Table Of ContentContributors
srebmuN in parentheses indicate eht segap no which eht "srohtua snoitubirtnoc .nigeb
David R. Bellwood ,5( 33), Centre for Coral Reef Biodiver- PhiUip .S Levin (377), National Marine Fisheries Service,
sity, School of Marine Biology and Aquaculture, James Cook Seattle, Washington 98112
University, Townsville, Queensland 4811, Australia
Mark I. McCormick (171, 221), School of Marine Biology
M. Julian Caley (81), School of Marine Biology and Aquacul- and Aquaculture, James Cook University, Townsville,
ture, James Cook University, Townsville, Queensland 4811, Queensland 4811, Australia
Australia
Philip .L Munday (81), Centre for Coral Reef Biodiversity,
J. Howard Choat (57), School of Marine Biology and Aqua- School of Marine Biology and Aquaculture, James Cook
culture, James Cook University, Townsville, Queensland
University, Townsville, Queensland 4811, Australia
4811, Australia
Arthur A. Myrberg, Jr. (123), Rosensteil School of Marine and
Robert K. Cowen (149), Rosensteil School of Marine and
Atmospheric Science, University of Miami, Miami, Florida
Atmospheric Science, University of Miami, Miami, Florida
33149
33149
Christopher .W Petersen (103), College of the Atlantic,
Peter .J Doherty 23( ,)7 Australian Institute of Marine ,ecneicS
Bar Harbor, Maine 04609
Cape Ferguson, Townsville, Queensland 4810, Australia
Graham E. Forrester (275), Department of Biological Sci- Serge Planes (201), Centre de Biologie et d'Ecologie Tropi-
ences, University of Rhode lsland, Kingston, Rhode Island cale et Mdditerran~e, EPHE ESA 8046 CNRS, Universitd de
02881 Perpignan, Perpignan 66860, France
Lee A. Fuiman (123), Department of Marine Science, Uni- D. Ross Robertson (57), Smithsonian Tropical Research In-
versity of Texas at Austin, Marine Science Institute, Port stitute, Balboa, Panama
Aransas, Texas 78373
Garry R. Russ (421), School of Marine Biology and Aquacul-
Churchill B. Grimes 3( 77), National Marine seirehsiF ,ecivreS ture, James Cook University, Townsville, Queensland 4811,
Southwest Fisheries Science Center, Santa Cruz Laboratory, Australia
Santa Cruz, California 95060
Yvonne .J Sadovy (391), Department of Ecology & Biodiver-
Jonathan A. Hare (243), National Oceanic and Atmospheric
sity, The University of Hong Kong, Hong Kong
Administration, National Ocean Service, National Centers
for Coastal Ocean Science, Center for Coastal Fisheries and Peter .F Sale (361), Department of Biological Sciences
Habitat Research, Beaufort, North Carolina 28516 and Great Lakes Institute for Environmental Research,
University of Windsor, Windsor, Ontario, Canada N9B
Mireille L. Harmelin-Vivien (265), Centre d'Ocdanologie de
3P4
Marseille, CNRS UMR 6540, Universit8 de al -e~narretid~M
Station Marine d'Endoume 13007 Marseille, France Mark A. Steele (275), Department of Biological Sci-
ences, University of Rhode Island, Kingston, Rhode Island
Mark A. Hixon (303), Department of Zoology, Oregon State
02881
University, Corvallis, Oregon 97331
Geoffrey .P Jones (81,221), Centre for Coral Reef Biodiver- Simon R. Thorrold (243), Biology Department, Woods
sity, School of Marine Biology and Aquaculture, James Cook Hole Oceanographic Institution, Woods Hole, Massachusetts
University, Townsville, Queensland 4811, Australia 02543
Jeffrey M. Lcis (171), Ichthyology and Center for Biodiver- Richard R. Vance (275), Department of Organismic Bio-
sity and Conservation Research, Australian Museum, Sydney logy, Ecology and Evolution, University of California at
NSW 2010, Australia Los Angeles, Los Angeles, California 90095
xi
xii Contributors
Amanda C. J. Vincent (391), Project ,esrohaeS Department Robert R. Warner (103), Department of ,ygolocE Evolution,
of Biology, McGill ,ytisrevinU Penfield, Montreal, Quebec and Marine ,ygoloiB University of ,ainrofilaC Santa ,arabraB
H3 A 1B1, adanaC atnaS ,arabraB California 93106
Peter C. Wainwright ,5( 33), Center for Population ,ygoloiB Michael .S Webster (303), Department of Zoology, Oregon
ytisrevinU of ainrofilaC at Davis, Davis, California 95616 State ,ytisrevinU Corvallis, Oregon 97331
eca:erP
T
he impetus to produce this book came in a brief demands, and for fulfilling my requests, usually in a
phone call in 1998. Chuck Crumly, of Academic timely manner.
Press, called with an invitation and a deadline. Either The book is divided into four sections with a brief
The Ecology of Fishes on Coral Reefs, published in introduction to each. While the sections group together
1991, would be allowed to lapse into out-of-print sta- chapters with thematic similarities, there are many in-
tus, or I would agree to produce a second edition. Look- stances where chapters in one section make points of
ing back on all the work, I suspect it might have been relevance to chapters in other sections. Nevertheless, a
wiser to say, "Let her lapse." But I didn't. During my de- sequence from Chapter 1 to Chapter 19 makes reason-
liberations, I thought about whether a new edition was able sense, and, if I used it in a graduate seminar, that's
worthwhile, whether further books on the topic were the sequence I would follow.
justified, and what my colleagues would say if I came I knew that growth in this field had been substan-
seeking chapter authors. I told Academic Press that a tial, but in finalizing the bibliography for this book, I
second edition was unrealistic, but that an entirely new realized just how great it had been. When Paul Ehrlich
book, that would visit some of the same topics, was a (1975) reviewed the population biology of reef fishes,
definite possibility. Then began the search for willing he did a thorough job in 36 pages and cited 313 ref-
contributors. erences going back to 1908, including a handful or
My intention from the beginning was to produce a two from prior to 1950. In Sale (1980), I reviewed
book that would speak to graduate students, to scien- the field in 54 pages, citing 318 references, nearly all
tists in the field, to reef managers and others interested of which were from the 1960s and 1970s. The Ecol-
in coral reefs, and to the wider ecological and scientific ogy of Fishes on Coral Reefs required 754 pages, of
community. I am confident that this book will do so, which 87 pages comprised a bibliography of about
and will open new doors that attract new people to be- 1690 citations, mostly from the 1970s and 1980s. The
come direct participants in this exciting field. The 30 present book contains over 2580 citations, of which
contributors (including myself) include 15 based in the more than 60% are from 1990 or later, while just
United States, 9 in Australia, two in Canada, two in 14% are from the 1970s or earlier. Further, the present
France, one in Panama, and one in Hong Kong. (Lest book is less comprehensive than the former, and whole
my American colleagues read this as a sign of their fields of ecology are omitted to keep the book to man-
preeminence, 11 of 19 chapters include authors with ageable size. There are a lot more people doing reef
significant Australian experience while just 5 have ex- fish ecology now than there were as little as 10 years
clusively American parentage. And, to keep my Aussie ago.
friends under controlmsome of us have left your shores, The other change in this field has been the growing
mates.) The chapters provide comprehensive coverage awareness by reef fish ecologists that our study animals
of the major fields of ecology of reef fishes currently are not only wonderful, but valuable, rare, and be-
being investigated, essential reviews in several cognate coming rarer. I hope that this book will encourage still
areas, and four chapters devoted to science of manage- more ecologists to explore reef fishes as model organ-
ment issues. As they arrived over the last 18 months, isms with which to ask important and fundamental eco-
and I had a chance to read them, their quality provided logical questions, and to this end, most chapters close
the spur to ensure I did the things I had to do to get with questions for the future. But I hope, even more,
the book to press. There is some excellent work here, that this book will encourage ecologists to use their sci-
and I thank each of the contributors for working hard ence to contribute to much more effective management
to produce a quality product, for putting up with my of our impact on reef fish and the other components
xiii
xiv ecaferP
of coral reef systems. There is good, intellectually stim- summer of 2001. Each provided the careful attention
ulating science that is desperately needed if we are to that allowed me to assemble a pooled bibliography with
manage these systems sustainably in the future. I want minimal mistakes, and they did the work cheerfully. I
somebody to write a new book on reef fish ecology thank Caroline Lekic who came to my aid at a critical
in 10 years and to be able to keep it in the present point as we compiled the index. Finally, I cannot ade-
tense. quately thank two special people, Donna and Darian,
I have already thanked the contributors. In putting who make my life worthwhile, while somehow under-
the final manuscript together, I was helped by two standing that I sometimes neglect them, only because I
undergraduates in turn: Nick Kamenos, who worked do love what I do.
in my lab through the fall of 2000, and Allison
Pratt, who worked there through the spring and Peter .F Sale
Introduction
E cology si a holistic science that seeks a broad understanding of the relationships
among organisms, their environment, and, increasingly, humans. Though the science
of ecology has its own specialties, ecologists must remain well versed in the methods,
the goals, and the knowledge accumulated in other fields. In the past decade or so, rapid
developments in instrumentation, in techniques and analytical approaches, and in general
knowledge in fields as remote as molecular biology and physical oceanography have made
keeping up with progress in these allied fields ever more difficult. Yet, more than ever,
the challenges facing ecologists can be answered only by using the knowledge gained by
workers in other fields, and by building collaborations that go beyond the boundaries of
ecology. Several of these other fields are discussed in later parts of this book; however,
this first part deliberately attempts to provoke awareness and generate new questions.
The five chapters that make up Section I all focus on ecological science. Yet each chapter
also looks at an edge of ecological exploration, and will help fill in gaps in knowledge of
some cognate fields. I believe that each chapter has the capacity to stimulate new questions
and new approaches by those who will do the future work in reef fish ecology.
In Chapter 1, Dave Bellwood and Peter Wainwright begin at the beginning, with a
review of the origins of reef fishes and reef fish faunas, reminding us of how advanced
reef fish assemblages are, and how relatively recently derived are those species that now
dominate coral reef systems. (That they refer to this 50-million-year-ago history as a long
one confirms they are, at heart, ecologists rather than paleontologists!)
Bellwood and Wainwright review this history in a way that facilitates understanding
of the biogeographic features of coral reef systems. Given that ecological studies are in-
creasingly being done on larger spatial scales, their discussion of how reef fish assemblages
differ from place to place si particularly helpful. Their question of whether reef fishes
have played an important role in facilitating the development of coral reefs si particularly
provocative.
Wainwright and Bellwood, in Chapter 2, shift from phylogeny to functional morphol-
ogy, developing a picture of the feeding ecology of reef fishes as driven by morphological
possibilities and constraints. For those readers brought up with classical ichthyology, this
chapter will be a refreshing update, but not a surprise. But too many students of ecology
now manage to bypass "old-fashioned" courses, and, for them, this chapter may open a
new door to appreciating that ecology si the result of interactions of real organisms that
have physical limitations and possibilities.
In Chapter 3, Howard Choat and Ross Robertson steer a more narrowly defined
ecological path, but set out a strong argument for radically revising the way reef fish
ecologists and fisheries biologists have approached demographic questions. They argue,
convincingly, that it si possible to age coral reef species, and demonstrate that the results
of so doing are going to cause some significant revisions in the "conventional wisdom"
concerning longevity and growth rates in these animals. Given that so much fisheries science
depends on knowledge of age structures and growth rates, their argument has importance
for management as well as for ecology.
