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_Polar Lakes and Rivers_ This page intentionally left blank _Limnology of Arctic and Antarctic Aquatic Ecosystems_ EDITED BY Warwick F. Vincent and Johanna Laybourn-Parry Great Clarendon Street, Oxford OX2 6DP Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries. Published in the United States by Oxford University Press Inc., New York © Oxford University Press 2008 The moral rights of the authors have been asserted Database right Oxford University Press (maker) First published 2008 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, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above. You must not circulate this book in any other binding or cover and you must impose the same condition on any acquirer. British Library Cataloguing in Publication Data Data available Library of Congress Cataloging in Publication Data Data available Typeset by Newgen Imaging Systems (P) Ltd., Chennai, India Printed in Great Britain on acid-free paper by Antony Rowe, Chippenham, Wiltshire ISBN 978–0–19–921388–7 (Hbk) ISBN 978–0–19–921389–4 (Pbk) 10 9 8 7 6 5 4 3 2 1 Preface From the summit of the tumulus I saw the ice ahead of us in the same condition. a long blue lake or a rushing stream in every furrow. Peary, R.E. (1907). Nearest the Pole, p. 220. Hutchinson, London. We marched down a narrow gap, cut through a great bar of granite, and saw ahead of us a large lake, some three miles long. It was of course frozen, but through the thick ice covering we could see water plants, and below the steep cliffs the water seemed very deep. Taylor, G. (1913). The western journeys. In Huxley, L. (ed.), Scott’s Last Expedition, vol. II, p. 193. Smith Elder & Co., London. From the early explorers onwards, visitors to the Arctic and to Antarctica have commented with great interest on the presence of lakes, wetlands, and flowing waters. These environments encompass a spectacular range of conditions for aquatic life, from dilute surface melt ponds, to deep, highly stratified, hypersaline lakes. Many of these high-latitude ecosystems are now proving to be attractive models to explore fundamental themes in limnology; for example, landscape–lake interactions, the adaptation of plants, animals, and microbes to environmental extremes, and climate effects on ecosystem structure and functioning. Some of these waters also have direct global implications; for example, permafrost thaw lakes as sources of greenhouse gases, subglacial aquatic environments as a planetary storehouse of ancient microbes, and Arctic rivers as major inputs of fresh water and organic carbon to the world ocean. For more years than we care to admit, the two of us have talked about the need for a text on high-latitude lakes and rivers that compared and contrasted the two polar regions. Whereas the Arctic and Antarctic have much in common, they also have distinct differences. Within the polar research community, scientists typically work exclusively in either the Arctic or in Antarctica. Both of us have conducted research in both polar regions, and this has impressed upon us the remarkable diversity of high-latitude aquatic ecosystems, and their striking commonalities and differences. The Arctic and Antarctica are currently the focus of unprecedented public and political attention, not only for their natural resources and geopolitical significance, but also because they are continuing to provide dramatic evidence of how fast our global environment is changing. Moreover, 2007–2008 marked the fourth International Polar Year (IPY), so the time seemed right to turn the talk into action. However, if it had not been for Ian Sherman of Oxford University Press, and his persuasiveness and encouragement at the American Society for Limnology and Oceanography meeting in Santiago de Compostela in 2005, we would probably not have embarked on this book. One of the valuable opportunities provided by this book project has been to bring together groups of Arctic and Antarctic scientists for many of the chapters. People who ordinarily would not have found themselves collaborating have shared their knowledge and expertise from the two polar regions. We hope that this collaboration will foster further joint ventures among our colleagues, and that it will encourage a more pole-to-pole approach towards high-latitude ecosystems, in the spirit of IPY. This book is intended for both the specialist and the more general reader. To assist the latter we have included a glossary of terms. The color plates are also intended to provide a better picture of the habitats and organisms to those unfamiliar with them. We asked the authors to adopt a tutorial approach for nonspecialists, to limit their citations to be illustrative (rather than exhaustive) of key concepts and observations, and, where possible, to consider differences and similarities between the Arctic and Antarctic. We have greatly appreciated their willingness to be involved in this project and the excellence of their contributions. In addition to thanking the contributing authors to this volume, we express our gratitude to Ian Sherman, Helen Eaton, and other staff at Oxford University Press for their expert help in bringing this volume through to completion; the many reviewers of the manuscripts; Janet Drewery at Keele University and Tanya Adrych at the University of Tasmania for assistance in manuscript preparation; our students, postdoctoral fellows, and other researchers who have worked with us on the limnology of Arctic and Antarctic lakes and rivers; and our research funding and logistics agencies, including the Natural Sciences and Engineering Research Council (Canada), the Canada Research Chair program, the Canadian Network of Centers of Excellence program ArcticNet, Polar Shelf Canada, the Natural Environment Research Council (UK), the Engineering and Physical Sciences Research Council (UK), the Leverhulme Trust, and the Australian, UK, New Zealand, Spanish, and US Antarctic programmes. Warwick F. Vincent and Johanna Laybourn-Parry 2008 About International Polar Year The International Polar Year (IPY) 2007–2008 represents one of the most ambitious coordinated international science programmes ever attempted. Researchers from over sixty countries and a broad range of disciplines are involved in this two-year effort to study the Arctic and Antarctic and explore the strong links these regions have with the rest of the globe. Researchers in IPY 2007–8 have made a commitment to raising awareness about the polar regions and increasing the accessibility of science. This book is part of an internationally endorsed IPY outreach project. For more information, please visit www.ipy.org. Contents Preface v About international polar year vii Contributors xvii 1 Introduction to the limnology of high-latitude lake and river ecosystems 1 Warwick F. Vincent, John E. Hobbie, and Johanna Laybourn-Parry Outline 1 1.1 Introduction 1 1.2 History of polar limnology 4 1.3 Limnological diversity 5 1.4 Controlling variables for biological production 7 1.4.1 Water supply 7 1.4.2 Irradiance 8 1.4.3 Low temperature 8 1.4.4 Nutrient supply 9 1.4.5 Benthic communities 9 1.5 Food webs in polar lakes 10 1.6 Polar lakes and global change 12 1.6.1 Physical thresholds 12 1.6.2 Biogeochemical thresholds 13 1.6.3 Biological thresholds 13 1.7 Conclusions 14 Acknowledgements 14 References 14 Appendix 1.1 18 2 Origin and geomorphology of lakes in the polar regions 25 Reinhard Pienitz, Peter T. Doran, and Scott F. Lamoureux Outline 25 2.1 Introduction 25 2.2 Lake origins 26 2.2.1 Wetlands 26 2.2.2 Ice-dependent lakes 27 2.2.3 Postglacial lakes 28 2.2.4 Thermokarst lakes and ponds 30 2.2.5 Coastal uplift systems 31 2.2.6 Meteoritic impact crater lakes 32 2.2.7 Volcanic lakes 32 2.2.8 Karst systems 34 2.2.9 Tectonic lakes 34 2.2.10 Lakes of other origins 34 2.3 Geographical regions 35 2.3.1 The circumpolar Arctic 35 2.3.2 Coastal Antarctic lakes 35 2.3.3 Antarctic and Arctic subglacial lakes 36 2.4 Effects of landscape evolution and climate change on polar lakes 37 2.5 Conclusions 38 Acknowledgments 38 References 38 Ключевые слова: developmental stage, hydrobiologia, alaska, microbial loop, optical property, geophysical, diversity, smol, surface, oceanography, blake, bacterivorous flagellate, press, thermokarst disturbance, biomass, ecology, source, standing stock, victoria island, letters doi, island, bacteria, northern, subantarctic island, ecosystem, alpha-beta-, mat, positive correlation, concentration, time, lake, char, microbiology, van trappen, mcknight, water, global distillation, branchinecta gaini, uv-b radiation, group, northern coastline, internal wave, wa, multiple stressor, bunger hills, arctic lake, environmental stochasticity, disko island, dry valleys, dordrecht, antarctic, environment, gibson, ha, petroleum hydrocarbon, structure, parry, anaerobic condition, fairy shrimp, taymyr peninsula, nutrient, ecosysten dynamics, high, delayed maturation, gammaand epsilonproteobacteria, solar radiation, euphotic zone, antarctic lake, hawes, specie, valleys, antarctica, production, mcmurdo dry, organic matter, serge duchesneau, larsemann hills, hobbie, ellesmere island, van hove, permafrost, primary production, doran, sediment, process, primary, freezethaw cycle, dry, science, laybourn-parry, doc, tectonic framework, climate, form, laybourn parry, epilithic biofilm, seasonal progression, valley, glacial, change, freshwater biology, canada, royal society, thermal regime, environmental property, generalized depiction, mcmurdo, polar desert, springer, period, trophic cascade, los ros, cyanobacterial trichomes, depth, national academy, cover, hills, exciting frontier, transantarctic mountains, international conference, high-latitude, priscu, ice shelf, vestfold hills, community, journal, water-filled hole, biology, rate, trans-antarctic mountains, main body, north, study, region, canadian, cold condensation, temperature, food, greenland, organochlorine pesticide, glacier, davis station, signy island, habitat, data, ecological, superoxide dismutase, hodgson, year, environmental, peterson, annual cycle, sulphur dioxide, laybourn, area, microbial consortium, result, strong correlation, phytoplankton, limnology, river, american society, functional gene, douglas, mcmlter org, byers peninsula, clostridium laramiense, level, dowden hutchinson, carbon, structural weakness, aquatic, polar, limnology oceanography, isostatic rebound, stream, arctic, microbial, physical, kola peninsula, biogeochemical processes, polar biology, flow, siegert, freshwater, profundal zone, rapid alteration, cape royds, calcium carbonate, summer, microbial ecosystems, radiation, basal mechanic, long-term ecological, power, pond, ice, filose amoeba, blood falls, latitude, benthic, hudson bay, leaf litter, high arctic, growth, electron acceptor, condition, rrna gene, organic, shelf, marginal zone, limnological property, nature, junk publishers, finnish lapland, biogeochemical property, war, population, vincent, fish, subglacial, station infrastructure, lindell, primary producer, ultraviolet radiation, climate change, large