A New Ecology - Systems Perspective - Intro ppsx

vi Contents5.5 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 envi

A New Ecology: Systems Perspective

Front cover photo is by B.D Fath and shows Møns Klint, Denmark.

The back cover photos show (from left to right) (1) a danish beech forest (Ryget Skov), (2) Krimml Falls in Austria, (3) part of the shore of Namchu Lake in Tibet, (4) Crater Lake, Oregan, USA, and (5) Natron Lake, Tanzania and were taken by S.E Jørgensen (1 and 4), B.D Fath (2), and M.V Jørgensen (3 and 5)

A New Ecology

Systems Perspective

Sven E Jørgensen Environmental Chemistry Section Royal Danish School of Pharmacy DK-2100 Copenhagen, Denmark Simone Bastianoni Department of Chemical and Biosystems Sciences University of Siena

53100 Siena, Italy Felix Müller Ecology Centre University of Kiel

24118 Kiel, Germany Bernard C Patten Institute of Ecology University of Georgia Athens, GA 30602-2602, USA

Brian D Fath Biology Department Towson University Towson, MD 21252, USA João C Marques Department of Zoology Institute of Marine Research (IMAR), University of Coimbra 3004-517 Coimbra, Portugal Søren N Nielsen Environmental Chemistry Section Royal Danish School of Pharmacy DK-2100 Copenhagen, Denmark 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

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CONTENTS Preface ix

1.1 Environmental management has changed 1

2.1 Why must ecosystems be open? 7 2.2 An isolated system would die (maximum entropy) 8

2.4 The second law of thermodynamics interpreted for open systems 18

2.6 Quantification of openness and allometric principles 22

2.8 What about the environment? 31

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

4.1 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

4.8 Directionality in evolution? 76

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vi Contents

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

6.1 Variability in life conditions 103

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

7.1 The normality of disturbance 143 7.2 The risk of orientor optimization 151 7.3 The characteristics of disturbance 152 7.4 Adaptability as a key function of ecosystem dynamics 156 7.5 Adaptive cycles on multiple scales 160 7.6 A case study: Human disturbance and retrogressive dynamics 164

8.2 Do ecological principles encompass other proposed ecological theories?: Evolutionary theory 168 8.3 Do ecological principles encompass other proposed

ecological theories?: Island biogeography 176 8.4 Do ecological principles encompass other proposed

ecological theories?: Latitudinal gradients in biodiversity 180 8.5 Do ecological principles encompass other proposed

ecological theories?: Optimal foraging theory 184 8.6 Do ecological principles encompass other proposed

ecological theories?: Niche theory 187

8.7 Do ecological principles encompass other proposed ecological theories?: Liebig’s law of the minimum 191 8.8 Do ecological principles encompass other proposed

ecological theories?: The river continuum concept (RCC) 194 8.9 Do ecological principles encompass other proposed

ecological theories?: Hysteresis in nature 196

9.2 Entropy production as an indicator of ecosystem trophic state 200 9.3 The use of ecological network analysis (ENA) for the simulation

of the interaction of the american black bear and its environment 206 9.4 Applications of network analysis and ascendency to South Florida

9.5 The application of eco-exergy as ecological indicator for assessment of ecosystem health 218 9.6 Emergy as ecological indicator to assess ecosystem health 221 9.7 The eco-exergy to empower ratio and the efficiency of ecosystems 228 9.8 Application of eco-exergy and ascendency as ecological indicator

to the Mondego Estuary ( Portugal) 231

10.1 Are basic ecological properties needed to explain our observations? 243 10.2 Previous attempts to present an ecosystem theory 243 10.3 Recapitulation of the ecosystem theory 245 10.4 Are there basic ecosystem principles? 246

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The scope of this book is to demonstrate that we do have an ecosystem theory that can be used to describe ecosystem structure and function It was previously shown in the book,

Integration of Ecosystem Theories: A Pattern (3rd edition, 2002), that the various

contri-butions to systems ecology are consistent and together form a pattern of ecological

processes My book with Yuri Svirezhev, Toward a Thermodynamic Theory of Ecosystems

(2004), presented the thermodynamics of this pattern in a mathematical language This

book, A New Ecology: Systems Perspective, shows that the basic properties of ecosystems

(presented in Chapters 2–7) lead to or are consistent with ten tentative propositions for ecosystems (Chapter 10), which can be used to explain ecological observations (Chapter 8)

An ecosystem theory is a prerequisite for wider application of ecological sciences in envi-ronmental management because with the theory it becomes feasible to guide conservation

or environmental management Chapter 9 shows how the presented ecosystem theory can

be applied to assess ecosystem health, a facet of environmental management A thermody-namic interpretation of the evolution is under preparation in my other book with Yuri

Svirezhev, A Thermodynamic Theory of the Evolution, with expected publication in 2007 or early in 2008 The three books Toward a Thermodynamic Theory of Ecosystems, this book

A New Ecology: Systems Perspective, and the coming one, A Thermodynamic Theory of the Evolution form a troika that presents a useful ecological theory.

This book has nine authors The basic outline of the book was formulated during a one-week brainstorming meeting on the Danish island of Møn in June 2005 All nine authors have written parts of the book and have reviewed the contributions of the other authors The book is therefore a joint effort resulting from close teamwork I am the first author because the idea to produce a book about ecosystem theory and systems ecology was initi-ated by me based on a brainstorming meeting with system ecologists I edited this book with Brian Fath after all the authors had exchanged ideas and reviewed the ten chapters Brian Fath is therefore considered the second editor of the book Bai Lian Li (Larry) parti-cipated in the brainstorming meeting in Møn and he contributed significantly to the out-line of ideas making up the final book However, due to his engagement with the Eco-summit 2007 in China, he was unable to contribute written material for the book He

is, however, working on a Chinese edition of the book, which we all consider of great importance as China during the last few years has shown an increased interest in envi-ronmental problems This Chinese interest for envienvi-ronmentally sound management is expected to accelerate in the coming years, which makes a Chinese edition of this book even more important Bernie Patten and Enzo Tiezzi were unable to attend the brain-storming meeting, but they both contributed written material and comments on the chapters ( Photos 1 and 2)

Using my 2004 Stockholm Water Prize, I established a foundation to promote ecosys-tem theory and integrated environmental assessment The Foundation’s grants support brainstorming meetings and travel particularly for young scientists focusing on system ecology, ecological modelling, and lake management The foundation is named “William

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x Preface

Photo 1: From the brainstorming meeting at Møn, June 2005.

Photo 2: From the brainstorming meeting at Møn, June 2005.

Williams’ and Milan Straškraba’s Foundation” after two of my close scientist friends who passed away in 2002 and 2000 William Williams has contributed significantly to inte-grated lake management and Milan Straškraba has played a major role in system ecology

in the last two decades of the 20th century The nine authors express their appreciation to the foundation for the support that has made it possible to publish this book in the hope that it will enhance a wider application of ecosystem theory in ecology to explain obser-vations and to facilitate ecological sound conservation and environmental management

Sven Erik Jørgensen Copenhagen, July 2006

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