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
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 15 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, andf.s. Hu Ill 8.1 Introduction HI 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.C. Chapin, and J.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 Manipulations: 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 V Diversity 168 12.4 Effects of Global Warming on Diversity 173 12.5 Conclusions - 179 References 180
Contents XI 13 Anthropogenic Impacts on Biodiversity in the Arctic O.R. Young and F.S. Chapin III..' 183 13.1 Proximate Causes 183 13.2 Social Drivers 186 13.3 Arctic Interactions 188 13.4 Humans as Components of Arctic Ecosystems 192 13.5 Conclusions ' 194 References 195 Part III: Ecosystem Consequences of Diversity 14 Plant Functional Diversity and Resource Control of Primary Production in Alaskan Arctic Tundras G.R. Shaver 199 14.1 Introduction 199 14.2 Resource Uptake 200 14.3 Effects on Community Productivity 205 14.4 Conclusions. '..'.. 208 References 209 15 Direct and Indirect Effects of Plant Species on Biogeochemical Processes in Arctic Ecosystems S.E. Hobbie... 213 15.1 Introduction 213 15.2 Direct Influence over Biogeochemistry 215 15.3 Indirect Influence over Biogeochemistry 217 15.4 Predictions of Species'Effects with Climate Warming 220 15.5 Approaches to Studying the Role of Species in Biogeochemistry 221 15.6 Conclusions 222 References.. 222 16 Causes and Consequences of Plant Functional Diversity in Arctic Ecosystems F.S. Chapin III, S.E. Hobbie, 'M.S. Bret-Harte, and G. Bonan 225 16.1 Introduction 225 16.2 Predicting Species Response to Global Change 225
XII Contents 16.3 Effects of Species on Ecosystem Processes 231 16.4 Species Diversity as Insurance Against Loss of Function 234 16.5 Conclusions 235 References 235 17 Ecosystem Consequences of Microbial Diversity and Community Structure J. Schimel 239 17.1 Introduction 239 17.2 "Broad" Processes 241 17.3 "Narrow" Processes 243 17.4 Stress " 248 17.5 Implications for Tundra 249 17.6 Overall Considerations 250 17.7 Conclusions 250 References 251 18 Diversity of Biomass and Nitrogen Distribution Among Plant Species in Arctic and Alpine Tundra Ecosystems J. Pastor.. 255 18.1 Introduction 255 18.2 Concepts of Diversity 256 18.3 Patterns of Diversity of Carbon and Nitrogen Distribution 258 18.4 Experimental Tests of Controls over Diversity 262 18.5 Towards a Dynamic Model of Diversity in Tundra Ecosystems 265 18.6 Conclusions \267 References 268 19 The Plant-Vertebrate Herbivore Interface in Arctic Ecosystems R.L. Jefferies and J.P. Bryant 271, 19.1 Introduction 271 19.2 Patterns of Vegetation and Vertebrate Herbivory in Arctic Tundra 271 19.3 Vertebrate Herebivory in a Warming Climate: Implications for Tundra Biodiversity 272 19.4 Effects of Anthropogenic Disturbances on Trophic Interactions - A Potential Climatic Change Scenario 274
Contents. XIII 19.5 Conclusions 277 19.6 Summary 278 References 279 20 Insect Diversity, Life History, and Trophic Dynamics in Arctic Streams, with Particular Emphasis on Black Flies (Diptera: Simuliidae) A.E. Hershey, R.W. Merritt, and M.C. Miller 283 20.1 Introduction 283 20.2 Arctic Insect Diversity 283 20.3 Black Fly Life History Features 288 20.4 Trophic Dynamics 289 20.5 Organic Matter Processing by Black Flies in an Arctic Lake Outlet 290 20.6 Summary and Conclusions 292 References 293 21 Land-Water Interactions: The Influence of Terrestrial Diversity on Aquatic Ecosystems G.W. Kling 297 21.1 Introduction 297 21.2 Inputs from Land and the Regulation of Aquatic Systems 297 21.3 Terrestrial Diversity 300 21.4 Movement of Material from Land to Water 303 21.5 Conclusions 305 References... 308 Part IV: A Synthesis 22 Patterns, Causes, Changes, and Consequences of Biodiversity in Arctic and Alpine Ecosystems F.S. Chapin III and Ch. Korner 313 22.1 The Arctic and Alpine Biota 313 22.2 Past, Present, and Future Changes in Biodiversity.... 315 22.3 The Significance of Biodiversity /or Ecosystem Function 316 22.4 Conclusions 320 Subject Index. 321