Tài liệu Aquatic Food Webs An Ecosystem Approach docx

Aquatic Food Webs

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Aquatic Food Webs

An Ecosystem Approach

EDITED BY

Andrea Belgrano

National Center for Genome Resources (NCGR),

Santa Fe, NM, USA

Ursula M. Scharler

University of Maryland Center for Environmental Science,

Chesapeake Biological Laboratory (CBL),

Solomons, MD, USA and Smithsonian Environmental Research Center,

Edgewater, MD, USA

Jennifer Dunne

Pacific Ecoinformatics and Computational Ecology Lab, Berkeley,

CA USA; Santa Fe Institute (SFI) Santa FE, NM, USA; Rocky Mountain

Biological Laboratory, Crested Butte, CO USA

AND

Robert E. Ulanowicz

University of Maryland Center for Environmental Science,

Chesapeake Biological Laboratory (CBL),

Solomons, MD, USA

1

1

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British Library Cataloging in Publication Data

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Library of Congress Cataloging-in-Publication Data

Aquatic food webs : an ecosystem approach / edited by Andrea Belgrano ... [et al.].

p. cm.

Includes bibliographical references and index.

ISBN 0-19-856482-1 (alk. paper) – ISBN 0-19-856483-X (alk. paper) 1. Aquatic

ecology. 2. Food chains (Ecology) I. Belgrano, Andrea.

QH541.5.W3A68225 2004

577.60

16–dc22 2004027135

ISBN 0 19 856482 1 (Hbk) 9780198564829

ISBN 0 19 856483 X (Pbk) 9780198564836

10 9 8 7 6 5 4 3 2 1

Typeset by Newgen Imaging Systems (P) Ltd., Chennai, India

Printed in Great Britain

on acid-free paper by Antony Rowe, Chippenham

FOREWORD

CURRENT AND FUTURE PERSPECTIVES

ON FOOD WEBS

Michel Loreau

Food webs have been approached from two basic

perspectives in ecology. First is the energetic view

articulated by Lindeman (1942), and developed by

ecosystem ecology during the following decades.

In this view, food webs are networks of pathways

for the flow of energy in ecosystems, from its

capture by autotrophs in the process of photo￾synthesis to its ultimate dissipation by hetero￾trophic respiration. I would venture to say that the

ecological network analysis advocated by

Ulanowicz and colleagues in this book is heir to

this tradition. A different approach, rooted in

community ecology, was initiated by May (1973)

and pursued by Pimm (1982) and others. This

approach focuses on the dynamical constraints

that arise from species interactions, and empha￾sises the fact that too much interaction (whether in

the form of a larger number of species, a greater

connectance among these species, or a higher

mean interaction strength) destabilises food webs

and ecological systems. The predictions resulting

from this theory regarding the diversity and con￾nectance of ecological systems led to a wave of

comparative topological studies on the structure of

food webs. Thus, the two traditions converge in

the search for patterns in food-web structure

despite different starting points. This book results

from the confluence of these two perspectives,

which are discussed in a number of chapters.

Patterns, however, are generally insufficient to

infer processes. Thus, the search for explanations of

these patterns in terms of processes is still very much

alive, and in this search the energetic and dynamical

perspectives are not the only possible ones. Bio￾geochemical cycles provide a functional perspective

on food webs that is complementary to the energetic

approach (DeAngelis 1992). Material cycles are

among the most common of the positive feedback

loops discussed by Ulanowicz in his concluding

remarks, and may explain key properties of eco￾systems (Loreau 1998). The stoichiometry of ecolo￾gical interactions may further strongly constrain

food-web structure (Sterner and Elser 2002; Elser

and Hessen’s chapter). There has also been con￾siderable interest in the relationship between bio￾diversity and ecosystem functioning during the last

decade (Loreau et al. 2002).Merging the theories that

bear upon food webs and the maintenance of species

diversity is urgently needed today, and may provide

new insights into food-webs structure and ecosys￾tem functioning (Hillebrand and Shurin’s chapter).

The structure and functioning of ecological sys￾tems is determined not only by local constraints

and interactions, but also by larger-scale processes.

The importance of regional and historical influ￾ences has been increasingly recognised in com￾munity ecology (Ricklefs and Schluter 1993). The

extent to which they shape food webs, however,

has been relatively little explored. The recent

development of metacommunity theory (Leibold

et al. 2004) provides a framework to start exam￾ining spatial constraints on the structure and

functioning of local food webs (Melian et al.‘s

chapter). At even larger time scales, food webs are

the result of evolutionary processes which deter￾mine their current properties. Complex food webs

may readily evolve based on simple ecological

interactions (McKane 2004). The evolution of food￾web and ecosystem properties is a fascinating

topic for future research.

v

This book provides a good synthesis of recent

research into aquatic food webs. I hope this

synthesis will stimulate the development of new

approaches that link communities and ecosystems.

References

DeAngelis, D. L. 1992. Dynamics of nutrient cycling and

food webs. Chapman & Hall, London.

Leibold, M. A., M. Holyoak, N. Mouquet, P. Amarasekare,

J.M. Chase,M. F. Hoopes, R. D. Holt, J. B. Shurin, R. Law,

D. Tilman, M. Loreau, and A. Gonzalez. 2004. The

metacommunity concept: a framework for multi-scale

community ecology. Ecology Letters 7: 601–613.

Lindeman, R. L. 1942. The trophic-dynamic aspect of

ecology. Ecology 23: 399–418.

Loreau, M. 1998. Ecosystem development explained by

competition within and between material cycles. Pro￾ceedings of the Royal Society of London, Series B 265: 33–38.

Loreau, M., S. Naeem, and P. Inchausti. Eds. 2002. Bio￾diversity and ecosystem functioning: synthesis and per￾spectives. Oxford University Press, Oxford.

May, R. M. 1973. Stability and complexity in model ecosys￾tems. Princeton University Press, Princeton.

McKane, A. J. 2004. Evolving complex food webs. The

European Physical Journal B 38: 287–295.

Pimm, S. L. 1982. Food webs. Chapman & Hall, London.

Ricklefs, R. E., and D. Schluter. Eds 1993. Species diversity

in ecological communities: historical and geographical per￾spectives. University of Chicago Press, Chicago.

Sterner, R. W., and J. J. Elser. 2002. Ecological stoichiometry:

the biology of elements from molecules to the biosphere.

Princeton University Press, Princeton.

vi FOREWORD

Contents

Foreword v

Michel Loreau

Contributors ix

Introduction 1

Andrea Belgrano

PART I Structure and function 5

1 Biosimplicity via stoichiometry: the evolution of food-web

structure and processes 7

James J. Elser and Dag O. Hessen

2 Spatial structure and dynamics in a marine food web 19

Carlos J. Melia´n, Jordi Bascompte, and Pedro Jordano

3 Role of network analysis in comparative ecosystem

ecology of estuaries 25

Robert R. Christian, Daniel Baird, Joseph Luczkovich, Jeffrey C. Johnson,

Ursula M. Scharler, and Robert E. Ulanowicz

4 Food webs in lakes—seasonal dynamics and the impact

of climate variability 41

Dietmar Straile

5 Pattern and process in food webs: evidence from running waters 51

Guy Woodward, Ross Thompson, Colin R. Townsend, and Alan G. Hildrew

PART II Examining food-web theories 67

6 Some random thoughts on the statistical analysis of food-web data 69

Andrew R. Solow

7 Analysis of size and complexity of randomly constructed food

webs by information theoretic metrics 73

James T. Morris, Robert R. Christian, and Robert E. Ulanowicz

8 Size-based analyses of aquatic food webs 86

Simon Jennings

vii

9 Food-web theory in marine ecosystems 98

Jason S. Link, William T. Stockhausen, and Elizabeth T. Methratta

PART III Stability and diversity in food webs 115

10 Modeling food-web dynamics: complexity–stability implications 117

Jennifer A. Dunne, Ulrich Brose, Richard J. Williams, and Neo D. Martinez

11 Is biodiversity maintained by food-web complexity?—the

adaptive food-web hypothesis 130

Michio Kondoh

12 Climate forcing, food web structure, and community dynamics

in pelagic marine ecosystems 143

L. Ciannelli, D. Ø. Hjermann, P. Lehodey, G. Ottersen,

J. T. Duffy-Anderson, and N. C. Stenseth

13 Food-web theory provides guidelines for marine conservation 170

Enric Sala and George Sugihara

14 Biodiversity and aquatic food webs 184

Helmut Hillebrand and Jonathan B. Shurin

PART IV Concluding remarks 199

15 Ecological network analysis: an escape from the machine 201

Robert E. Ulanowicz

Afterword 208

Mathew A. Leibold

References 211

Index 255

viii CONTENTS

Contributors

Daniel Baird, Zoology Department, University of Port

Elizabeth, Port Elizabeth, South Africa.

Jordi Bascompte, Integrative Ecology Group, Estacio´n

Biolo´gica de Don˜ana, CSIC, Apdo. 1056, E-41080,

Sevilla, Spain. Email: [email protected]

Andrea Belgrano, National Center for Genome Resources

(NCGR), 2935 Rodeo Park Drive East, Santa Fe, NM

87505, USA. Email: [email protected]

Ulrich Brose, Technical University of Darmstadt,

Department of Biology, Schnittspahnstr. 3, 64287

Darmstadt, Germany.

Robert R. Christian, Biology Department, East Carolina

University, Greenville, NC 27858, USA. Email:

[email protected]

Lorenzo Ciannelli, Centre for Ecological and

Evolutionary Synthesis (CEES), Department of Biology

University of Oslo, Post Office Box 1066, Blindern,

N-0316 Oslo, Norway. Email: lorenzo.ciannelli@

bio.uio.no.

Janet T. Duffy-Anderson, Alaska Fisheries Science

Center, NOAA, 7600 Sand Point Way NE, 98115

Seattle, WA, USA.

Jennifer A. Dunne, Pacific Ecoinformatics and Computa￾tional Ecology Lab, P.O. Box 10106, Berkeley, CA 94709

USA; Santa Fe Institute, 1399 Hyde Park Road, Santa Fe,

NM 87501 USA; Rocky Mountain Biological Laboratory,

P.O. Box 519, Crested Butte, CO 81224 USA Email:

[email protected].

James J. Elser, School of Life Sciences, Arizona State

University, Tempe, AZ 85287, USA. Email: j.elser@

asu.edu

Dag O. Hessen, Department of Biology, University of

Oslo, P.O. Box 1050, Blindern, N-0316 Oslo, Norway.

Alan G. Hildrew, School of Biological Sciences, Queen

Mary, University of London, Mile End Road, London,

E1 4NS, UK. Email: [email protected]

Helmut Hillebrand, Institute for Botany, University of

Cologne, Gyrhofstrasse 15 D-50931 Ko¨ln, Germany.

Email: [email protected]

D.Ø. Hjermann, Centre for Ecological and Evolutionary

Synthesis (CEES), Department of Biology University of

Oslo, Post Office Box 1066 Blindern, N-0316 Oslo,

Norway.

Simon Jennings, Centre for Environment, Fisheries

and Aquaculture Science, Lowestoft Laboratory NR33

0HT, UK. Email: [email protected]

Jeffrey C. Johnson, Institute of Coastal and Marine

Resources, East Carolina University, Greenville, NC

27858, USA.

Pedro Jordano, Integrative Ecology Group, Estacio´n

Biolo´gica de Don˜ana, CSIC, Apdo. 1056, E-41080,

Sevilla, Spain.

Michio Kondoh, Center for Limnology, Netherlands

Institute of Ecology, Rijksstraatweg 6, Nieuwersluis,

P.O. Box 1299, 3600 BG Maarssen, The Netherlands.

Email: [email protected]

P. Lehodey, Oceanic Fisheries Programme, Secretariat

of the Pacific Community, BP D5, 98848 Noumea

cedex, New Caledonia.

Mathew Leibold, Section of Integrative Biology, The

University of Texas at Austin, 1 University Station,

C0930 Austin, TX 78712, USA. Email: mleibold@

mail.utexas.edu

Jason S. Link, National Marine Fisheries Service,

Northeast Fisheries Science Center, 166 Water St.,

Woods Hole, MA 02543, USA. Email: jlink@

whsunl.wh.whoi.edu

Michel Loreau, Laboratoire d’Ecologie, UMR 7625

Ecole Normale Superieure 46, rue d’ Ulm F-75230,

Paris Cedex 05, France. Email: [email protected]

Joseph Luczkovich, Biology Department, East Carolina

University, Greenville, NC 27858, USA.

Neo D. Martinez, Pacific Ecoinformatics and

Computational Ecology Lab, P.O. Box 10106,

Berkeley, CA 94709 Rocky Mountain Biological

Laboratory, P.O. Box 519, Crested Butte, CO 81224

USA.

Carlos J. Melia´n, Integrative Ecology Group, Estacio´n

Biolo´gica de Don˜ana, CSIC, Apdo. 1056, E-41080,

Sevilla, Spain.

Elizabeth T. Methratta, National Marine Fisheries

Service, Northeast Fisheries Science Center,

166 Water St., Woods Hole, MA 02543, USA.

James T. Morris, Department of Biological Sciences,

University of South Carolina, Columbia, SC 29208, USA.

Email: [email protected]

ix

Geir Ottersen, Institute of Marine Research, P.O. Box

1870 Nordnes, 5817 Bergen, NORWAY Current

Address: Centre for Ecological and Evolutionary

Synthesis, Department of Biology, P.O Box 1050

Blindern, N-0316 Oslo, NORWAY.

Enric Sala, Center for Marine Biodiversity and

Conservation, Scripps Institution of Oceanography,

La Jolla, CA 92093-0202, USA. Email:

[email protected]

Ursula M. Scharler, University of Maryland,

Center for Environmental Science, Chesapeake

Biological Laboratory (CBL), Solomons, MD 20688,

USA; Smithsonian Environmental Research

Center, Edgewater, MD, USA. Email: scharler@

cbl.umces.edu

Jonathan B. Shurin, Department of Zoology,

University of British Columbia, 6270 University Blvd.

Vancouver, BC V6T 1Z4, Canada.

Nils Chr. Stenseth, Centre for Ecological and

Evolutionary Synthesis (CEES), Department of Biology

University of Oslo, Post Office Box 1066 Blindern, N￾0316 Oslo, Norway.

Dietmar Straile, Dietmar StraileLimnological

Institute, University of Konstanz, 78457 Konstanz,

Germany. Email: dietmar.straile@

uni-konstanz.de

William T. Stockhausen, National Marine Fisheries

Service, Northeast Fisheries Science Center,

166 Water St., Woods Hole, MA 02543, USA.

Andrew R. Solow, Woods Hole Oceanographic

Institution, Woods Hole, MA 02543, USA. Email:

[email protected]

Geroge Sugihara, Center for Marine Biodiversity and

Conservation, Scripps Institution of Oceanography,

La Jolla, CA 92093-0202, USA.

Ross Thompson, Biodiversity Research Centre,

University of British Columbia, Vancouver, Canada.

Colin R. Townsend, Department of Zoology,

University of Otago, New Zealand.

Robert E. Ulanowicz, University of Maryland, Center

for Environmental Science, Chesapeake Biological

Laboratory (CBL), Solomons, MD 20688, USA. Email:

[email protected]

Richard J. Williams, Pacific Ecoinformatics and

Computational Ecology Lab, P.O. Box 10106, Berkeley,

CA 94709 USA; Rocky Mountain Biological Laboratory,

P.O. Box 519, Crested Butte, CO 81224 USA; San Fran￾cisco State University, Computer Science Department,

1600 Holloway Avenue, San Francisco, CA 94132 USA.

Guy Woodward, Department of Zoology, Ecology

and Plant Science, University College, Cork,

Ireland.

x CONTRIBUTORS

INTRODUCTION

Aquatic food-webs’ ecology:

old and new challenges

Andrea Belgrano

Looking up ‘‘aquatic food web’’ on Google provides

a dizzying array of eclectic sites and information

(and disinformation!) to choose from. However,

even within this morass it is clear that aquatic

food-web research has expanded greatly over the

last couple of decades, and includes a wide array

of studies from both theoretical and empirical

perspectives. This book attempts to bring together

and synthesize some of the most recent perspec￾tives on aquatic food-web research, with a parti￾cular emphasis on integrating that knowledge

within an ecosystem framework.

It is interesting to look back at the pioneering

work of Sir Alister Hardy in the early 1920s at

Lowestoft Fisheries Laboratory. Hardy studied the

feeding relationship of the North Sea herring with

planktonic assemblages by looking at the species

distribution patterns in an attempt to provide

better insights for the stock assessment of the

North Sea fisheries. If we take a look in his food￾web scheme (Figure 1), it is interesting to note that

he considered species diversity in both phyto￾plankton and zooplankton, and also specified

body-size data for the different organisms in the

food web. Thus, it appears that already almost

100 years ago the concept of constructing and

drawing links among diverse species at multiple

trophic levels in a network-like fashion was in the

mind of many aquatic researchers.

In following decades, researchers began to

consider links between food-web complexity and

ecological community stability. The classic, and still

contentious MacArthur hypothesis that ‘‘Stability

increases as the number of link increase’’ (1955)

gave rise to studies such as that by Paine (1966)

that linked latitudinal gradients in aquatic species

diversity, food-web complexity, and community

stability.

Following that early MacArthur hypothesis, we

find it timely to also ask, How complex are aquatic

food webs?

The first book on theoretical food-web ecology

was written by May (1973), followed by Cohen

(1978). Since then, Pimm (1982) and Polis and

Winemiller (1996) have revisited some of the ideas

proposed by May and Cohen and discussed them

in different contexts, and trophic flow models have

been proposed and used widely for aquatic and

particularly marine ecosystems (e.g. Wulff et al.

1989; Christensen and Pauly 1993). However,

recent advances in ecosystem network analysis

(e.g. Ulanowicz 1996, 1997; Ulanowicz and Abarca￾Arenas 1997) and the network structure of food

webs (e.g. Williams and Martinez 2000; Dunne

et al. 2002a,b; Williams et al. 2002) in relation to

ecosystem dynamics, function, and stability clearly

set the path for a new, complementary research

agenda in food-web analysis. These and many

other studies suggest that a new synthesis of

available information is necessary. This new

synthesis is giving rise to novel basic research that

generalizes across habitats and scales, for example,

the discovery of universal scaling relations in food￾web structure (Garlaschelli et al. 2003), and is also

underpinning new approaches and priorities for

whole-ecosystem conservation and management,

particularly in marine systems.

Aquatic food-web research is also moving beyond

an exclusive focus on taxa from phytoplankton

to fish. A new look at the role that marine microbes

1

Figure 2 The microbial loop: impressionist version.

A bacteria-eye view of the ocean’s euphotic layer.

Seawater is an organic matter continuum, a gel of

tangled polymers with embedded strings, sheets,

and bundles of fibrils and particles, including living

organisms, as ‘‘hotspots.’’ Bacteria (red) acting

on marine snow (black) or algae (green) can

control sedimentation and primary productivity;

diverse microniches (hotspots) can support high

bacterial diversity. (Azam, F. 1998. Microbial

control of oceanic carbon flux: the plot thickens.

Science 280: 694–696.) (See Plate1)

Figure 1 The food web of herring Clupea harengus Hardy (1924). From Parables of Sea & Sky—The life, work and art of Sir Alister

Hardy F. R. S. Courtesy of SAHFOS—The CPR Survey, Plymouth, UK.

2 INTRODUCTION

play in the global ocean (Azam and Worden 2004)

suggests that oceanic ecosystems can be character￾ized as a complex dynamic molecular network.

The role of microbial food webs (Figure 2—see

also, Plate 1—Azam 1998) needs to be considered

to understand the nonlinearities underlying the

relationship between the pelagic and benthic

domains.

Emerging challenges in aquatic food-web research

include integrating genomic, biogeochemical,

environmental, and economic data in a modeling

effort that will elucidate the mechanisms govern￾ing the ecosystem dynamics across temporal and

spatial scales at different levels of organization

and across the whole variety of species diversity,

including humans. Aquatic food webs may pro￾vide a particularly useful empirical framework for

developing and testing an information theory of

ecology that will take into account the complex

network of interactions among biotic and abiotic

components of ecosystems.

Acknowledgments

This work was funded in part or in full by the US

Dept of Energy’s Genomes to Life program (www.

doegenomestolife.org) under the project ‘‘Carbon

Sequestration in Synechococcus sp.: From Mole￾cular Machines to Hierarchical Modeling’’ (www.

genomes-to-life.org).

INTRODUCTION 3

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