Mountain Timberlines: Ecology, Patchiness, and Dynamics

375 Pages·2003·17.783 MB·English

by  Friedrich-Karl Holtmeier (auth.)

MOUNTAIN TIMBERLINES
ADVANCES IN GLOBAL CHANGE RESEARCH
VOLUME 14
Editor-in-Chief
Martin Beniston, Institute ofGeography, University of Fribourg, Perolles,
Switzerland
Editorial Advisory Board
B. Allen-Diaz, Department ESPM-Ecosystem Sciences, University ofCalifornia, Berkeley,
CA, U.S.A.
R.S. Bradley, Department of Geosciences, University of Massachusetts, Amherst, MA,
U.S.A.
w. eramer, Department of Global Change and Natural Systems, Potsdam Institute for
Climate Impact Research, Potsdam, Germany.
H.F. Diaz, Climate Diagnostics Center, Oceanic andAtmospheric Research, NOAA,
Boulder, CO, U.S.A.
S. Erkman, Institute for Communication and Analysis of Science and Technology -ICAST,
Geneva, Switzerland.
M. LaI, Centre for Atmospheric Sciences, Indian Institute ofTechnology, New Delhi, India.
U. Luterbacher, The Graduate Institute oflnternational Studies, University ofGeneva,
Geneva, Switzerland.
I. Noble, CRC for Greenhouse Accounting and Research School ofB iological Sciences,
Australian National University, Canberra, Australia.
L. Tessier, Institut Mediterraneen d'Ecologie et PaLeoecologie, Marseille, France.
F. Toth, International Institute for Applied Systems Analysis, Laxenburg, Austria.
M.M. Verstraete, Space Applications Institute, EC Joint Research Centre, Ispra (VA),
Italy.
The tit/es published in this series are listed at the end afthis va/urne.
MOUNTAIN TIMBERLINES
Ecology, Patchiness, and Dynamics
by
Friedrich-Karl Holtmeier
Institute of Landscape Ecology,
University of Münster,
Münster, Germany
KLUWER ACADEMIC PUBLISHERS
DORDRECHT I BOSTON I LONDON
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Printed on acid-free paper
First published 2000 under the tide "Die Höhengrenze der Gebirgswälder" ISBN 3-927889-85-7,
by © Verlag Natur & Wissenschaft, P.O. Box 170209,42624, Solingen, Germany
[email protected]
ISBN 978-94-015-1256-5 ISBN 978-94-015-1254-1 (eBook)
DOI 10.1007/978-94-015-1254-1
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© 2003 Kluwer Academic Publishers and copyright holders
Softcover reprint of the hardcover 1s t edition 2003
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TADLE OF CONTENTS
PREFACE vii
ACKNOWLEDGEMENTS IX
1. INTRODUCTION 1
2. mSTORY AND PRESENT STATE 5
OF TIMBERLINE RESEARCH
2.1 Early timberline research 5
2.2 Modem timberline research 7
3. DEFINITIONS, TERMINOLOGY 11
4. PHYSIOGNOMIC AND ECOLOGICAL
DIFFERENTIATION OF TIMBERLINE 29
4.1 Tree species at timberline 29
4.2 Relationship of timberline elevation to macroclimate,
climate character, and the mass-elevation effect 46
4.3 Ecological conditions and processes at the timberline 55
4.3.1 Reat deficiency 55
4.3.2 Carbon balance 58
4.3.3 Freezing and frost drought 61
4.3.3.1 Frosttolerance and damages 61
4.3.3.2 Frost drought 71
4.3.4 Soil temperature 79
4.3.5 Wind 95
43.6 Snow cover 98
4.3.6.1 Distribution and characteristics of snow cover 98
4.3.6.2 Effects ofthe snow cover on sites 100
4.3.7 Soils 113
vi Mountain Timberlines
4.3.8 Topography 125
4.3.8.1 Slope gradient and geomorphic structure 125
4.3.8.2 Exposure 153
4.3.9 Regeneration 157
4.3.9.1 Seed-produced regeneration 157
4.3.9.2 Vegetative reproduction 180
4.3.10 Influence ofsite conditions on growth form 187
4.3.11 Influence oftrees and tree stands on site conditions 218
4.3.12 Anthropogenic impact on timberline 236
5. TIMBERLINE FLUCTUATIONS 257
5.1 Approach to past and present changes 257
5.2 Conclusions and perspectives 279
REFERENCES 287
INDEX 349
PREFACE
For about 40 years 1 have been engaged in timberline research. Thus, one
could suppose that writing this book should not have been too difficult. It
was harder, however, than expected, and in the end 1 feIt that more questions
had arisen than could be answered within its pages. Perhaps it would have
been easier to write the book twenty years aga and then leave the subject to
mature. But the late Prof. Heinz Ellenberg convinced me to portray a much
needed and complete pieture of what we know of the timberline with special
respect to its great physiognomie, structural and ecological variety.
The first version of this book was published in the German language (Holt
meier, 2000). An extensive summary and translated figure and photo cap
tions and table headings were added to enable non-German speaking people
to make use of the book as weIl. Nevertheless, 1 was very delighted when
Prof. Martin Beniston encouraged me to prepare an English edition for the
series "Advances in Global Change Research", which will guarantee a wider
circulation.
Timberline is a worldwide and very heterogeneous phenomenon, which can
only be presented by way of examples and analyzing different types of tim
berline. My own field experience is necessarily limited to certain timberline
areas, such as the Alps, northern Scandinavia, northern Finland and many
high mountain ranges in the western United States and Canada. However,
my own observations and the results of my and my collaborators research
were essential for developing the concept of the book and became integrated
into the picture of timberline that is presented in the following chapters.
Since the most thorough-going study of the literature cannot compensate for
lack of one' s own field experience and observation, the main discussion is
focused on the timberline regions in the northern hemisphere where 1 have
carried out extensive field research.
Timberline is influenced by many abiotic and biotic factors, often interre
lated in a manifold way. Thus, some factors, such as temperature, wind or
soil, are addressed in >separate chapters, while other agents are described in
the context of more complex ecological processes. Thus, for example, the
role of avian seed dispersal in the forest-alpine tundra ecotone is discussed in
the chapter on seed-produced regeneration (Chapter 4.3.8.1). The influence
viii Mountain Timberlines
of cyclic mass-outbreaks of leaf-eating insects is considered in relation to
timberline tluctuations (Chapter 5).
This book is a conclusive synthesis of my own and my collaborators studies,
and the evaluation of a wealth of literature. Thus, an ample reference list (al
most 1300 titles) is included in this volume. However, in view of the world
wide existence of mountain timberline and of the many possible scientific
approaches to timberline (botany, ecology, climatology, soil science, for
estry, historical science, etc.) this list cannot be expected to be complete, and
surely the reader will miss inclusion of one or another familiar publication.
On the other hand, the references include many papers and books that were
written in German. Anglophone researchers seldom or never refer to these
papers. But these publications are retlecting a long tradition in timberline re
search by German speaking scientists and provide much useful information
that I have tried to make available to those researehers not acquainted with
the extensive 19th and 20th century publications written in German.
Moreover, I was very anxious to provide sufficient illustrations, in particular
photos and sketches out of my field notebooks, for conveying a good visual
impression of the great physiognomie variety of timberline, which also mir
rors its heterogeneity and ecological variety. Instructive illustration is part of
the conception of this book. Most of the photos were taken in my special re
search areas. In addition, friends and colleagues provided a few pictures
from timberline regions that I did not visit by myself. All my photos were
taken exclusively for documentation of timberline, which means that they do
not show landscapes where also timberline was portrayed more or less acci
dentally. There is no other comparable timberline-specific photo collection. I
hope that this collection and the book will contribute to a better understand
ing of mountain "timberline" and its dynamics.
Münster, February 2003
Friedrich-Karl Holtmeier
ACKNOWLEDGEMENTS
This book could not have been written without revising by friends, col
leagues and collaborators. Although being myself close to retirement I still
feel deeply obliged to my academic teacher Prof. Or. Or. h. c. Carl Troll
(1899-1975, Institute of Geography, University of Bonn) and also to Prof.
Or. Ulrich Schweinfurth (Institute of Geography, South Asia Institute, Uni
versity of Heidelberg). Both introduced me into the wide field of landscape
ecology and particularly into timberline problems already when I was a stu
dent. My thanks also go to Prof. Or. Or. h.c. Joachim Blüthgen (1912-1973,
Institute of Geography, University of Münster), himself a pioneer in northern
tree line research, who gave me the first chance to carry out field research in
northern Finnish Lapland in the late 1960s. Moreover, I gratefully remember
my friend Prof. Or. Paavo Kallio (1914-1992, Kevo Subarctic Research
Institute, University of Turku) who watched my tree line studies in Lapland
with great interest and provided use of all facilities at the Kevo Research In
stitute in northern Finnish Lapland.
My special thanks go to those friends and colleagues who essentially contri
buted to developing the conception of this book or supported my fieldwork
by their great hospitality and good company in the field. First of all I men
tion Prof. Or. Gabriele Broll (Division of Geoecology, University of Vechta,
Germany), Or. James B. Benedict (Center for Mountain Archeology, Ward,
Colorado), Mrs. Audrey OeLella-Benedict (Cloudy Ridge Naturalists, Ward,
Colorado) and Or. Wyman C. Schmidt (Research Scientist Emeritus, Inter
mountain Forestry Sciences Laboratory, Montana State University, Boze
man). I am also obliged to Oipl.-Geogr. R. Brandes (Institute for Geography,
University of Erlangen), Prof. Or. Frank Klötzli (Institute of Geobotany,
University of Zürich), Prof. Or. Ernst Löffler (Physical Geography, Univer
sity Saarbrücken), Prof. Or. Hermann Mattes (Institute of Landscape Ecolo
gy, University of Münster), Prof. Or. Richard Pott (Institute of Geobotany,
University of Hannover), Or. M. Oaud Rafiqpoor (Institute of Geography,
University of Bonn), Prof. Or. Michael Richter (Institute of Geography, Uni
versity of Erlangen), Prof. Or. Udo Schickhoff (Institute of Geography and
Botanical Garden, University of Greifswald), Or. Hans-Uwe Schütz (Schöp
pingen), Or. Andreas Vogel (Institute of Landscape Ecology, University of
Münster) and Prof. Or. Masatoshi Yoshino (Institute of Geography, Auchi
University, Toyohashi-City, Japan). Not least I want to thank my collabora
tors Mrs. Oipl.-Landschaftsökologin Kerstin Anschlag, Mrs. Lisette van der
x Mountain Timberlines
Giessen and Dr. Andreas Müterthies (EFTAS, Münster), who were a great
help to me in search for literature and preparation of the many figures and
graphs. Finally my thanks go to my wife Waltraud Holtmeier and my son
Jan-Ulrich Holtmeier for occasional assistance in the field and their patience
with the "fellow lost to timberline". I am also very grateful to the Deutsche
Forschungsgemeinschaft, who funded most of the fieldwork abroad. More
over, I thank Prof. Dr. Martin Beniston (University of Fribourg) and the KIu
wer Acadernic Publishing for publishing the book in trus book series, and
Mrs. Marie Johnson and Mr. Edwin Beschler for help in editing.