Arctic and Alpine Biodiversity: Patterns, Causes and Ecosystem Consequences

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1 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

2 Contents Part I: Patterns and Causes of Diversity 1 Patterns and Causes of Arctic Plant Community Diversity M.D.Walker Background and Definitions Arctic Species Diversity: The First Filter Biogeographical Patterns Within the Arctic: The Second Set of Filters Distribution of Species in Communities: The Third Set of Filters Global Change and Community Dynamics Conclusions 15 References 16 2 Causes of Arctic Plant Diversity: Origin and Evolution D.F. Murray Introduction ' Historical Factors of Arctic Plant Diversity Evolution of Diversity Breeding Mechanisms Polyploidy Ecotypes Conclusions,. 28 References 29 3 Patterns and Causes of Genetic Diversity in Arctic Plants J.B. McGraw Introduction Present Patterns of Genetic Diversity Causes of Present Genetic Diversity Patterns 39

3 VIII Contents 3.4 Genetic Response to Future Climate Change Conclusions 41 References 42 4 Alpine Plant Diversity: A Global Survey and Functional Interpretations Ch. Korner Introduction How Much Land Is Covered by Alpine Ecosystems? Plant Species Diversity in the Alpine Life Zone Plant Functional Groups in Alpine Ecosystems Causes of Alpine Plant Diversity Effects of Biodiversity on Alpine Ecosystem Functioning Alpine Biodiversity and Climate Change 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 Introduction Number of Species in the Mountains Effective Factors for Evolution Models for Mountain Florogenesis The History of Vegetation Conclusions 77 References 78 6 Diversity of the Arctic Terrestrial Fauna Yu.I. Chernov Number of Species in the Arctic Fauna Patterns and Causes of Biodiversity Changes in the Arctic Taxonomic Composition and Functional Groups Biotagenesis Species Structure of Arctic Communities Intraspecies Diversity Correlation Between Forms of Diversity in Arctic Biomes Conclusions 93 References 93

4 Contents IX 7 Animal Diversity at High Altitudes in the Austrian Central Alps E. Meyer and K. Thaler Introduction " Altitudinal Zonation of the Main Groups of Animals Species Numbers of Invertebrates and Altitudinal Zonation Animal Communities Above the Timberline Altitude-Related Changes in Abundance and Biomass of the Soil Fauna Endemism and Vicariance in the Alpine Fauna 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 What Were the Composition and Regional Patterns of Tundra During Periods of Different Late Quaternary Climate? What Were the Patterns of Shrub and Tree Invasions into Tundra When Climate Warmed at the End : ; of the Last Glacial Period? / What Major Changes in Ecosystem Processes Accompanied the Conversion of Tundra to Forest at the End of the Last Glacial Period? Conclusion 121 References \ 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 Introduction Mammalian Effects on Ecosystem Processes 128

5 X Contents 9.3 Hypotheses for the Steppe-Tundra Transition Conclusions 132 References Palaeorecords of Plant Biodiversity in the Alps B. Ammann Introduction Possibilities and Limitations of the Fossil Record Temporal Changes in Biodiversity in the Alps - Three Examples Needs for Future Research Conclusions 146 References Implications for Changes in Arctic Plant Biodiversity from Environmental Manipulation Experiments T.V. Callaghan and S. Jonasson Introduction Mechanisms of Change in Biodiversity Responses of Soils to Environmental Manipulations: Implications for Plant Nutrition and Biodiversity Direct Responses of Plants to Environmental Perturbations and Implications for Changes in Populations, Communities and Biodiversity \ Conclusions 163 References Patterns and Current Changes in Alpine Plant Diversity G. Grabherr, M. Gottfried, A. Gruber, and H. Pauli Introduction The Altitudinal Limits of Plant Life Patterns V Diversity Effects of Global Warming on Diversity Conclusions References 180

6 Contents XI 13 Anthropogenic Impacts on Biodiversity in the Arctic O.R. Young and F.S. Chapin III..' Proximate Causes Social Drivers Arctic Interactions Humans as Components of Arctic Ecosystems 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 Introduction Resource Uptake Effects on Community Productivity Conclusions. '..' References Direct and Indirect Effects of Plant Species on Biogeochemical Processes in Arctic Ecosystems S.E. Hobbie Introduction Direct Influence over Biogeochemistry Indirect Influence over Biogeochemistry Predictions of Species'Effects with Climate Warming Approaches to Studying the Role of Species in Biogeochemistry Conclusions 222 References Causes and Consequences of Plant Functional Diversity in Arctic Ecosystems F.S. Chapin III, S.E. Hobbie, 'M.S. Bret-Harte, and G. Bonan Introduction Predicting Species Response to Global Change 225

7 XII Contents 16.3 Effects of Species on Ecosystem Processes Species Diversity as Insurance Against Loss of Function Conclusions 235 References Ecosystem Consequences of Microbial Diversity and Community Structure J. Schimel Introduction "Broad" Processes "Narrow" Processes Stress " Implications for Tundra Overall Considerations Conclusions 250 References Diversity of Biomass and Nitrogen Distribution Among Plant Species in Arctic and Alpine Tundra Ecosystems J. Pastor Introduction Concepts of Diversity Patterns of Diversity of Carbon and Nitrogen Distribution Experimental Tests of Controls over Diversity Towards a Dynamic Model of Diversity in Tundra Ecosystems Conclusions \267 References The Plant-Vertebrate Herbivore Interface in Arctic Ecosystems R.L. Jefferies and J.P. Bryant 271, 19.1 Introduction Patterns of Vegetation and Vertebrate Herbivory in Arctic Tundra Vertebrate Herebivory in a Warming Climate: Implications for Tundra Biodiversity Effects of Anthropogenic Disturbances on Trophic Interactions - A Potential Climatic Change Scenario 274

8 Contents. XIII 19.5 Conclusions Summary 278 References 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 Introduction Arctic Insect Diversity Black Fly Life History Features Trophic Dynamics Organic Matter Processing by Black Flies in an Arctic Lake Outlet Summary and Conclusions 292 References Land-Water Interactions: The Influence of Terrestrial Diversity on Aquatic Ecosystems G.W. Kling Introduction Inputs from Land and the Regulation of Aquatic Systems Terrestrial Diversity Movement of Material from Land to Water Conclusions 305 References Part IV: A Synthesis 22 Patterns, Causes, Changes, and Consequences of Biodiversity in Arctic and Alpine Ecosystems F.S. Chapin III and Ch. Korner The Arctic and Alpine Biota Past, Present, and Future Changes in Biodiversity The Significance of Biodiversity /or Ecosystem Function Conclusions 320 Subject Index. 321

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