Pugnaire F.I. Valladares F. Functional Plant Ecology
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Library of Congress Cataloging-in-Publication Data
Functional plant ecology / edited by Francisco Pugnaire and Fernando Valladares. -- 2nd ed.
p. cm. -- (Books in soils, plants, and the environment ; 120)
Rev. ed. of: Handbook of functional plant ecology / edited by Francisco I. Pugnaire, Fernando
Valladares. c1999.
Includes bibliographical references and index.
ISBN 978-0-8493-7488-3
1. Plant ecology. 2. Plant ecophysiology. I. Pugnaire, Francisco I., 1957- II. Valladares, Fernando,
1965- III. Handbook of functional plant ecology. IV. Title. V. Series.
QK901.H295 2007
581.7--dc22 2007000591
Visit the Taylor & Francis Web site at
http://www.taylorandfrancis.com
and the CRC Press Web site at
http://www.crcpress.com
Francisco Pugnaire/Functional Plant Ecology 7488_C000 Final Proof page x 12.5.2007 2:00pm Compositor Name: JGanesan
Table of Contents
Preface.................................................................................................................................xiii
Editors ..................................................................................................................................xv
Contributors ........................................................................................................................xvii
Chapter 1 Methods in Comparative Functional Ecology ....................................................1
Carlos M. Duarte
Chapter 2 Opportunistic Growth and Desiccation Tolerance: The Ecological
Success of Poikilohydrous Autotrophs................................................................7
Ludger Kappen and Fernando Valladares
Chapter 3 Ecological Significance of Inherent Variation in Relative Growth Rate
and Its Components ..........................................................................................67
Hendrik Poorter and Eric Garnier
Chapter 4 The Architecture of Plant Crowns: From Design Rules to Light
Capture and Performance ............................................................................... 101
Fernando Valladares and U
¨
lo Niinemets
Chapter 5 Structure and Function of Root Systems........................................................ 151
Robert B. Jackson, William T. Pockman, William A. Hoffmann, Timothy M. Bleby,
and Cristina Armas
Chapter 6 Water Relations and Hydraulic Architecture.................................................. 175
Melvin T. Tyree
Chapter 7 Responses of Plants to Heterogeneous Light Environments........................... 213
Robert W. Pearcy
Chapter 8 Acquisition, Use, and Loss of Nutrients ......................................................... 259
Frank Berendse, Hans de Kroon, and Wim G. Braakhekke
Chapter 9 Functional Attributes in Mediterranean-Type Ecosystems ............................. 285
Richard Joffre, Serge Rambal, and Claire Damesin
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Chapter 10 Tropical Forests: Diversity and Function of Dominant Life-Forms............. 313
Ernesto Medina
Chapter 11 Plant Diversity in Tropical Forests ...............................................................351
S. Joseph Wright
Chapter 12 Arctic Ecology............................................................................................... 369
Sarah E. Hobbie
Chapter 13 Plant Life in Antarctica ................................................................................. 389
T.G. Allan Green, Burkhard Schroeter, and Leopoldo G. Sancho
Chapter 14 Facilitation in Plant Communities ................................................................ 435
Ragan M. Callaway and Francisco I. Pugnaire
Chapter 15 Plant Interactions: Competition .................................................................... 457
Heather L. Reynolds and Tara K. Rajaniemi
Chapter 16 Plant–Herbivore Interaction: Beyond a Binary Vision.................................. 481
Elena Baraza, Regino Zamora, Jose
´
A. Ho
´
dar, and Jose
´
M. Go
´
mez
Chapter 17 Ecology of Plant Reproduction: Mating Systems and Pollination................ 515
Anna Traveset and Anna Jakobsson
Chapter 18 Seed and Seedling Ecology ............................................................................ 549
Kaoru Kitajima
Chapter 19 Biodiversity and Interactions in the Rhizosphere: Effects
on Ecosystem Functioning ............................................................................581
Susana Rodrı
´
guez-Echeverrı
´
a, Sofia R. Costa, and Helena Freitas
Chapter 20 Resistance to Air Pollutants: From Cell to Community ............................... 601
Jeremy Barnes, Alan Davison, Luis Balaguer, and Esteban Manrique-Reol
Chapter 21 Canopy Photosynthesis Modeling .................................................................627
Wolfram Beyschlag and Ronald J. Ryel
Chapter 22 Ecological Applications of Remote Sensing at Multiple Scales .................... 655
John A. Gamon, Hong-Lie Qiu, and Arturo Sanchez-Azofeifa
Chapter 23 Generalization in Functional Plant Ecology: The Species-Sampling
Problem, Plant Ecology Strategy Schemes, and Phylogeny .......................... 685
Mark Westoby
Index................................................................................................................................... 705
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xii Table of Contents
Preface
Diversity of plant form and life history and their distribution onto different habitats suggest
that plant functions should underlie this diversity, providing tools to successfully and differ-
entially thrive in every habitat. The knowledge of these functions is then the key to under-
stand community and ecosystem structure and functioning, something that attracted the
interest and effort of many plant ecologists trying to establish patterns of adaptive special-
ization in plants.
This volume on Functional Plant Ecology is an updated version of a successful first edition
in which we tried to put together chapters from all areas of plant ecology to provide readers
the broadest view of functional approaches to plant ecology. Our aim was to gather original
reviews with an attractive presentation, giving a comprehensive overview of the topic with a
historical perspective when needed. The book is intended for a broad audience, from plant
ecologists to students, with characteristics of both a textbook and an essay book. We were not
interested in presentation of new experimental data, novel theoretical interpretations, or
hypotheses, but rather asked the authors to provide easy-to-read, up-to-date, and suggestive
introductions to each topic.
Deciding the book composition was not an easy task, as many attractive, substantial
topics emerged at first glimpse. Finally, only a short number made their way into the book,
and we are aware that many important questions have been left out, but practical and
technical reasons limited the extent of the volume. The book follows a bottom-up approach,
from the more specific, detailed studies focusing on plant organs to the broadest ecosystem
approaches, each gathering chapters on the most outstanding aspects.
The history, aims, and potentials of functional approaches are established in the first
chapter, which also sets the limits of functional plant ecology, a science centered in the study
of whole plants and that attempts to predict responses in plant functioning caused by envir-
onmental clues, emphasizing plant influence on ecosystem functions, services, and products,
and aiming to extract patterns and functional laws from comparative analyses. The search for
these patterns is likely to be most effective if driven by specific hypotheses tested on the basis of
comparative analyses at the broadest possible scale. Functional laws thus developed may hold
predictive power irrespective of whether they represent direct cause–effect relationships. Yet,
the nested nature of the control of functional responses implies uncertainties when scaling
functional laws, either toward lower or higher levels of organization.
We would like to express our sincere thanks to the authors who contributed to this
volume for their efforts in updating their chapters and for meeting the deadlines over already
busy timetables. Finally, we want to thank John Sulzycki for his support throughout and Pat
Roberson for her help and patience. All of them made possible and greatly improved the
quality of this work.
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Editors
Francisco I. Pugnaire is a research scientist at the Arid Zones Experimental Station in
Almerı
´
a, Spain, an Institute of the Spanish National Council for Scientific Research. Author
of numerous publications, his research interests focus on physiological ecology and commu-
nity dynamics and on the effects of current environmental changes on biodiversity. He is
actively involved in several EU and national initiatives in the science–policy interface.
Centered on semiarid environments, he has also worked in desert ecosystems, tropical forests,
and high mountain environments. Member of the editorial board of several international
journals, he is also a member of the ecological societies of Spain, UK, and USA and vice
president of the European Ecological Federation.
Fernando Valladares is currently a senior scientist of the Spanish Research Council (CSIC) at
the Centre of Environmental Sciences and associate professor at the Rey Juan Carlos
University of Madrid. He has published more than 150 scientific articles, and is the author
or coauthor of 5 books. Dr. Valladares is a member of the editorial board of three of the most
prestigious international journals of plant ecology and physiology, and he is an active
member of the main ecological societies of Spain, UK, and USA. Dr. Valladares is involved
in several panels and expert committees addressing global change issues and transferring
ecological knowledge to society and policy makers at both national and international levels.
His research on functional plant ecology and evolution is focused on phenotypic plasticity
and on the physiological and morphological strategies of photosynthetic organisms to cope
with changing and adverse environmental conditions. The research has been carried out in a
range of ecosystems, spanning from Antarctica to the tropical rain forest, although his
current research objectives are centered on low-productivity Mediterranean ecosystems.
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Contributors
Cristina Armas
Department of Biology
Duke University
Durham, North Carolina
Luis Balaguer
Department of Plant Biology
Complutense University
Madrid, Spain
Elena Baraza
Laboratory of Ecology of Communities
Institute of Ecology
UNAM, Mexico D.F.
Jeremy Barnes
Institute for Research on the Enviornment
and Sustainability
Newcastle University
Newcastle upon Tyne, England
Frank Berendse
Centre for Ecosystem Studies
Wageningen University
Wageningen, The Netherlands
Wolfram Beyschlag
Department of Experimental and
Systems Ecology
University of Bielefeld
Bielefeld, Germany
Timothy M. Bleby
Department of Biology
Duke University
Durham, North Carolina
Wim G. Braakhekke
Centre for Ecosystem Studies
Wageningen University
Wageningen, The Netherlands
Ragan M. Callaway
Division of Biological Sciences
University of Montana
Missoula, Montana
Sofia R. Costa
Department of Botany
University of Coimbra
Coimbra, Portugal
Claire Damesin
Plant Ecophysiology
Paris-Sud University
Orsay, France
Alan Davison
School of Biology and Psychology:
Division of Biology
Newcastle University
Newcastle upon Tyne, England
Carlos M. Duarte
Mediterranean Institute for
Advanced Studies
Spanish Council for Scientific
Research
Blanes, Spain
Helena Freitas
Department of Botany
University of Coimbra
Coimbra, Portugal
John A. Gamon
Department of Biology and
Microbiology
California State University
Los Angeles, California
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xvii
Eric Garnier
Centre for Functional and Evolutionary
Ecology
CNRS
Montpellier, France
Jose
´
M. Go
´
mez
Department of Ecology
University of Granada
Granada, Spain
T.G. Allan Green
Department of Biological Sciences
Waikato University
Hamilton, New Zealand
Sarah E. Hobbie
Department of Ecology, Evolution,
and Behavior
University of Minnesota
St. Paul, Minnesota
Jose
´
A. Ho
´
dar
Terrestrial Ecology Group
Department of Ecology
University of Granada
Granada, Spain
William A. Hoffmann
Department of Plant Biology
North Carolina State University
Raleigh, North Carolina
Robert B. Jackson
Department of Biology
Duke University
Durham, North Carolina
Anna Jakobsson
Mediterranean Institute for
Advanced Studies
Esporles, Mallorca
Balearic Islands, Spain
Richard Joffre
Centre for Functional and Evolutionary
Ecology
CNRS
Montpellier, France
Ludger Kappen
Department of Botany
University of Kiel
Kiel, Germany
Kaoru Kitajima
Department of Botany
University of Florida
Gainesville, Florida
Hans de Kroon
Department of Experimental Plant Ecology
Institute of Water and Wetland Research
Radboud University
Nijmegen, The Netherlands
Esteban Manrique-Reol
Department of Agricultural and
Environmental Science
University of Newcastle
Newcastle upon Tyne, England
Ernesto Medina
Center of Ecology
Venezuelan Institute for Scientific
Investigations
Caracas, Venezuela
U
¨
lo Niinemets
Institute of Environment and
Agriculture
Estonian University of Life Sciences
Tartu, Estonia
Robert W. Pearcy
Division of Biological Sciences
University of California
Davis, California
William T. Pockman
Department of Biology
University of New Mexico
Albuquerque, New Mexico
Hendrik Poorter
Institute of Environmental Biology
Utrecht University
Utrecht, The Netherlands
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xviii Contributors
Francisco I. Pugnaire
Arid Zones Research Station
Spanish Council for Scientific Research
Almeria, Spain
Hong-Lie Qiu
Department of Geography and Urban
Analysis
California State University
Los Angeles, California
Tara K. Rajaniemi
Biology Department
University of Massachusetts
Dartmouth, Massachusetts
Serge Rambal
Centre for Functional and Evolutionary
Ecology
CNRS
Montpellier, France
Heather L. Reynolds
Department of Biology
Indiana University
Bloomington, Indiana
Susana Rodrı
´
guez-Echeverrı
´
a
Department of Botany
University of Coimbra
Coimbra, Portugal
Ronald J. Ryel
Department of Wildland Resources
and the Ecology Center
Utah State University
Logan, Utah
Arturo Sanchez-Azofeifa
Department of Earth and Atmospheric
Sciences
University of Alberta
Edmonton, Alberta, Canada
Leopoldo G. Sancho
Department Vegetal II
Complutense University
Madrid, Spain
Burkhard Schroeter
Botanical Institute
University of Kiel
Kiel, Germany
and
IPN – Leibniz Institute for Science
Education
University of Kiel
Kiel, Germany
Anna Traveset
Mediterranean Institute for Advanced
Studies
Esporles, Mallorca
Balearic Islands, Spain
Melvin T. Tyree
Aiken Forestry Sciences Laboratory
Burlington, Vermont
Fernando Valladares
Centre for Environmental Studies
Spanish Council for Scientific
Research
Madrid, Spain
Mark Westoby
Department of Biological Sciences
Macquarie University
Sydney, Australia
S. Joseph Wright
Smithsonian Tropical Research
Institute
Balboa, Republic of Panama
Regino Zamora
Department of Ecology
University of Granada
Granada, Spain
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Contributors xix