Alpine Ecosystems in the Northwest Caucasus

421 Pages·2004·13.139 MB·English

by  V. G. Onipchenko (auth.), Vladimir G. Onipchenko (eds.)

Alpine Ecosystems in the Northwest Caucasus
Geobotany 29
Series Editor
M.l.A. WERGER
The titles published in this series are listed at the end of this volume.
Alpine Ecosystems
in the Northwest Caucasus
Edited by
Vladimir G. Onipchenko
Department of Geobotany,
Moscow State University, Moscow, Russia
Springer-Science+Business Media, B. V.
A c.I.P. Catalogue record for this book is available from the Library of Congress
ISBN 978-90-481-6649-7 ISBN 978-1-4020-2383-5 (eBook)
DOI 10.1007/978-1-4020-2383-5
Printed on acid-free paper
All Rights Reserved
© 2004 Springer Science+Business Media Dordrecht
Originally published Kluwer Academic Publishers in 2004
Softcover reprint of the hardcover I st edition 2004
No part of this work may be reproduced, stored in a retrieval system, or transmitted
in any form or by any means, electronic, mechanical, photocopying, microfilming, recording
or otherwise, without written permission from the Publisher, with the exception
of any material supplied specifically for the purpose ofbeing entered
and executed on a computer system, for exclusive use by the purchaser of the work.
To the memory ofP rofessor Tikhon A.Rabotnov
Contents
Contributors X111
Acknowledgement XVll
Preface xix
1. Geography, geology, climate and the communities studied
(V.G.Onipchenko) I
1.1. Geographical position I
1.2. Relief 1
1.3. Geology 3
1.4. Climate 3
1.5. Studied communities 4
2. Soils and nutrient turnover (T.L Malysheva, V.G. Onipchenko,
Makarov M.I., A.V. Volkov, LV. Bulatnikova) 9
2.1. Main soil types of the Teberda Reserve 9
2.1.1. Forest soils 10
2.1.2. Subalpine and alpine soils 10
2.1.3. Variability of soil properties under different subalpine
and alpine communities 12
2.2. Soils ofthe four studied alpine communities 15
2.3.Nutrient turnover in the alpine communities 19
2.3.1. Element accumulation in communities 19
2.3.2. Coefficients of biological uptake (CBU) 21
2.3.3. Relative concentration of elements (RCE) 22
Vll
Vlll Contents
3. Plant geographical description of the area (V.N. Pavlov) 25
3.1. History of floristic study 25
3.2. Steppe belt 28
3.3. Deciduous (broad-leaved) forest belt 30
3.4. Coniferous forest belt 35
3.5. Timberline 40
3.6. Subalpine belt 43
3.7. Alpine belt 47
3.8. Subnival belt 50
4. Alpine plant communities: syntaxonomy, floristic composition
and richness (V.G.Onipchenko) 55
4.1. Floristic sources 55
4.2. Syntaxonomy of high-mountain communities 56
4.2.1. Screes and pebbles 56
4.2.2. Rocks 57
4.2.3. Fens 58
4.2.4. Cold water springs 58
4.2.5. Kobresia grasslands 59
4.2.6. Alpine heaths 60
4.2.7. Alpine grasslands 62
4.2.8. Snow bed communities 64
4.2.9. Subalpine meadows 65
4.2.10. Elfin dwarf schrub 67
4.2.11. Elfin birch woodland 68
4.3. Comparison of the floristic richness of alpine communities
in the Northwest Caucasus and the Central Alps 69
4.3.1. Introduction and object description 69
4.3.2. Compared communities 70
4.3.3. Methods and results 70
4.3.4. Discussion 74
5. Structure and dynamics of closed alpine communities 77
5.1. Plant life forms (O.V. Cherednichenko) 77
5.1.1. Morphological analysis of alpine communities 77
5.1.2. Experimental study of the potential ability of
taproot-rhizomatous plants to vegetative propagation 86
5.2. Biomass and production (V.G. Onipchenko) 92
5.2.1. Introduction 92
5.2.2. Methods 92
5.2.3. Aboveground biomass 93
5.2.4. Belowground biomass 96
Contents IX
5.2.5. Production estimation 98
5.3. Spatial structure (V.G. Onipchenko, O.V. Cherednichenko,
A.A. Zakharov) 99
5.3.1. Introduction 99
5.3.2. Spatial heterogeneity of plant communities and soil
properties according "mass-effect" approach 99
5.3.3. Alpine lichen heath spatial structure: hypothesis
and experiments 105
5.3.4. Festuca varia grasslands 111
5.3.5. Geranium gymnocaulon-Hedysarum caucasicum meadow 117
5.3 .6. Spatial structure of snowbed communities 121
5.4. Soil-plants relationships (V.G. Onipchenko, M.1. Makarov) 125
5.4.1. Introduction and methods 125
5.4.2. Effects by species 127
5.4.3. Variation within and between taxa and samples 131
5.4.4. Comparison with natural community soil properties 133
5.4.5. Cluster analysis of the species 135
5.5. Seasonal dynamics (V.G. Onipchenko) 136
5.5.1. Introduction 136
5.5.2. Methods 136
5.5.3. Phenological curves and periods 137
5.5.4. Plant phenological types 141
5.6. Fluctuations and trends in the dynamics of alpine lichen heath
and snobed communities (V.G. Onipchenko, K.A. Golikov,
O.V. Blinkova, A.A. Zakharov) 144
5.6.1. Introduction 144
5.6.2. Methods 145
5.6.3. Variation in shoot density 146
5.6.4. Species groups according to their dynamic behaviour
and weather conditions 149
5.6.5. Trends in species dynamics 151
5.6.6. Autocorrelation analysis 153
5.6.7. ALH revegetation after wild boar digging 154
5.7. Successions and historical dynamics during the Holocene
(M.S. Blinnikov) 158
5.7.1. Introduction 158
5.7.2. Methods 158
5.7.3. Pollen analysis results and discussion 160
5.7.4. Phytolith analysis: results and discussion 172
5.7.5.Conc1usions 178
x Contents
6. Population biology of alpine plants 181
6.1. Age structure and adaptation of the populations of three
non-clonal alpine perennial plants (I.E. Sizov, N.V.Mikhailova,
A.V. Mikhailov, V.G. Onipchenko, A.S. Komarov) 181
6.1.1. Introduction 181
6.1.2. Materials and methods 183
6.1.3. Age-state composition 184
6.1.4. Life spans of age stages 186
6.1.5. Frequency of flowering 189
6.1.6. Analysis of the populations' dynamics using matrix
models 192
6.1. 7. Conclusions 194
6.2. Soil seed banks (G.V. Semenova) 194
6.2.1. Introduction 194
6.2.2. Methods 195
6.2.3. Results and discussion 197
6.3. Population strategies (V.G.Onipchenko, G.V. Semenova) 212
6.3.1. Introduction 212
6.3.2. Methods 215
6.2.3. Results and discussion 216
7. Experimental study of plant relationships 223
7.1. Reciprocal transplantations in alpine plant communities
(E.V. Volkova, V.G. Onipchenko, M.S. Blinnikov) 223
7.1.1. Introduction 223
7.1.2. Methods 224
7.1.3. Results and discussion 225
7.2. Dominant removals 236
7.2.1. Alpine lichen heath (A.A. Aksenova,
V.G. Onipchenko, M.S. Blinnikov) 236
7.2.2. Removals in the Geranium gymnocaulon -Hedysarum
caucasicum meadow (O.V.Cherednichenko) 244
7.3. Shading experiments (A.A. Aksenova,
V.G. Onipchenko, M.S.Blinnikov) 250
7.3.1. Introduction 250
7.3.2. Field experiments 251
7.3.3. Influence of light deficiency on the floristic
richness of the four alpine communities 252
7.3.4. More and less shade-tolerant species in the four
communities 253
7.3.5. Difference in species reaction to early versus
late-season shading 257
Contents Xl
7.3.6. Discussion 261
8. Soil algae and cyanobacteria in the alpine plant communities
(L.Rasran) 263
8.1. Introduction and methods 263
8.2. Floristic composition 264
8.3. Biomasss 269
9. Fungi, mycorrhiza, and microbiological processes in the
alpine ecosystems 271
9.1. Soil micro fungal communities (M.V. Sogonov,
L.L. Velikanov) 271
9.1.1. Introduction 271
9.1.2. Study sites 272
9.1.3. Methods 273
9.1.4. Results and discussion 273
9.1.5. Conclusion and perspectives 282
9.2. Phytopathogenic fungi (K.L. Tarasov, V.G. Onipchenko) 283
9.3. Mycorrhiza (V.G. Onipchenko) 284
9.3.1. Introduction 284
9.3.2. Methods of sampling and microscoping 284
9.3.3. Types ofmycorrhiza 285
9.3.4. Comparison between communities 286
9.3.5. Seasonal dynamics ofmycorrhiza 289
9.3.6. Ontogenetic dynamics 290
9.3.7. Mycorrhizal infection and vegetative mobility 291
9.3.8. Disturbance and mycorrhizal infection 295
9.4. Soil fungal biomass and microbial activity
(T.N. Leinsoo, M.V.Sogonov, A.L. Stepanov) 296
9.4.1 Introduction 296
9.4.2 Methods 297
9.4.3 Results 298
10. Animals and their influence on plant communities 303
10.1. Wild ungulates (M.N. Ivanenko, V.G. Onipchenko) 303
10.1.1. The Caucasian goat (ibex) (Capra caucasica Guldenst.) 303
10.1.2. Chamois (Rupicapra rupicapra L.) 309
10.1.3. Wild boar (Sus scrofa L.) 311
10.2. Burrowing and trophic activity of small mammals
(S.V.Fomin, V.G. Onipchenko, M.A. Gerasimova) 312
10.2.1. Species diversity 312
10.2.2. Pine vole population density and spatial distribution 315