Alpine Biodiversity in Europe
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1 L. Nagy G. Grabherr Ch. Korner D.B.A. Thompson (Eds.) Alpine Biodiversity in Europe With 89 Figures, 17 in Color, and 61 Tables Springer
2 I Europe s Alpine Areas 1 An Outline of Europe s Alpine Areas... 3 G. GRABHERR. L. NAGY and D.B.A. THOMPSON 1.1 The Mountains of Europe An Ecological Classification of Europe s Alpine Areas The Regional Accounts (Chaps ) A Bioclimatic Characterisation of Europe s Alpine Areas.. 13 CH. KÖRNER. J. PAULSEN and S. PELAEZ-RIEDL 2.1 Introduction Site Selection and Methods What Was Measured and Where? Sensors and Sensor Positioning Data Analysis Results Between-Year and Between-Site Variation Temperature Extremes The Daily Course of Ground Temperature The Seasonal Course of Ground Temperature Seasonal Mean Temperatures Growing Season Length Thermal Sums Alpine Biota Share a Common Climate Across Europe
3 3 The Regional Accounts 3.1 The High Mountain Vegetation of the Scandes R. VIRTANEN Geography. Geology. and Ecological Conditions Climate TheFlora The Alpine Zone and the Treeline Ecotone Number of Plant Taxa Rare Taxa Altitude Zones Raunkiaer s Biological Types Vegetation and Its Responses to a Changing Environment The High Mountain Vegetation of Scotland L. NAGY Introduction Geology, Soils and Climate TheFlora The Present-Day Flora Vegetation Species and Habitat Diversity and Causes of Change Vegetation of the Giant Mountains. Central Europe J. JENIK and J. STURSA Introduction Physical-Geographical Setting Flora and Vegetation Refugia of the Polygenous Flora The Alpine Vegetation of the Alps P. OZENDA and J.-L. BOREL Physiography Floristic Diversity Species Richness. Altitude Endemism... 58
4 3.4.5 Taxonomic Diversity Life Forms Comparison of Different Mountain Ranges - Species Richness. Floristic and Community Similarities The Alpine Flora and Vegetation of the South-Eastern Carpathians G. COLDEA Geography and Geology The Treeline Ecotone (Subalpine) and the Alpine Zone Climate Flora History of the Flora Number of Taxa Vegetation Human Impacts vs. Diversity The High Mountain Flora and Vegetation of the Apennines and the Italian Alps... F. PEDROTTI and D. GAFTA Geography Geology and Climate History of the Flora Today s Flora Altitudinal Vegetation Zonation Floristic Diversity and Distinctiveness of Communities Causes of Change and Conservation The Vegetation of the Alpine Zone in the Pyrenees D. GOMEZ. J.A. SESE and L. VILLAR Geography History, Geology. Soils and Climate History of the Flora Today s Flora Vegetation
5 3.8 High Mountain Vegetation of the Caucasus Region... G. NAKHUTSRISHVILI The Caucasus Geography and Geology Climate TheFlora History of the Flora Human Impacts Number of Taxa High Mountain Vegetation The Treeline Ecotone Tall-Herb Vegetation Meadows, Feather-Grass Steppes and Scrub at the Treeline The Alpine Zone Proper The Subnival Zone A Comparative Analysis of the High Mountain Vegetation of the Caucasus and the Alps The Vegetation of the Corsican High Mountains... J. GAMISANS Geography Climate TheFlora History of the Flora Human Impact The Number of Taxa Vegetation Altitude Zones The High Mountain Vegetation of the Balkan Peninsula A. STRID. A. ANDONOSKI and V. ANDONOVSKI Geography Climate Soils The Flora Endemism and Speciation Vegetation
6 Plant and Vegetation Diversity 4 Overview: Patterns in Diversity U. MOLAU 4.1 Scale and Diversity Temporal Diversity Diversity at Landscape and Continent Scales Taxonomic Diversity of Vascular Plants in the European Alpine Areas H. VARE. R. LAMPINEN. C. HUMPHRIES and P. WILLIAMS 5.1 Introduction Methods The Selection of Taxa Taxon Richness Range-Size Rarity Area Selection Near-Minimum Area Set to Represent All Hotspot Areas Results and Discussion Taxon Richness Regional Mountain Range Richness Area Selections Conclusions Patterns in the Plant Species Richness of European High Mountain Vegetation R. VIRTANEN. T. DIRNBÖCK. S. DULLINGER. G. GRABHERR. H. PAULI. M. STAUDINGER and L. VILLAR 6.1 Introduction The Data Geographical and Ecological Trends in Plant Species Richness Discussion Latitudinal Trends The Size of Mountain Areas
7 XVI Soil Substratum Snow Cover Local and Regional Species Richness Conclusions Altitude Ranges and Spatial Patterns of Alpine Plants in Northern Europe J. I. HOLTEN 7.1 Introduction Study Area Methods Altitude Ranges and Patterns Altitudinal Ranges of Alpine Plant Species Boreal-Alpine (Montane) Group Widespread (Ubiquitous) Group Centric Group Alpine Vascular Plant Species Richness vs. Altitude What Limits the Altitudinal Ranges and Patterns in the Alpine Zone? Limiting Factors High Summer Temperatures and Drought Winter Temperatures Alpine Permafrost Conclusions Vascular Plant and Bryophyte Diversity Along Elevational Gradients in the Alps J.-P. THEURILLAT. A. SCHLÜSSEL. P. GEISSLER. A. GUISAN. C. VELLUTI and L. WIGET Introduction Study Areas Methodology The Transects Analyses Patterns of Distribution Elevation and Species Richness Life Forms. Growth Forms and Elevation
8 Distribution of Elevational Species Groups Similarity and Elevation Cumulative Species Richness and Elevation Discussion Theoretical Background The Temperature-Physiography Hypothesis Conclusions Assessing the Long-Term Dynamics of Endemic Plants at Summit Habitats... H. PAULI. M. GOTTFRIED. T. DIRNBÖCK. S. DULLINGER and G. GRABHERR 9.1 Introduction Study Areas Methods Results Vegetation Cover and Vascular Plant Species Cover Species Richness and Endemic Species Discussion Orographic Isolation and the Cryophilic Nature of Endemics Climate Warming and the Vulnerability of Cryophilic Endemic Species Conclusions Mapping Alpine Vegetation... T. DIRNBÖCK. S. DULLINGER. and G. GRABHERR M. GOTTFRIED. C. GINZLER 10.1 Introduction Study Site Methods Results and Discussion The Map Accuracy Assessment Remotely Sensed Maps and Biodiversity Research Conclusions
9 11 A GIS Assessment of Alpine Biodiversity at a Range of Scales... J. KLING, K. LINDBLAD. R. BJÖRK. J. DANHARDT U. MOLAU. and A. LIESS 11.1 Introduction GIS in Ecology The Latnjajaure Catchment Biodiversity Assessment by GIS Field Sampling GIS Database and Modelling Results and Conclusions... Plants... Animals Invertebrates 12 Overview: Invertebrate Diversity in Europe s Alpine Regions... P. BRANDMAYR. R. PIZZOLOTTO and S. SCALERCIO 12.1 Macrolepidoptera Carabids A Comparison of the Diversity Patterns of Arachnids. Carabids and Lepidoptera Conclusions The Geographical Distribution of High Mountain Macrolepidoptera in Europe... Z. VARGA 13.1 Introduction Faunal Types in the European Orobiomes The Alpine Faunal Type Speciation and Endemism in the Alpine Faunal Type Biogeographical Characterisation of the Xeromontane Faunal Type Biogeographical Connections and Similarities of the European High Mountains
10 13.7 Geographical and Phylogenetic Patterns in the European High Mountain Macrolepidoptera Conclusions High Altitude Invertebrate Diversity in the Ural Mountains 259 Y.E. MIKHAILOV and V.N. OLSCHWANG 14.1 Introduction Species Diversity: State of Knowledge Historical Origins of Zoological Research A Brief Inventory of the Fauna The Composition and Origin of the High Altitude Fauna Dynamics of Species Diversity: Long-Term Observations Altitudinal Gradients in Species Diversity and Abundance of Soil Fauna Herbivore-Vegetation Interactions in the Alpine Zone Phytophagous Insects as Potential Bioindicators in Alpine Ecosystems Conclusions The Diversity of High Altitude Arachnids (Araneae Opiliones Pseudoscorpiones) in the Alps K. THALER 15.1 Introduction High Altitude Arachnids of the Alps: a Systematic Overview Opiliones and Pseudoscorpiones Araneae Regional Distribution Biology Vertical Distribution Spatial Distribution Phenology, Winter Activity, Life Cycles Biotic Interactions Various Aspects Conclusions
11 16 Patterns of Butterfly Diversity Above the Timberline in the Italian Alps and Apennines... L. TONTINI. S. CASTELLANO. S. BONELLI and E. BALLETTO 16.1 Introduction Materials and Methods Assemblage Description Grazing Impacts Results Richness and Density of Species Species Richness Along Latitude and Altitude Population Densities and Assemblage Structure Variation Exclusively Alpine Species Richness and Densities Grazing Impacts Conclusions Diversity Patterns of Carabids in the Alps and the Apennines... P. BRANDMAYR. R. PIZZOLOTTO. S. SCALERCIO. M.C. ALGIERI and T. ZETTO 17.1 Introduction Overall Features of Alpine Carabid Communities in the Alps 17.3 Species Recruitment in Alpine Habitats Diversity Patterns in the Alps Carabid Assemblages of the Central Apennines Alpine Carabid Assemblages and Climate Change Conclusions Vertebrates 18 Overview: Alpine Vertebrates. Some Challenges for Research and Conservation Management D.B.A. THOMPSON Species Diversity Variation Population Cycling and Herbivores Habitat Use
12 18.4 Conclusions Breeding Bird Assemblages and Habitat Use of Alpine Areas in Scotland D.B.A. THOMPSON. D.P. WHITFIELD. H. GALBRAITH. K. DUNCAN. R.D. SMITH. S. MURRAY and S. HOLT Introduction Study Area and Methods Results The Breeding Bird Assemblages Habitat Use Discussion Breeding Bird Assemblages Habitat Use Consequences of Habitat Modification Conclusions Rodents in the European Alps: Population Ecology and Potential Impacts on Ecosystems... D. ALLAINE and N.G. Yoccoz 20.1 Introduction Is There an Alpine Rodent? Population Dynamics and Demography Alpine Rodents and Plant Communities Consequences of Changes in the Management of Alpine Ecosystems and in Climate Needs for Further Work Conclusions Large Herbivores in European Alpine Ecosystems: Current Status and Challenges for the Future A. LOISON. C. TOIGO and J.-M. GAILLARD Introduction The Species Concerned and Their Distribution in Europe
13 21.3 Ecological Features Morphology and Physiology Habitat Feeding Ecology Social and Mating System Spatial Features Dynamics in Space and Time Demographic Patterns Density Dependence and Effects of Climate The Role of Large Herbivores on the Mountain Landscape and Plant Communities Changes in Mountain Ecosystems and New Challenges for Large Herbivores... Interaction with Domestic Ungulates, Diseases and Parasites The Increase of Lowland Ungulates and Inter-specific Competition The Return of Large Carnivores... Habitat Fragmentation Recreational Use of the Mountains and Disturbance Conclusions and Future Research Needs Diversity of Alpine Vertebrates in the Pyrenees and Sierra Nevada. Spain... J.P. MARTINEZ RICA 22.1 Introduction Alpine Vertebrate Species Species Richness vs. Diversity Indices Altitudinal Gradients and Biodiversity Conclusions The Impacts of Vertebrate Grazers on Vegetation in European High Mountains... B. ERSCHBAMER. R. VIRTANEN and L. NAGY 23.1 Introduction The Historical Perspective The Grazers and Their Diet Grazing Impacts
14 23.5 The Effects of Grazing Exclusion on the Vegetation Biomass and Species Composition Dead Standing Plant Material, Litter. and Soil Microbes Evidence from Outside Exclosures Canopy Structure. Growth Form. Flowering and Seed Production Succession After the Abandonment of Pasturing Nutrient Enrichment and Community Changes Grazing in the Alpine Zone: Ecological Problems and Prospect Conclusions V Long-Term Vegetation Dynamics 24 Overview: Alpine Vegetation Dynamics and Climate Change. a Synthesis of Long-Term Studies and Observations... G. GRABHERR 24.1 Introduction Treeline Ecotone Trees and Forest Line Dynamics Vegetation Dynamics Within the Alpine Zone 24.4 Dynamic Processes at the Limits of Plant Life Conclusions Long-Term Changes in Alpine Plant Communities in Norway and Finland... R. VIRTANEN. A. ESKELINEN and E. GAARE 25.1 Introduction Study Areas Data Collection and Analysis Changes in Species Composition and Abundance Heath Vegetation Snowbed Vegetation Alpine Soligenous Mires Discussion Conclusions
15 26 Vegetation Dynamics at the Treeline Ecotone in the Ural Highlands. Russia P.A. MOISEEV and S.G. SHIYATOV 26.1 Introduction Materials and Methods Study Areas Long-Term Weather Data Repeat Photography and Image Analysis Four Examples of Photo Pairs Discussion Conclusions Recent Increases in Summit Flora Caused by Warming in the Alps M. BAHN and CH. KÖRNER 27.1 Introduction The Mount Glungezer Summit Floristic Changes Between Conclusions The Piz Linard (341 1 m). the Grisons. Switzerland. Europe s Oldest Mountainvegetation Study Site H. PAULI. M. GOTTFRIED and G. GRABHERR 28.1 Introduction Method Results Changes in Species Richness Changes in Species Abundance Discussion Conclusions
16 29 Alpine Biodiversity in Space and Time: a Synthesis L. NAGY. G. GRABHERR. CH. KÖRNER and D.B.A. THOMPSON 29.1 Spatial Trends in Animal and Plant Species Richness Latitudinal Trends Altitude Trends Animals Temporal Changes Environment Grazers Outlook Subject Index
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