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A New Ecology: Systems Perspective Front cover photo is by B.D. Fath and shows M?ns Klint, Denmark. Back cover photos show (from left to right) (1) a Danish beech forest (Ryge Skov), (2) Krimml Falls in Austria, (3) part of the shore of Namchu Lake in Tibet, (4) Crater Lake, Oregon, USA, and (5) Natron Lake, Tanzania. These were taken by S.E. J?rgensen (1 and 4), B.D. Fath (2), and M.V. J?rgensen (3 and 5). A New Ecology: Systems Perspective Sven E. J?rgensen Environmental Chemistry Section Royal Danish School of Pharmacy DK-2100 Copenhagen, Denmark Brian D. Fath Biology Department Towson University Towson, MD 21252, USA Simone Bastianoni Department of Chemical and Biosystems Sciences University of Siena 53100 Siena, Italy Jo?o C. Marques Department of Zoology Institute of Marine Research (IMAR), University of Coimbra 3004-517 Coimbra, Portugal Felix M?ller Ecology Centre University of Kiel 24118 Kiel, Germany S?ren N. Nielsen Environmental Chemistry Section Royal Danish School of Pharmacy DK-2100 Copenhagen, Denmark Bernard C. Patten Institute of Ecology University of Georgia Athens, GA 30602-2602, USA Enzo Tiezzi Department of Chemical and Biosystems Sciences University of Siena 53100 Siena, Italy Robert E. Ulanowicz Chesapeake Biological Laboratory P.O. Box 38, 1 Williams Street, Solomons, MD 20688-0038, USA Amsterdam • Boston • Heidelberg • London • New York • Oxford Paris • San Diego • San Francisco • Singapore • Sydney • Tokyo Elsevier Linacre House, Jordan Hill, Oxford OX2 8DP, UK Radarweg 29, PO Box 211, 1000 AE Amsterdam, The Netherlands First edition 2007 Copyright © 2007 Elsevier 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. Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone (+44) (0) 1865 843830; fax (+44) (0) 1865 853333; email: permissions@elsevier.com. Alternatively you can submit your request online by visiting the Elsevier web site at http://elsevier.com/locate/permissions, and selecting Obtaining permission to use Elsevier material. Notice No responsibility is assumed by the publisher for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions or ideas contained in the material herein. Because of rapid advances in the medical sciences, in particular, independent verification of diagnoses and drug dosages should be made. Library of Congress Cataloging in Publication Data A catalog record is available from the Library of Congress British Library Cataloguing in Publication Data A catalogue record is available from the British Library ISBN: 978-0-444-53160-5 For information on all Elsevier publications visit our website at books.elsevier.com Printed and bound in The Netherlands 07 08 09 10 11 10 9 8 7 6 5 4 3 2 1 Contents Preface ix 1 Introduction: A New Ecology is Needed 1 1.1 Environmental management has changed 1 1.2 Ecology is changing 2 1.3 Book outline 3 2 Ecosystems have Openness (Thermodynamic) 7 2.1 Why must ecosystems be open? 7 2.2 An isolated system would die (maximum entropy) 8 2.3 Physical openness 13 2.4 The second law of thermodynamics interpreted for open systems 18 2.5 Dissipative structure 20 2.6 Quantification of openness and allometric principles 22 2.7 The cell 30 2.8 What about the environment? 31 2.9 Conclusion 32 3 Ecosystems have Ontic Openness 35 3.1 Introduction 35 3.2 Why is ontic openness so obscure? 36 3.3 Ontic openness and the physical world 39 3.4 Ontic openness and relative stability 49 3.5 The macroscopic openness: Connections to thermodynamics 50 3.6 Ontic openness and emergence 53 3.7 Ontic openness and hierarchies 55 3.8 Consequences of ontic openness: a tentative conclusion 56 4 Ecosystems have Directionality 59 4.1 Since the beginnings of ecology 59 4.2 The challenge from thermodynamics 60 4.3 Deconstructing directionality? 62 4.4 Agencies imparting directionality 63 4.5 Origins of evolutionary drive 66 4.6 Quantifying directionality in ecosystems 68 4.7 Demystifying Darwin 74 4.8 Directionality in evolution? 76 4.9 Summary 77 5 Ecosystems have Connectivity 79 5.1 Introduction 79 5.2 Ecosystems as networks 80 5.3 Food webs 82 5.4 Systems analysis 84 5.5 Ecosystem connectivity and ecological network analysis 86 5.6 Network environ analysis primer 86 5.7 Summary of the major insights cardinal hypotheses (CH) from network environ analysis 92 5.8 Conclusions 101 6 Ecosystems have Complex Dynamics (Growth and Development) 103 6.1 Variability in life conditions 103 6.2 Ecosystem development 105 6.3 Orientors and succession theories 112 6.4 The maximum power principle 115 6.5 Exergy, ascendency, gradients, and ecosystem development 120 6.6 Support for the presented hypotheses 125 6.7 Toward a consistent ecosystem theory 133 6.8 Exergy balances for the utilization of solar radiation 139 6.9 Summary and conclusions 141 7 Ecosystems have Complex Dynamics – Disturbance and Decay 143 7.1 The normality of disturbance 143 7.2 The risk of orientor optimization 151 7.3 The characteristics of disturbance 152 References 265 Ostwald W. 1931. Gedanken zur Biosph?re. BSB B. G. Teubner Verlagsgesellschaft, Leipzig, Germany, p. 1978. Pahl-Wostl C. 1998. Ecosystem organization across a continuum of scales: A comparative analysis of lakes and rivers. In: Peterson DL, Parker VT (eds.), Ecological Scale: Theory and Applications. Columbia University Press, New York, NY, pp. 141–170. Paine RT. 1980. Food webs: linkage, interaction strength and community infrastructure. J Anim Ecol. 49, 667–685. Patricio J, Sala F, Pardal MA, J?rgensen MA, Marques JC. 2006. Ecological Indicators performance during a re-colonization field experiment and its compliance with ecosystem theories. Ecol. Model. 19, 46–59. Patricio J, Ulanowicz R, Pardal MA, Marques JC. 2004. Ascendency as an ecological indicator: a case study of estuarine pulse eutrophication. Est. Coast. Shelf Sci. 60, 23–35. Patten BC. 1978. Systems approach to the concept of environment. Ohio J. Sci. 78, 206–222. Patten BC. 1981. Environs: the super-niches of ecosystems. Am. Zool. 21, 845–852. Patten BC. 1982. Environs: relativistic elementary particles or ecology. Am. Nat. 119, 179–219. Patten BC. 1983. Linearity enigma in ecology. Ecol. Model. 18, 155–170. Patten BC. 1985. Energy cycling in the ecosystem. Ecol. Model. 28, 1–71. Patten BC. 1991. Network ecology: Indirect determination of the life-environment relationship in ecosystems. In: Higashi M, Burns TP (eds.), Theoretical Ecosystem Ecology: The Network Perspective. Cambridge University Press, London pp. 288–351. Patten BC. 1992. Energy, emergy and environs. Ecol. Mod. 62, 29–69. Patten BC, Auble GT. 1981. System theory of the ecological niche. Am. Nat. 117, 893–922. Patten BC, Bosserman RW, Finn JT, Cale WG. 1976. Propagation of cause in ecosystems. Read pp. 458–471. In: Patten, BC (ed.), Systems Analysis and Simulation in Ecology, vol. 4. Academic Press, NY, pp. 457–579. Patten BC, Higashi M, Burns TP. 1990. Trophic dynamics in ecosystem networks: significance of cycles and storages. Ecol. Model. 51, 1–28. Patten BC. Holoecology. The Unification of Nature by Network Indirect Effects. Complexity in Ecological Systems Series, Columbia University Press, New York, NY (in preparation). Patten BC, J?rgensen SE. 1995. Complex Ecology: The Part-Whole Relation in Ecosystems. Prentice Hall PTR, New Jersey, 706 pp. Patten BC, Straskraba M, J?rgensen SE. 1997. Ecosystem emerging 1: conservation. Ecol. Model. 96, 221–284. Peters RH. 1983. The Ecological Implications of Body Size. Cambridge University Press, Cambridge, MA, 329 pp. Peterson RT. 1963. A field guide to the birds of Texas and adjacent states. The Peterson Field Guide Series. Houghton Mifflin, Boston, MA. 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