ECOSYSTEMS OF THE WORLD Editor in Chief: David W. Goodall CSIRO, Midland, W.A. (Australia) I. TERRESTRIAL ECOSYSTEMS II. AQUATIC ECOSYSTEMS A. Natural Terrestrial Ecosystems A. Inland Aquatic Ecosystems 1. Wet Coastal Ecosystems 22. River and Stream Ecosystems 2. Dry Coastal Ecosystems 23. Lakes and Reservoirs 3. Polar and Alpine Tundra 4. Mires: Swamp, Bog, Fen and Moor B. Marine Ecosystems 5. Temperate Deserts and Semi-Deserts 24. Intertidal and Littoral Ecosystems 6. Coniferous Forests 25. Coral Reefs 7. Temperate Deciduous Forests 26. Estuaries and Enclosed Seas 8. Natural Grasslands 27. Ecosystems of the Continental Shelves 9. Heathlands and Related Shrublands 28. Ecosystems of the Deep Ocean 10. Temperate Broad-Leaved Evergreen Forests 11. Mediterranean-Type Shrublands C. Managed Aquatic Ecosystems 12. Hot Deserts and Arid Shrublands 29. Managed Aquatic Ecosystems 13. Tropical Savannas 14. Tropical Rain Forest Ecosystems III. UNDERGROUND ECOSYSTEMS 15. Wetland Forests 16. Ecosystems of Disturbed Ground 30. Cave Ecosystems B. Managed Terrestrial Ecosystems 17. Managed Grasslands 18. Field Crop Ecosystems 19. Tree Crop Ecosystems 20. Greenhouse Ecosystems 21. Bioindustrial Ecosystems ECOSYSTEMS OF THE WORLD 3 POLAR AND ALPINE TUNDRA Edited by F.E. Wielgolaski Department of Biology, Division of Botany, University of Oslo, P.O. Box 1045, Blindern N-0316 Oslo, Norway ELSEVIER Amsterdam - Lausanne - New York - Oxford - Shannon - Singapore - Tokyo ELSEVIER SCIENCE B.V. Sara Burgerhartstraat 25 P.O. Box 211 1000 AE Amsterdam The Netherlands Library of Congress Cataloging in Publication data Polar and alpine tundra / edited by F.E. Wielgolaski. p. cm. -- (Ecosystems of the world ; 3) Includes bibliographical references and index. ISBN 0-444-88265-0 1. Tundra ecology. I. Wielgolaski, F.E., 1931- . II. Series. QH541.5.T8P65 1997 577.5'86--dc21 97-19344 CIP First edition: 1997 Second impression: 1998 ISBN 0-444-88265-0 (volume) ISBN 0-444-41702-8 (series) © 1997 Elsevier Science B.V. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the publisher, Elsevier Science B.V., Copyright and Permissions Department, P.O. Box 521, 1000 AM Amsterdam, The Netherlands. Special regulations for readers in the U.S.A. - This publication has been registered with the Copyright Clearance Center Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923. Information can be obtained from the CCC about conditions under which photocopies of parts of this publication may be made in the U.S.A. All other copyright questions, including photocopying outside of the U.S.A., should be referred to the publisher. © The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Printed in The Netherlands. PREFACE Polar and alpine tundra have many similarities, areas more knowledge has been available on the but also differences; particularly towards the equa- ecology of plants than on other organisms. tor, high mountains show strong differences from polar regions. Therefore, workers in tropical The Arctic tundra of the world is divided between mountains normally do not treat the highest ele- only few countries. Two of the chapters - those vated areas studied as tundra. Towards and in the covering North America and Russia - are there- subpolar regions there are generally more similari- fore particularly long. ties between alpine belts and real polar tundra, although permafrost may often be missing in the Work on this volume started many years ago. mountains even there. However, these similarities Some of the authors were able to complete their justify the presentation of polar and alpine tundra assignments relatively fast; others took considera- in one volume of this series. bly longer. Apologies were therefore offered to the former group, who saw their writing go out of The chapters of the volume are mainly arranged date, which, in some cases, necessitated considera- geographically, starting with some general intro- ble re-writing, when the volume as a whole was ductory chapters, followed by the alpine tundra of ready for press. As editor, I want to thank the various regions, and ending with chapters on the various authors for their willingness to contribute, polar regions. their patience, cooperation and acceptance of sug- gestions by the editor. There have been problems to find authors for some alpine areas, particularly smaller mountains. Gen- I also want to thank Prof. D.W. Goodall, Editor- erally, this reflects the fact that only little ecologi- in-Chief of the series, for all his help both in cal work has been carried out in the area. It is also professional matters and in linguistic corrections. obvious from the chapters written, that in some F.E. WIELGOLASKl Professor of Botanical Ecology University of Oslo, Norway Chapter 16 ARCTIC ECOSYSTEMS IN RUSSIA Yu.I. CHERNOV and N.V. MATVEYEVA INTRODUCTION dra which formed the basis of the collection of the Botanical Institute of the Academy of Science for Biological research in the Eurasian Arctic the northeast of Europe. In his book, Kertselli started at the beginning of the last century. Right (1911) gives much interesting ecological informa- from the dawn of investigations on nature in tion. northern Europe and Siberia, there were attempts In the first decade of this century, the literature at some ecological generalizations to embrace and devoted to the plant and animal kingdoms of analyze the natural phenomena of life in the Eurasian tundras had become rather extensive. tundra. Schrenk (1848, 1854), for instance, gave Common features in the composition and distribu- many interesting and deep thoughts on the life of tion of many taxa, particularly birds, mammals, tundra plants and animals, and interesting ecologi- insects, and vascular plants, were outlined (see cal studies were made by Baer (1838), who visited reviews by Gorodkov, 1938; Aleksandrova, 1956; Novaya Zemlya in 1837. The famous work of Rebristaya, 1977; Chernov, 1978b). For instance, Middendorf (1869), which played a very large role in 1904 a detailed summary by Schalow on birds in investigations on nature in arctic Siberia, should of the Arctic was published. Other issues of the be considered as the starting point of scientific Fauna Arctica published in Jena sum up the results ecology in the Arctic. from studies on various groups of animals at high At the end of the 19th and the beginning of the latitudes. There, also, were published the first data 20th centuries, interest in studying the organic on the microflora of the Arctic (Severin, 1909; world of the extreme North was dramatically Isachenko, 1914) and results are given from inten- increased. At that time, numerous expeditions sive studies of cryptogamic plants of the tundra were undertaken with special purposes: zoological, zone (Elenkin, 1909; Brotherus, 1910). botanical, and entomological. Rich collections Of great significance for the development of were built up which served as a basis of knowledge biological investigations in the Arctic were the of the arctic flora and fauna; for example, in numerous expeditions undertaken in the 1920s to Russia, the collections of Jacobson in 1896 on Novaya Zemlya, which soon became in some way Novaya Zemlya, and entomological and botanical a standard of arctic nature. In that period, one materials from the expedition of the Kuznetsov Norwegian and several Soviet expeditions worked brothers to the polar Ural and the Yugorskiy there, and great numbers of botanical and zoologi- peninsula (Yugorskiy Poluostrov) in 1909. Vast cal papers were published. An especially important materials on many groups of plants and animals role was played by a series of papers by Tolmachev were obtained by the Russian Polar Expedition (1930, 1931) in which he gave an analysis of the headed by Toll in 1900-1903 on Taymyr and in composition of flora on Novaya Zemlya and, some other regions of arctic Siberia. One of the using this region as an example, raised the problem participants of this expedition, Birulya (1907), of floristic origins in the Arctic. studied many aspects of the biology of arctic birds Yet, even now the qualitative composition of the and biocenotic relations. A veterinarian and trav- arctic flora and fauna is far from being completely eller, Kertselli, collected a thoroughly documented established. The vascular plant flora has been most herbarium of the flora of Bol'shezemel'skaya Tun- thoroughly studied, as is reflected in the 10-volume 361 362 Yu.I. CHERNOV AND N.V. MATVEYEVA edition of the Arctic Flora of the USSR. Important In particular, analyses of the tundra yeast flora contributions to the study of the flora of arctic have proved the necessity of revising many tradi- Eurasia were made by the founder of that series, tional ideas on the taxonomy of unicellular fungi Tolmachev, and some of his successors from the (I. Chernov and Bab'eva, 1988). school created by him (Yurtsev, 1966, 1974; Pet- Information on the soil microflora of the Rus- rovskyi, 1973, 1985; Rebristaya, 1977; Yurtsev et sian arctic landscapes is also at an initial stage, al., 1978; Tolmachev, 1986). At present, the com- with earlier work having been limited to some position of the vascular plant flora is one of the episodic works by Isachenko and Simakova most advanced lines of biological investigation at (1934), Kriss (1947, 1952), and Sushkina (1960). At high latitudes. the end of the 1960s, Parinkina started her investi- In general, the taxonomy and the zonal and gations on Taymyr. Owing to her systematic work, regional distribution of mosses and lichens, the studies on the composition of the microflora most important cenosis-formers in the Arctic, have and on the bacterial ecology of tundra soils have also been clarified. A basic handbook of arctic made noticeable progress (Parinkina, 1971, 1973, mosses was published in 1961 (Abramova et al., 1979a-c, 1986).