KEPLIN INSTITUTE OF LANDSCAPE ECOLOGY UNIVERSITY OF MUENSTER ROBERT-KOCH-STR. 26-28 48149 MUENSTER GERMANY E-mail: [email protected] [email protected] ISBN 3-540-24325-9 Springer Berlin Heidelberg New York Library of Congress Control Number: 2004117857 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, broad- casting, reproduction on microfilm or in any other way, and storage in data banks. 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Oelschläger Typesetting: Camera-ready by the Authors Printing: Krips, Meppel Binding: Litges+Dopf, Heppenheim Printed on acid-free paper 30/2132/AO 5 4 3 2 1 0 Preface Mountain ecosystems belong to the most endangered ecosystems in the world. Especially, the treeline ecotone acts as an indicator for environmental change. However, ecological processes in the treeline ecotone are not yet completely understood. The studies provided in this book may contribute to a better understanding of the interactions between vegetation, climate, fauna, and soils in the treeline ecotone. An introductory chapter is given on plants living under extreme conditions, climate change aspects, and methods for characterization of alpine soils. The following articles focus on mountainous areas in America, Europe and Asia. The Working Group on Mountain and Northern Ecosystems at the Institu- te of Landscape Ecology, University of Münster (Germany), has been working on topics related to the treeline ecotone for several decades. This period under the chairmanship of Friedrich-Karl Holtmeier has come to an end now when he retired in 2004. He initiated numerous studies in high mountains and in the North. Many of his students, who became infected by the ‘mountain virus’, will continue these investigations on ecological processes in the altitudinal and northern treeline ecotones. With this compilation of studies in mountain ecosystems we want to thank Friedrich-Karl Holtmeier for his excellent guidance in these cold and fascinating environments. This book could not have been edited without much valuable help of many people. We gratefully acknowledge the interesting contributions of the authors and also the constructive comments from those colleagues who reviewed earlier versions of the manuscripts. We are grateful to Dr. Hans-Jörg Brauckmann, Maja Masanneck and Marta Jacuniak (University of Vechta, Germany) for the careful preparation of the final version of the papers. Not last our thanks go to Dr. Christian Witschel and his staff (Geosciences, Springer Publishers) for the very good cooperation. Gabriele Broll and Beate Keplin Contents General Aspects of Vegetation and Soils in Cold Environments Guideline for Describing Soil Profiles in Mountain Ecosystems 1 Gabriele Broll, Bettina Hiller, Frank Bednorz, Gerald Müller and Thomas Reineke Peripheral Plant Population Survival in Polar Regions 43 Robert M.M. Crawford Climate Change and High Mountain Vegetation Shifts 77 Gian-Reto Walther, Sascha Beißner and Richard Pott Regional Treeline Studies in America Regeneration of Whitebark Pine in the Timberline Ecotone of the Beartooth Plateau, U.S.A.: Spatial Distribution and Responsible Agents 97 Sabine Mellmann-Brown Structure and the Composition of Species in Timberline Ecotones of the Southern Andes 117 William Pollmann and Renate Hildebrand Pocket Gopher – Actor under the Stage. Studies on Niwot Ridge, Colorado Front Range, U.S.A. 153 Hans-Uwe Schütz VIII Contents The Impact of Seed Dispersal by Clark’s Nutcracker on Whitebark Pine: Multi-scale Perspective on a High Mountain Mutualism 181 Diana F. Tomback Regional Treeline Studies in Europe Humus Forms and Reforestation of an Abandoned Pasture at the Alpine Timberline (Upper Engadine, Central Alps, Switzerland) 203 Bettina Hiller and Andreas Müterthies A Discontinuous Tree-ring Record AD 320-1994 from Dividalen, Norway: Inferences on Climate and Treeline History 219 Andreas Joachim Kirchhefer Woodland Recolonisation and Postagricultural Development in Italy 237 Pietro Piussi Regional Treeline Studies in Asia Isolated Mountain Forests in Central Asian Deserts: A Case Study from the Govi Altay, Mongolia 253 Jan Cermak, Lars Opgenoorth and Georg Miehe The Upper Timberline in the Himalayas, Hindu Kush and Karakorum: a Review of Geographical and Ecological Aspects 275 Udo Schickhoff Editors Gabriele Broll Department of Geo- and Agroecology (ISPA) University of Vechta P.O. Box 1553 D-49364 Vechta Germany [email protected] Beate Keplin Institute of Landscape Ecology University of Münster Robert-Koch-Str. 26 D-48149 Münster Germany [email protected] Authors XI Authors Frank Bednorz Institute of Landscape Ecology University of Münster Robert-Koch-Str. 26 D-48149 Münster Germany [email protected] Sascha Beißner Institute of Geobotany University of Hannover Nienburger Str. 17 D-30167 Hannover Germany Gabriele Broll Department of Geo- and Agroecology (ISPA) University of Vechta P.O. Box 1553 D-49364 Vechta Germany [email protected] Jan Cermak Faculty of Geography University of Marburg Deutschhausstraße 10 D-35032 Marburg Germany [email protected] Robert M. M. Crawford Plant Science Laboratory Sir Harold Mitchell Building University of St Andrews St Andrews KY16 9A1 U.K. [email protected] XII Authors Renate Hildebrand Zum Hüggel 3 D-49205 Hasbergen Germany [email protected] Bettina Hiller Centre for Environmental Research University of Münster Mendelstr. 11 D-48149 Münster Germany [email protected] Andreas J. Kirchhefer Department of Biology University of Tromsø N-9037 Tromsø Norway [email protected] Sabine Mellmann-Brown P.O. Box 1044 Cooke City, MT 59020 At time of research affiliated with: U.S.D.A. Forest Service Montana State University Bozeman, MT 59717-0278 U.S.A. [email protected] Georg Miehe Faculty of Geography University of Marburg Deutschhausstraße 10 D-35032 Marburg Germany [email protected] Gerald Müller Haus Vogelsang GmbH Umweltconsulting Vogelsangweg 21-23 D-45711 Datteln-Ahsen Germany [email protected] Authors XIII Andreas Müterthies EFTAS Remote Sensing Ostmarkstr. 92 D-48145 Münster Germany [email protected] Lars Opgenoorth Faculty of Geography University of Marburg Deutschhausstraße 10 D-35032 Marburg Germany [email protected] Pietro Piussi Dipartimento di Scienze e Tecnologie Ambientali Forestali Via S. Bonaventura, 13 50145 Firenze Italy [email protected] William Pollmann Department of Geography University of Colorado Boulder, CO 80309-0260 U.S.A. [email protected] Richard Pott Institute of Geobotany University of Hannover Nienburger Str. 17 D-30167 Hannover Germany [email protected] Thomas Reineke Institute of Geography RWTH-Aachen Templergraben 55 D-52056 Aachen Germany [email protected] XIV Authors Udo Schickhoff Institute of Geography University of Bonn Meckenheimer Allee 166 D-53115 Bonn Germany [email protected] Hans-Uwe Schütz Biologische Station Rieselfelder Münster Coermühle 181 D-48157 Münster Germany [email protected] Diana F. Tomback Department of Biology CB 171 University of Colorado at Denver P.O. Box 173364 Denver, CO 80217 – 3364 U.S.A. [email protected] Gian-Reto Walther Institute of Geobotany University of Hannover Nienburger Str. 17 D-30167 Hannover Germany [email protected] Mountain Ecosystems Studies in Treeline Ecology General Aspects of Vegetation and Soils in Cold Environments Guideline for Describing Soil Profiles in Mountain Ecosystems 1 Guideline for Describing Soil Profiles in Mountain Ecosystems Gabriele Broll, Bettina Hiller, Frank Bednorz, Gerald Müller and Thomas Reineke 1 Introduction This guideline for describing soil profiles in mountainous ecosystems is intended to provide scientists around the world with other than soil science expertise to collect useful soil data such as soil profile descriptions and soil sampling. From the collected soil profile descriptions the scientists should be able to calculate important parameters such as field capacities. In addition, the main objective of this guideline is to streamline methods for soil data collection in mountainous terrain throughout the world, which would result in comparable soil data. The field book Schoeneberger et al. (2002) is recommended as basis for the guideline. This manual was used providing a minimum data set