Table Of ContentEcological Studies, Vol. 113
Analysis and Synthesis
Edited by
O.L. Lange, Wiirzburg, FRG
H.A. Mooney, Stanford, USA
Ecological Studies
Volumes published since 1989 are listed at the end of this book
F. Stuart Chapin III Christian Korner (Eds.)
Arctic and Alpine Biodiversity:
Patterns, Causes
and Ecosystem Consequences
With 68 Figures
Springer-Verlag
Berlin Heidelberg New York
London Paris Tokyo
Hong Kong Barcelona
Budapest
Prof. Dr. F. Stuart Chapin III
Department of Integrative Biology
University of California
Berkeley, CA 94720-3140
USA
Prof. Dr. Christian Korner
Department of Botany
University of Basel
SchonbeinstraBe 6
4056 Basel
Switzerland
Design of the cover illustration by Mark W. Chapin
ISBN-13: 978-3-642-78968-7 e-ISBN-13: 978-3-642-78966-3
DOl: 10.1007/978-3-642-78966-3
Library of Congress Cataloging-in-Publication Data. Arctic and alpine biodiversity: patterns, causes,
and ecosystem consequences/F. Stuart Chapin III and Christian Korner/eds.) p. cm. - (Ecological
studies; vol. 113) Based on papers from a workshop held in Kongsvold, Norway, Aug. 17-20, 1993.
Includes bibliographical references and index. ISBN-B: 978-3-642-78968-7
I. Biotic communities -Arctic regions - Congresses. 2. Mountain ecology - Congresses.
3. Biological diversity - Congresses. 4. Plant communities - Arctic regions - Ecology - Congresses. 5.
Mountain plants - Ecology - Congresses. I. Chapin, F. Stuart (F. Stuart), III. II. Komer, Christian.
III. Series: Ecological studies; v. 113. QH84. l.A7 1995574.5' 2621 -dc20 94-40221
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Preface
As human populations expand and have increasing access to technol
ogy, two general environmental concerns have arisen. First, human pop
ulations are having increasing impact on the earth system, such that we
are altering the biospheric carbon pools, basic processes of elemental
cycling and the climate system of the earth. Because of time lags and
feedbacks, these processes are not easily reversed. These alterations are
occurring now more rapidly than at any time in the last several million
years. Secondly, human activities are causing changes in the earth's biota
that lead to species extinctions at a rate and magnitude rivaling those of
past geologic extinction events. Although environmental change is
potentially reversible at some time scales, the loss of species is irrevo
cable. Changes in diversity at other scales are also cause for concern.
Habitat fragmentation and declines in population sizes alter genetic di
versity. Loss or introduction of new functional groups, such as nitro
gen fixers or rodents onto islands can strongly alter ecosystem processes.
Changes in landscape diversity through habitat modification and frag
mentation alter the nature of processes within and among vegetation
patches.
Although both ecological changes altering the earth system and the
loss of biotic diversity have been major sources of concern in recent
years, these concerns have been largely independent, with little concern
for the environmental causes the ecosystem consequences of changes in
biodiversity. These two processes are clearly interrelated. Changes in
ecological systems cause changes in diversity. Unfortunately, we know
much less about the converse. What types and magnitudes of change in
diversity alter the way in which ecosystems and the earth system func
tion? What are the processes and circumstances under which this
occurs?
The Scientific Committee on Problems in the Environment (SCOPE)
and the United Nations Environment Programme (UNEP) have re
cently initiated a Global Biodiversity Assessment. An important com
ponent of this assessment is a series of studies on the causes and
consequences of changes in biodiversity. An initial meeting sought to
define the general issues and principles required to predict the nature of
interaction between global change and biodiversity (Schulze and
Mooney 1993, Ecol. Stud. vol. 99). Because it was clear that the nature
VI Preface
of this interaction differs strikingly among ecosystem types, an addi
tional series of meetings was planned to assess the causes and conse
quences of changes in biodiversity in 14 major ecosystems. This book
addresses this issue in arctic and alpine ecosystems, covering about 8%
of the global land area, an area similar to that occupied by boreal forests
or by all the crop land on earth. Arctic and alpine ecosystems are crit
ical because:
1. high latitudes are predicted to undergo more pronounced warming
than other regions of the globe;
2. cold regions are the areas where climatic warming would have the
greatest ecological consequences;
3. high altitudes, due to reduced pressure, are regions where CO should
2
be particularly limiting and where rising CO might strongly stimu
2
late plant growth;
4. arctic ecosystems with their large frozen pools of carbon and methane
may exert strong feedbacks to global climate; and
5. due to their relative simplicity, these ecosystems may show clear
effects of species on ecosystem processes and may, therefore, be
strongly affected by loss or gain of species.
Hence, arctic and alpine ecosystems provide unique insights into
causes and consequences of diversity in general. Furthermore, arctic and
alpine ecosystems are the only biome with a global distribution, mak
ing them ideal for global monitoring of environmental change.
The book is subdivided into three sections, whose objectives are to:
1. describe the patterns of arctic and alpine diversity and suggest causes
for these patterns;
2. develop a framework for predicting how biodiversity may change;
and
3. discuss the ecosystem consequences of changes in biodiverisity.
Most of the chapters of this book are the product of a workshop held
17-20 August 1993 in Kongsvold, Norway. Kongsvold is situated in
Dovre Fjell National Park, where arctic and alpine biomes merge. The
workshop was sponsored by the US National Science Foundation, the
Swiss IGPB National Committee, and various personal travel grants.
The Kongsvold field station, which is one of Europe's oldest ecological
field stations, provided a blend of a long tradition of nature conserva
tion and a natural diversity oflandscapes that inspired many long discus
sions of the future of biodiversity and its consequences in cold
dominated ecosystems.
Berkeley, USA F.S. Chapin III
Basel, Switzerland Ch. Korner
December 1994
Contents
Part I: Patterns and Causes of Diversity
1 Patterns and Causes of Arctic Plant Community Diversity
M.D. Walker . . . . . . . . . . . . . . . . . . . .... 3
1.1 Background and Definitions . . . . . . . . . 3
1.2 Arctic Species Diversity: The First Filter. . . 5
1.3 Biogeographical Patterns Within the Arctic:
The Second Set of Filters . . . . . . . . . 6
1.4 Distribution of Species in Communities:
The Third Set of Filters. . . . . . . . . . 11
1.5 Global Change and Community Dynamics IS
1.6 Conclusions. 15
References. . . . . . . . . . . . . . . . . . 16
2 Causes of Arctic Plant Diversity: Origin and Evolution
D.F. Murray ..................... . 21
2.1 Introduction. 21
2.2 Historical Factors of Arctic Plant Diversity 22
2.3 Evolution of Diversity. 23
2.4 Breeding Mechanisms. 24
2.5 Polyploidy. . 25
2.6 Ecotypes ... 27
2.7 Conclusions . 28
References. . 29
3 Patterns and Causes of Genetic Diversity in Arctic Plants
J.B. McGraw ...................... . 33
3.1 Introduction.................. 33
3.2 Present Patterns of Genetic Diversity ..... 33
3.3 Causes of Present Genetic Diversity Patterns 39
VIII Contents
3.4 Genetic Response to Future Climate Change 40
3.5 Conclusions . 41
References. . . . . . . . . . . . . . . . 42
4 Alpine Plant Diversity: A Global Survey
and Functional Interpretations
Ch. Korner. 45
4.1 Introduction 45
4.2 How Much Land Is Covered by Alpine Ecosystems? 46
4.3 Plant Species Diversity in the Alpine Life Zone 49
4.4 Plant Functional Groups in Alpine Ecosystems . . . 53
4.5 Causes of Alpine Plant Diversity. . . . . . . . . . . 55
4.6 Effects of Biodiversity on Alpine Ecosystem Functioning . 57
4.7 Alpine Biodiversity and Climate Change. 59
4.8 Conclusions . 59
References. . . . . . . . . . . . . . . . . 60
5 Origin and Evolution of the Mountain Flora
in Middle Asia and Neighbouring Mountain Regions
O. Agakhanjanz and S.-W. Breckle 63
5.1 Introduction ............ 63
5.2 Number of Species in the Mountains. 63
5.3 Effective Factors for Evolution. . . 66
5.4 Models for Mountain Florogenesis 68
5.5 The History of Vegetation 74
5.6 Conclusions. 77
References. . . . . . . . . 78
6 Diversity of the Arctic Terrestrial Fauna
Yu.I. Chernov .............. 81
6.1 Number of Species in the Arctic Fauna 81
6.2 Patterns and Causes of Biodiversity Changes
in the Arctic. . . . . . . . . . . . . . . . . . 83
6.3 Taxonomic Composition and Functional Groups . 88
6.4 Biotagenesis. . . . . . . . . . . . . . . . 89
6.5 Species Structure of Arctic Communities 90
6.6 Intraspecies Diversity. . . . . . . . . . . 91
6.7 Correlation Between Forms of Diversity
in Arctic Biomes 92
6.8 Conclusions . 93
References. . . . 93
Contents IX
7 Animal Diversity at High Altitudes
in the Austrian Central Alps
E. Meyer and K. Thaler 97
7.1 Introduction .............. . 97
7.2 Altitudinal Zonation of the Main Groups
of Animals ........... . 97
7.3 Species Numbers of Invertebrates
and Altitudinal Zonation . . . . . 98
7.4 Animal Communities Above the Timberline. 100
7.5 Altitude-Related Changes in Abundance
and Biomass of the Soil Fauna ....... . 102
7.6 Endemism and Vicariance in the Alpine Fauna 104
7.7 Conclusions. 105
References. . . . . . . . . . . . . . . . . . . . 106
Part II: Past, Present, and Future Changes in Diversity
8 Arctic Tundra Biodiversity: A Temporal Perspective
from Late Quaternary Pollen Records
L.B. Brubaker, P.M. Anderson,
and F.S. Hu ....... 111
8.1 Introduction........ . . . . . . . . III
8.2 Late Quaternary Climate History. ........ 112
8.3 What Were the Composition and Regional Patterns
of Tundra During Periods
of Different Late Quaternary Climate? . . . . . . . 113
8.4 What Were the Patterns of Shrub and Tree Invasions
into Tundra When Climate Warmed at the End
of the Last Glacial Period? . . . . . . . . . . . . . 116
8.5 What Major Changes in Ecosystem Processes
Accompanied the Conversion of Tundra to Forest
at the End of the Last Glacial Period? 119
8.6 Conclusion 121
References. . . . . . . . . . . . . . . 122
9 Effects of Mammals on Ecosystem Change
at the Pleistocene-Holocene Boundary
S.A. Zimov, V.I. Chuprynin, A.P. Oreshko,
F.S. Chapin III, M.e. Chapin, and 1.F. Reynolds. 127
9.1 Introduction.................. 127
9.2 Mammalian Effects on Ecosystem Processes. 128
x Contents
9.3 Hypotheses for the Steppe-Tundra Transition 131
9.4 Conclusions 132
References . . . . . . . . . . . . . . . . . . . 133
10 Palaeorecords of Plant Biodiversity in the Alps
B. Ammann 137
10.1 Introduction . 137
10.2 Possibilities and Limitations
of the Fossil Record . . . . . 138
10.3 Temporal Changes in Biodiversity in the
Alps - Three Examples .. 138
10.4 Needs for Future Research 145
10.5 Conclusions 146
References . . . . . . . . . 146
11 Implications for Changes in Arctic Plant Biodiversity
from Environmental Manipulation Experiments
T.V. Callaghan and S. Jonasson. 151
11.1 Introduction . . . . . . . . . . . 151
11.2 Mechanisms of Change in Biodiversity 152
11.3 Responses of Soils to Environmental
Man,ipu1ations: Implications for Plant
Nutrition and Biodiversity. . . . . . . 153
11.4 Direct Responses of Plants to Environmental
Perturbations and Implications for Changes
in Populations, Communities and Biodiversity . 158
11.5 Conclusions 163
References . . . . . . . . . . . . . . . . . . . . 164
12 Patterns and Current Changes in Alpine Plant Diversity
G. Grabherr, M. Gottfried, A. Gruber,
and H. Pauli . 167
12.1 Introduction . 167
12.2 The Altitudinal Limits of Plant Life. 168
12.3 Patterns of Diversity. . . . . . . . . 168
12.4 Effects of Global Warming on Diversity 173
12.5 Conclusions 179
References . . . . . . . . . . . . . . . . 180
Description:As human populations expand and have increasing access to technol ogy, two general environmental concerns have arisen. First, human pop ulations are having increasing impact on the earth system, such that we are altering the biospheric carbon pools, basic processes of elemental cycling and the c
