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About the Duke Center for Marine Conservation

The Duke Center for Marine Conservation was established to develop a na-
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reach on a wide variety of coastal and marine topics and issues, including ma-
rine fisheries, marine mammals and threatened and endangered species, coastal
development, water quality, and ocean policy and planning.
Marine Conservation Biology
Marine
Conservation
Biology
THE SCIENCE OF MAINTAINING
THE SEA’S BIODIVERSITY
Edited by Elliott A. Norse

and Larry B. Crowder
Foreword by Michael E. Soulé
Marine Conservation Biology Institute
WA S H I N G T O N • COVELO • LONDON
Copyright © 2005 Island Press
All rights reserved under International and Pan-American

Copyright Conventions. No part of this book may be re-
produced in any form or by any means without permis-
sion in writing from the publisher: Island Press, 1718
Connecticut Ave., Suite 300, NW, Washington, DC 20009.
ISLAND PRESS is a trademark of The Center for Resource

Economics.
COVER IMAGE : Steller’s sea cow (Hydrodamalis gigas), a

giantic North Pacific sirenian, in its kelp forest ecosystem
(extinct 1798). © 2005 Ray Troll, www.trollart.com
Seahorse © 2004 Jamie Kelley
The authors and editors have made good faith efforts to

secure permission to reprint the following figure, and
have been unsuccessful in doing so. If the copyright hold-
ers will contact the publisher, any necessary corrections
can be made to future printings.
Figure 12.2, page 200, from Sainsbury, J.C. (1996). Com-

mercial Fishing Methods: An Introduction to Vessels and
Gears. 3rd ed. Fishing New Books, Oxford (UK)
No copyright claim is made in the work of Gregory M.

Ruiz, an employee of the federal government.
Library of Congress Cataloging-in-Publication Data
Marine conservation biology : the science of maintaining

the sea’s biodiversity / edited by Elliot A. Norse and Larry
B. Crowder ; foreword by Michael E. Soulé
p. cm.
Includes bibliographical references and index.
ISBN 1-55963-661-0 (cloth : alk. paper)—
ISBN 1-55963-662-9 (pbk. : alk. paper)
1. Marine biological diversity conservation.
I. Norse, Elliot A. II. Crowder, Larry B. III. Title.
QH91.8.B6M37 2005
333.95'616—dc22 2004029346
British Cataloguing-in-Publication data available.
Printed on recycled, acid-free paper
Design by BookMatters, Berkeley
Manufactured in the United States of America

10 9 8 7 6 5 4 3 2 1
For the evolving Rachel Carsons and Aldo Leopolds, whose vision, understanding,
and action will make all the difference for our blue planet
Contents
FOREWORD 5. Extinction Risk in Marine Species

Michael E. Soulé . xi Ransom A. Myers and C. Andrea Ottensmeyer . 58
P R E FA C E
Elliott A. Norse and Larry B. Crowder . xvii 6. Behavioral Approaches to Marine Conservation
Julia K. Parrish . 80
ACKNOWLEDGMENTS
Elliott A. Norse . xxi
PA R T T W O . Threats to Marine Biological Diversity
Larry B. Crowder . xxv
Elliott A. Norse and Larry B. Crowder . 105
1. Why Marine Conservation Biology?

7. The Potential for Nutrient Overenrichment to
Diminish Marine Biodiversity
Nancy N. Rabalais . 109
2. Back to the Future in Marine Conservation
Larry B. Crowder . 19
8. The Magnitude and Consequences of Bioinvasions
in Marine Ecosystems: Implications for Conservation
PA R T 0 N E . Marine Populations: The Basics
Biology
Larry B. Crowder and Elliott A. Norse . 31
James T. Carlton and Gregory M. Ruiz . 123
3. The Life of the Sea: Implications of Marine

9. Diseases and the Conservation of Marine Biodiversity
Population Biology to Conservation Policy
Kiho Kim, Andy P. Dobson, Frances M.D. Gulland,
Stephen R. Palumbi and Dennis Hedgecock . 33
and C. Drew Harvell . 149
4. The Allee Effect in the Sea

10. Multiple Stressors in Marine Systems
Don R. Levitan and Tamara M. McGovern . 47
Denise L. Breitburg and Gerhardt F. Riedel . 167
ix
x Contents
PA R T T H R E E . The Greatest Threat: Fisheries 19. Metapopulation Structure and Marine Reserves
Larry B. Crowder and Elliott A. Norse . 183 Romuald N. Lipcius, Larry B. Crowder,
and Lance E. Morgan . 328
11. Global Fisheries and Marine Conservation:

PA R T F I V E . Human Dimensions
Is Coexistence Possible?
Dave Preikshot and Daniel Pauly . 185
20. Developing Rules to Manage Fisheries:

12. The Global Destruction of Bottom Habitats by
A Cross-Cultural Perspective
Mobile Fishing Gear
James M. Acheson . 351
Les Watling . 198
21. The Role of Legal Regimes in Marine Conservation

13. Effects of Fishing on Long-Lived Marine Organisms
Alison Rieser, Charlotte Gray Hudson,
Selina S. Heppell, Scott A. Heppell, Andrew J. Read,
and Stephen E. Roady . 362
and Larry B. Crowder . 211
22. Uncertainty in Marine Management

14. Evolutionary Impacts of Fishing on Target
Populations Louis W. Botsford and Ana M. Parma . 375
Richard Law and Kevin Stokes . 232

23. Recovering Populations and Restoring Ecosystems:
Restoration of Coral Reefs and Related Marine
15. Are Sustainable Fisheries Achievable?
Communities
Ray Hilborn . 247
Robert H. Richmond . 393
Place-Based Management of
PA R T F O U R .
Marine Ecosystems 24. Toward a Sea Ethic
Larry B. Crowder and Elliott A. Norse . 261 Dorinda G. Dallmeyer . 410
16. Marine Protected Areas and Biodiversity 25. Ending the Range Wars on the Last Frontier:
Conservation Zoning the Sea
Callum M. Roberts . 265 Elliott A. Norse . 422
17. Marine Reserve Function and Design for Fisheries A B O U T T H E E D I T O R S . 445

Management
C O N T R I B U T O R S . 447
Joshua Sladek Nowlis and Alan Friedlander . 280
I N D E X . 451
18. Place-Based Ecosystem Management in the

Open Ocean
Elliott A. Norse, Larry B. Crowder, Kristina Gjerde,
David Hyrenbach, Callum M. Roberts, Carl Safina,
and Michael E. Soulé . 302
Foreword
Conservation biology emerged as a recognized field of

Connections and Obstacles—
mission-oriented scholarship about a quarter-century
An Allegorical Fancy
ago when many terrestrial ecologists, geneticists, and
systematists were horrified by the gathering momen- By the year 2012 in California, where sea otters had
tum of a great planetary extinction event. At the been rendered morbid by a host of human-generated
same time, they recognized that responsible govern- insults—from pathogen-rich sewage effluents to en-
mental institutions and nongovernmental organiza- trapment in feral fishing gear—the three-dimensional
tions alike were largely ignorant of knowledge from kelp forests were clear-cut by predation-released sea
several fields—including systematics, biogeography, urchins and converted to near two-dimensional sea
evolution, genetics, and ecology—that might inform urchin barrens. The fish and invertebrate diversity was
effective responses to the crisis. What was needed, vastly reduced by this habitat simplification. Soon,
they saw, was an organized movement to enhance human beings were saddened that recreational and
communication between concerned scientists on the fishing opportunities had diminished. Finally, coastal
one hand and professional conservationists on the real estate values sank because the kelp forests no
other. longer shielded the fragile coastline from wave damage.
Since then, communication of scientific knowl- The reach of these events was continental in scale;
edge about biodiversity has improved, but clouds of indeed, a piece of eroding shore-front real estate was
psychological, ideological, and economic denial — owned by your mother-in-law in Cleveland, affecting
public and private—have generally obscured the ac- both her retirement options and your life style when
celerating, planetwide catastrophe and have impeded she moved in with you.
the necessary actions that might save much of the bi- Such stories illustrate that every marine perturba-
ological beauty and grandeur that took billions of tion, from the functional extirpation of evolving enti-
years to evolve. Some rays of light, however, shine ties like sea otters, whales, sea turtles, cod, or wild
through. This book is such a beam, and a declaration salmon, to the smothering sprawl of eutrophic dead
by a prestigious cadre of marine scientists that life in zones (Rabalais, Chapter 7) or a shift in the Gulf Stream
the oceans is as vulnerable as it is on the lands. I offer will change the forests and rivers around Colombo,
only a few, brief comments based on my interactions Oslo, and Moscow. We can also say that all climate is
with scientists and activists during the last 25 years. ultimately maritime; in other words, that every change
xi
xii Foreword
in the oceans, whether natural or anthropogenic, re- teractive and biggest individuals in the sea, particularly
flects back into the atmosphere, affecting all the land, large fishes, reptiles, birds, and mammals. Jeremy Jack-
sooner than later. Finally, we can say that every ma- son and others have documented many such conse-
terial constellation that emerges on the land, whether quential pathologies, including the effects occurring
a mountain, a mansion, or a microchip, will bow to when big, algae-eating fish are removed from coral
entropy and be swept into the seas. Such is the unity reefs. Discoveries like these exemplify an ongoing rev-
of the biosphere on the short time scale of centuries to olution in ecology: the salience of top-down, cascading
millennia. Saying all this another way: Earth is a sin- interactions, particularly those initiated by large her-
gularity of pulsating land masses, gyrating waters, bivores and predators. More and more we see that the
and a tossing blanket of gases, all of which are mas- consequences of absent top-down interactions are vor-
sively perturbed by billions of rampaging, large- tices of degradation and simplification. A problem is
brained primates. that critical interactions are virtually impossible to
What are the implications of such global interac- discover when long-term, spatially extensive research
tions for marine conservation biology? The emer- is eschewed and when scientists ignore history. One re-
gence of conservation biology in the late 1970s was a sult of such ignorance is the professional malfeasance
timely reaction by scientists to the obvious destruc- dubbed by Daniel Pauly as the “shifting baseline syn-
tion of ecosystems worldwide by pesticides, bulldoz- drome” (Crowder, Chapter 2).
ers, and chain saws. At the same time, most people A terrestrial example is illustrative. Wolves were ex-
viewed the oceans as nearly inexhaustible. Only a tirpated from Yellowstone National Park in the 1920s.
decade or two later, however, marine biologists were Among the long-term sequelae of this policy was the
stunned by the vulnerability of life in the oceans, and failure of aspen seedlings or suckers to reach the for-
the field of marine conservation biology was born. In est canopy during the following eight decades; an-
celebrating the maturation of marine conservation bi- other was the disappearance from the northern
ology, however, it is helpful to reflect on processes reaches of the park of a critical ecosystem in the arid
that transcend biogeographic realms and dissolve dis- West—beaver wetlands and the beavers themselves.
ciplinary barriers that artificially segregate thinkers. Similar landscape changes are occurring throughout
Knowledge of spatial and temporal connectivity that the Rocky Mountain region. The cause of these land-
help modulate both planetary anabolism and catabo- scape changes has been overbrowsing on aspen and
lism has exploded, but the public’s ignorance of this willows by an exploding population of wolf prey;
knowledge is an impediment to the deepening aware- namely, elk or moose. Since the return of the wolves
ness of planetary articulations and, hence, a major im- in 1995, however, aspen saplings are shooting up and
pediment to the conservation of life. The habit of willows are flourishing, particularly in areas where
thinking of oceans as remote but enduring, and land wolves abound. As expected, beavers are returning in
as familiar but fragile, is unsound. large numbers and restoring the critical wetland habi-
tat, thereby increasing local species diversity and rais-
ing water tables.
Loss of Interactions between Species
The ecological renaissance in Yellowstone, however
Contributing hugely to the planetwide crisis of biotic profound it is as a case study, is only the first step to-
dissipation is a failure to synthesize across temporal ward continental restoration. The wolf is still confined
and spatial scales. This is particularly apparent with re- to only 5 percent of its original US range. The restora-
spect to the deliberate extermination of top predators tion of the wolf and other highly interactive species
on the land and the overkilling of the most highly in- must be replicated everywhere, not only in flagship lo-
Foreword xiii
cations such as Yellowstone. The same principle ap- ogy/management, and marine biology. Graduate stu-
plies to the oceans. dents rarely are taught even the fundamentals of other
The symbolic persistence of interactive species such sister disciplines; I doubt that many oceanographers or
as otters, wolves, whales, billfishes, groupers, and fisheries managers have read Robert Paine’s work. Sec-
sharks in just a few places is a fool’s solace. At best, it ond, grants and promotions demand the frequent
is an expedient means or a first step toward recovery, publication of rigorous, hypothesis-driven, experi-
and it should never be seen as an end in itself. Con- mental results; this discourages research on long-term
gratulating ourselves for the local persistence (or the and spatially extensive questions, including the top-
presence of juveniles only) of a once widespread, in- down effects of long-lived species, on which highly
teractive species is like claiming that most cities don’t controlled experiments are difficult if not impossible.
need emotionally complex, adult music because Third, scientific advance is trammeled by adherence to
Philadelphia, Tokyo, and Vienna already have sym- a particularly nonadaptive trait of modern ecology,
phony orchestras. the stranglehold of statistical methods developed for
The point is that ecological function and biological agronomic science that require a high degree of repli-
diversity in the sea and on the land have always de- cation, controlled environments, and the testing of
pended on large, potent animals, and that their deci- narrowly constructed hypotheses. This strange, hege-
mation during the last two or three centuries repre- monic paradigm should be applied to only a few as-
sents a decapitation of ecosystems. Globally, the pects of marine science. Fourth, conservatism is exac-
general killing of large interactive species, whether in erbated by the slow turnover of senior scientists
forests, reef systems, steppes, or pelagic regions, will be relative to the fast turnover of ideas, assuming that
seen as a decisive episode of the global biotic extinc- most intellectuals rarely discard ideas imprinted on
tion catastrophe. Those who relate to such animals as them during their apprenticeships. Finally, politics
mere tonnage, or as ecologically irrelevant epiphe- militates against the repatriation over large areas of
nomena of bottom-up driven food webs, will one day predator-driven, top-down dynamics because the gov-
be accused of a lethal, ecological myopia. ernment agencies that administer conservation laws
Nearly all dualisms in ecology are eventually such as the Endangered Species Act are strongly influ-
proven to be the two arms of an integrated theory. enced by conservative politicians hostile to predators.
Top-down dynamics complement, but do not com-
pete for, the gold medal with bottom-up, production-
Certification and Campaigning
driven dynamics. It is a tragedy that many scientists
are reluctant to embrace higher-order, complex theo- Market-based goals and the religion of economic
ries of ecological determination, preferring, like cow- growth currently dominate nearly every aspect of
boys, the single-barreled smoking gun. There are at modern society. An unfortunate corollary of this
least five interrelated factors contributing to this fail- reigning worldview is the belief in unrestrained com-
ure of imagination. One is the narrowness of graduate petition, which requires short-term profits and un-
education; the second is the academic reward system; sustainable, often unregulated, use of resources. It be-
the third is fashion in statistics; the fourth is the hooves marine conservationists, therefore, to make
longevity of politically powerful personalities and an better use of market-based defenses. A terrestrial ex-
inability of most human beings to grow and expand ample is “forest certification.” This is a labeling tool by
intellectually; the fifth is politics. which consumers can determine if the practices of the
First, the marine sciences are balkanized into iso- wood products industry are minimally destructive
lated disciplines, such as oceanography, fisheries biol- and have any redeeming environmental or social
xiv Foreword
value. It should be noted, however, that antedating teracting tsunami of overpopulation, perverse eco-
forest certification was the “dolphin-safe tuna” cam- nomic incentives, technomania, globalized corrup-
paign. Such campaigns are based on the sociological tion, and market mechanisms such as free trade that
premise that the public generally resonates better to currently preclude the competetiveness of environ-
arguments that appeal to their compassion and their mentally sustainable practices. In addition, the repa-
love of life than to promises for new cancer cures. At triation of large, pelagic predators at ecologically ef-
least it seems clear that the discovery of life-extending fective densities requires oceanwide protection and
pharmaceuticals in rain forest organisms has not management. Moreover, anything less bold and dra-
slowed the rate of forest destruction. matic than a whole-of-seas vision will be ignored by a
It bodes well that this technique is being applied dazed and laconic public, given the titanic levels of
more broadly by ocean advocates. Still, an insignifi- distraction and corporate/government propaganda.
cant proportion of shoppers and diners are aware that Finally, the scale of solutions must match the scales of
the consumption of farmed salmon and the capture of the underlying, dissipative forces—including open-
long-lived, slow-reproducing fish are unsustainable. ocean fisheries, industrially produced climate change,
The good news is that science-based certification pro- the transport across oceans of invasive marine species,
grams are being broadened to include prawns, bill- and chemical pollution. Whole-of-oceans strategies,
fishes, sharks, and old-growth bottom fish. Organiza- however, complement but do not substitute for local
tions like the Blue Ocean Institute and the Sustainable conservation efforts, including the protection of
Seafood Alliance deserve our support and money. coastal and reef systems.
Whole-of-oceans visions must be rigorous as well as
inspiring. And “rigor” means the erection of a seawall
Whole-of-Oceans Conservation
between science and politics—not allowing the jet-
When conservation biologists first called for conti- sam of human greed and desire to spill over and
nental-scale conservation projects based on the criti- smother the murmurings of ocean life. A mistake that
cal importance of regional- and continental-scale many scientists and advocates make when dealing
flows and processes, most conservationists were be- with issues like the sufficiency of protected habitat on
mused, thinking that such an audacious scheme land and sea is to be overly conciliatory and to do the
would never be taken seriously. They were wrong. developer’s/exploiter’s work for him by prematurely
Plans are nearly completed for a continuous network factoring in economic and political considerations
of protected lands along the spine of the North Amer- before the biological needs are known. This might be
ican Continent from the Sierra Madre Occidental to called “the neurotic need to appear reasonable in the
the Yukon and Brooks Range in Alaska. Similar proj- eyes of the exploiter,” the political analogue of psy-
ects are being developed in central and eastern Eu- chological codependency. Instead of clearly stating
rope, in southern Africa, and in Australia, to name a what is necessary to protect all living beings, includ-
few. Hundreds of governmental and nongovernmen- ing extensive processes in the sea, we often censor
tal organizations are involved. Such is the power of a ourselves, indulging in the calculus of naive reckon-
compelling, science-based, vision. ings of social, economic, and political “realities.” This
The sea still lacks such a vision, though similar is self-defeating. Policy makers need to know what is
bold, science-based ideas are being proposed for the biologically necessary for long-term, geographically
coastal seas and even for open oceans (Norse et al. extensive restoration of ocean life, not what scientists
Chapter 18). Scientifically rigorous whole-of-oceans believe to be politically feasible.
visions are the only practical way to respond to the in- Candor and boldness have power. A vision based
Foreword xv
on both compassion for all life and solid science vides the foundation for such a declaration of inter-
evokes respect and wonder and is far more effective dependence of all life, oceanic and terrestrial, nonhu-
than the mincing, qualified, soulless recommenda- man and human.
tions of committees and expert panels. True, there are
times (like these) when it appears that nothing will
Acknowledgments
stop the industrial and population juggernauts that
are destroying life on this planet, not to mention the I thank Jim Estes, John Terborgh, Elliott Norse, Larry
dignity and diversity of human cultures; such times Crowder, and Arty Wolfson for their helpful comments.
require patience, humor, and solidarity between con-
servationists and humanitarians. This volume pro- Michael E. Soulé
Preface: A New Science for a New Century
Just two human life spans ago, North America’s skies Mars, the only place in the universe where we know
thundered with the wingbeats of 5 billion passenger that life exists has rapidly been losing its distinguishing
pigeons (Ectopistes migratorius), its prairies shook with characteristic, its biological diversity, the diversity of
the hoofbeats of 30 million American bison (Bison genes, species, and ecosystems (Heywood 1995; Norse
americanus). Now these are fading memories: the pi- and McManus 1980; Norse et al. 1986; Office of Tech-
geons are extinct; the buffalo, having edged up from nology Assessment 1987; Wilson 1988). As the perva-
a population minimum of 600, live mainly as small siveness of this worldwide loss became apparent, peo-
herds in fenced enclosures. Our species ate them into ple from a diversity of sciences coalesced to study ways
history. But in 1883, even as American bison and pas- to prevent this loss in a new multidisciplinary science
senger pigeons plummeted toward extinction, fishes named conservation biology (Soulé and Wilcox 1980).
seemed so abundant that eminent British biologist By applying perspectives from systematics, ecology,
Thomas Huxley (1883) declared, “I believe that the biogeography, genetics, evolutionary biology, physiol-
cod fishery, the herring fishery, the pilchard fishery, ogy, behavior, wildlife biology, forestry, horticulture,
the mackerel fishery, and probably all the great sea- veterinary medicine, epidemiology, ecotoxicology,
fisheries are inexhaustible; that is to say, nothing we archeology, history, anthropology, sociology, econom-
can do seriously affects the number of fish.” ics, political science, law, and ethics, this new discipline
Now Jackson et al. (2001), Pauly and Maclean built the intellectual foundation for conserving biodi-
(2002), Roman and Palumbi (2003), and a growing versity. Most important is the realization that under-
list of others show that Huxley was wrong even then, standing humans—who are both the cause and the
that humankind has been systematically eating the victims of biodiversity loss—is as integral to conserva-
sea’s wildlife into history. In less than a lifetime, mation biology as is understanding biology.
rine fisheries have reduced populations of large preda- Yet, during its formative decades, conservation bi-
tory fishes by 90 percent (Myers and Worm 2003). It ology had a silent antecedent: nonmarine. None of the
seems that those entrusted with protecting marine chapters in Soulé and Wilcox’s (1980) landmark first
life never learned the lessons of the passenger pigeons conservation biology book, and only one ( Johannes
and buffalo. and Hatcher 1986) in Soulé’s second book, concern bio-
Even as humankind spends billions of dollars in diversity loss in the sea. In the sole marine chapter in
the hope of detecting the faintest echoes of life on Wilson’s biodiversity book, Ray (1988) highlights in-
xvii
xviii Preface
attention to the sea with the telling example of a (SCB), a nonprofit advocacy organization, Marine Con-
conservation-oriented world biome map that simply servation Biology Institute (MCBI) was founded with
leaves the oceans blank. Conservation biologists’ in- the goal of advancing the new science of marine con-
difference to marine species and ecosystems was de- servation biology. MCBI’s preliminary objective was or-
cried in an early issue of Conservation Biology (Kauf- ganizing the first Symposium on Marine Conservation Bi-
man 1988) and quantified by Irish and Norse (1996), ology at SCB’s 1997 annual meeting in Victoria, British
who found that terrestrial papers outnumbered ma- Columbia (Canada). Its 44 marine paper sessions were
rine papers 13:1 during the journal’s first nine years. a quantum advance from the total of 2 marine paper
In a more detailed study, Kochin and Levin (2003) sessions in the 10 previous SCB annual meetings. In
quantified our impression that conservation biology 1998, 1,605 conservation biologists and marine scien-
has largely overlooked the largest of the Earth’s bio- tists joined in signing an unprecedented statement
logical realms. called Troubled Waters: A Call for Action (Box 1.1),
Moreover, while the sea was terra incognita for which urges citizens and governments worldwide to
conservation biology, biodiversity conservation was “provide sufficient resources to encourage natural and
largely overlooked by marine scientists. Marine sci- social scientists to undertake marine conservation bi-
ences have tended to treat the diversity of marine life ology research needed to protect, restore and sustain-
as either (1) largely irrelevant (oceanography), (2) im- ably use life in the sea.” The Second Symposium on Ma-
portant only because it is fascinating (marine biology), rine Conservation Biology (San Francisco, California,
or (3) important only to the extent that it is edible USA) took place in 2001, the year in which SCB estab-
(fisheries biology). Kochin and Levin (2003) show lished a Marine Section. The 2004 annual meeting of
that marine science papers that examine conservation the American Association for the Advancement of Sci-
are scarce. Similarly, a book on marine biodiversity by ence (AAAS) in Seattle, Washington (USA) had more
Ormond et al. (1997) is so focused on distribution pat- marine conservation sessions than any previous AAAS
terns of species that it largely overlooks their conser- meeting. Marine biodiversity loss has become an
vation status and trends. Those who study conserva- agenda item within the scientific community.
tion have—until very recently—been a tiny minority The messages that scientists generate have drawn
in the marine sciences. Conservation biologists have the attention of the public and decision makers. In the
overlooked the sea while marine scientists have over- United States, public opinion polls for SeaWeb by the
looked conservation, which seems, from our perspec- Mellman Group (1999) showed that 87 percent of
tive, a rather large gap for a blue planet. Americans consider the condition of the ocean very
There is compelling reason to fill this gap. By the important or somewhat important to them personally,
end of the 20th century (Butman and Carlton 1995; and 92 percent feel the responsibility to preserve the
Norse 1993; Thorne-Miller and Catena 1991), it be- ocean and restrict human activities necessary to do so.
came apparent that the sea—including estuaries, semi- In 1999, a bipartisan group of US Congressmembers
enclosed seas, coastal waters, and the open oceans—is founded the House Oceans Caucus to frame new fed-
rapidly losing its biological diversity as the human eral laws concerning marine issues. In 2000, President
population increases, technologies become more pow- Clinton issued Executive Order 13158, calling for fed-
erful, and people seek the last places with exploitable eral agencies to cooperate in establishing a national
biomass to replace those denuded by overexploitation. system of marine protected areas. The country with
Now, at last, attention to marine biodiversity loss is the most marine scientists—the United States—has
increasing. In 1996, 18 years after Michael Soulé’s sem- lagged behind Australia, a country with only 5 percent
inal Conservation Biology Conference, 10 years after the of the US GDP, in producing comprehensive national
founding of the Society for Conservation Biology ocean policies. Now, however, a third of a century
Preface xix
after the last comprehensive report on US ocean pol- tions, are so underrepresented. We hope, in years to
icy, the blue-ribbon Pew Oceans Commission (2003) come, that both the utility of this book and our errors
issued its visionary report, followed by the recom- and omissions will encourage others to do better. But
mendations of the US Commission on Ocean Policy for now, to help stop and reverse the loss of the sea’s
(2004). Concern about declining marine biodiversity biodiversity, to ensure that bountiful seas do not be-
loss has spread from the scientific community to the come another fading memory, to empower the next
decision-making community. generation of leaders in marine conservation, we offer
Coalescing a new science requires bringing to- this book.
gether people and their ideas. Symposia on marine
conservation biology are one way of doing this. This Elliott A. Norse and Larry B. Crowder
book is another. We have assembled it because we be-
lieve that humans can be wise enough to see the con-
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xx Preface
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Acknowledgments
If it takes a village to raise a child, then surely it takes ology teacher in high school; Priscilla Pollister and
a world to raise a book. In examining a topic that is Elizabeth Worley, my mentors and biology professors
fascinating, painful, and rewarding, I have been in college; John Garth, my major professor in graduate
buoyed by a globe-spanning community of old school; Bill Beller, who introduced me to the realities of
friends and new friends who gave generously of government in my first job in marine conservation;
themselves so that this book could happen. First Malcolm Baldwin, the conservation visionary who
among them is my coeditor Larry Crowder, whose tasked me with writing the chapter that first defined
prodigious knowledge, expansive vision, penetrating the idea of maintaining biological diversity; and Roger
insight, wisdom, humor, enthusiasm, energy, self- McManus, my coauthor in that endeavor.
lessness, doggedness, and patience have been a lesson I thank my longstanding friends, mentors, and
in how to be a better scientist and person. He made MCBI Board Members: Gary Fields, Michael Soulé, Jim
this effort both an adventure and a pleasure. He is the Carlton, Irene Norse, Alison Rieser, John Twiss, Bob
best of the best. Kerr, Jim Greenwood, and Larry Crowder, who shared
Of course, no idea is really new; they all have their wisdom (even when I resisted hearing it!) and that
wellsprings. My motivation to foster the growth of ultimate limiting factor, their time. I thank the talented
marine conservation biology began with my family. and dedicated staff of Marine Conservation Biology In-
My uncle, Elliott Albert, so deeply loved nature that stitute, who encouraged me to work on the book when
he might well have devoted his life to conservation I would otherwise have been helping them with their
had he not sacrificed it as an 18-year-old Ranger on a scientific and conservation tasks or working to raise the
beach in Italy in 1944. He bequeathed me his name money to pay our salaries. I particularly thank MCBI
and his unfinished mission. My mother, Harriett Sig- Chief Scientist Lance Morgan, who never once hesi-
man, and my father, Larry Norse, fed me love, envi- tated to pick up the mess I made as this book proceeded
ronmental ethics, and fascinating facts about wildlife in fits and starts, as well as Bill Chandler, Fan Tsao, Sara
along with my ABCs. Although neither went to col- Maxwell, Hannah Gillelan, Mary Karr, Julia Christ-
lege, they encouraged me to learn all I could to help man, John Guinotte, Katy Balatero, Amy Mathews-
make the world a better place. They entrusted my Amos, Jocelyn Garovoy, Peter Etnoyer, Aaron Tinker,
training to people who shared belief in what I might and Caroline Gibson, whose fingerprints are thickly
become. These include Norman Scovronick, my zo- scattered throughout the pages.
xxi
xxii Acknowledgments
This book benefited enormously from my fellow Moore Family Foundation, Rockefeller Brothers Fund,
Pew Fellows in Marine Conservation and many others Russell Family Foundation, Sandler Family Supporting
in the growing community of senior and student ma- Foundation, Sun Hill Foundation, Surdna Foundation,
rine conservation biologists from Townsville to Mi- William H. and Mattie Wattis Harris Foundation, Vidda
lano. They are too numerous to thank individually, Foundation, Weinstein Family Charitable Foundation,
yet thank them I must, for they generated many of the and another special friend and funder who requests
insights that I now call my own. anonymity.
I thank Ann Lin and Linda Lyshall, whose assis- The richness of this book comes to us courtesy of
tance under trying circumstances saved the readers the authors (see Table of Contents), including many of
from errors, inconsistencies, and awkward phrases, the brightest stars in our field, who integrated lifetimes
and to Diane Ersepke and David Peattie for their deft of insights while enduring my repeated requests for
copyediting and organizational skills. Few people are quick action and, too often, my slow responses. The re-
more deserving of gratitude than Island Press’s Barbara sults—their chapters—speak with astounding power
Youngblood, Barbara Dean, Dan Sayre, Todd Baldwin, and eloquence. Other people do their heavy lifting be-
and Chuck Savitt, who waited so long for this book. I hind the scenes, performing the painstaking, anony-
hope that the result merited their patient guidance. mous task of critically reviewing chapters, and I espe-
And knowing that many people do judge books by cially thank Peter Auster, Felicia Coleman, Paul Epstein,
their covers, I thank Ray Troll, the Charles R. Knight Jocelyn Garovoy, Michael Hellberg, Ray Hilborn,
of our era and a champion of the beautiful and bizarre Graeme Kelleher, Jim Kitchell, Marc Miller, Lance Mor-
life in oceans present and past, for his remarkable gan, Jack Musick, John Ogden, Gail Osherenko, Hans
cover art commemorating Steller’s sea cow in its North Paerl, Pete Peterson, Bob Richmond, Dan Rubenstein,
Pacific ecosystem, and Jamie Kelley for her lovely sea- Dan Simberloff, Bob Steneck, Rob Stevenson, Gordon
horse drawing. Thayer, and Rob Wilder for doing so. Life is music; I
Funding for this book has come from people who thank the late Stan Rogers, who taught me how much
give to make the world a better place. I am deeply it hurts fishermen when they lose their way of life, and
grateful to the Pew Charitable Trusts, which had the Paul Peña, whose fusion of Tuvan throat-singing and
faith and foresight to fund MCBI to hold the first (Vic- gutbucket scratchy blues in Genghis Blues kept me
toria, British Columbia, 1997) and second (San Fran- going until the book was done. This book could not
cisco, California, 2001) Symposia on Marine Conservation have been completed without the staff of Friday Har-
Biology. These were thrilling meetings of leading bor Laboratories of the University of Washington, who
thinkers and eager learners. I thank those who trusted welcomed me to use their library and the beautiful fa-
us enough to gave MCBI that rarest of gifts; namely, cilities of the Helen Riaboff Whiteley Center to read,
unrestricted support, which my staff and I used to rethink, write, and edit away from ringing telephones.
search, write, and edit this book during the years be- In contemplating what it takes to be a marine con-
tween its conception in 1996 and its birth in 2005. servation biologist in a world where better knowledge
These wonderful people and foundations include Mark and funding are needed, the odds of winning are
and Sharon Bloome, Jennifer and Ted Stanley, Bertram daunting and too few people care enough, I might
Cohn, Anne Rowland, Sally Brown, Ben Hammett, seem a traitor to my generation for not agreeing that
the Bay Foundation, Bullitt Foundation, Curtis and “All you need is love.” If you want to save the oceans,
Edith Munson Foundation, David and Lucile Packard you need intellect, physical endurance, and an un-
Foundation, Educational Foundation of America, Ed- shakable belief in something much, much larger than
wards Mother Earth Foundation, Geraldine R. Dodge yourself. But you definitely also need love, of which
Foundation, Henry Foundation, Marisla Foundation, there are many sources.
Acknowledgments xxiii
One ongoing source of love in my life stands out. I despaired that my tasks were just too much to han-
My wife, Irene Norse, opened our home to a stream of dle, it was Irene who urged me to finish the book so I
jetlagged, hungry, fascinating, and impassioned ma- could fulfill some of my need to “pay it forward.”
rine conservation biologists; accepted the economic Last and most of all, I thank all of the people—es-
and emotional risk that I might fail in starting a new pecially the young people all around the world—who
organization dedicated to building a new science; put in decades to come will apply their minds, bodies,
up with too many trips that separated us; gave me the hearts, and spirits to the uniquely noble challenge of
time to close my office door to read, think, write, and saving life in the sea and elsewhere on our planet.
edit; served as my trusted counsel and critic; and
taught me by example that goodness is something Elliott Norse
you do through acts small and large every day. When Redmond, Washington
Acknowledgments
First, I must thank my friend and fellow editor, Elliott onates in my mind. I thank him and my mother, Jean
Norse, for 25 years of dialogue and debate. Elliott had Crowder, for encouraging me to find my gifts and to
begun this book before I became involved, but he wel- prepare myself for a career built on passion and com-
comed my input and allowed me to make it mine as mitment. I grew up in central California, a land where
well as his. This book has undergone a long gestation water was (and is) in short supply, but I seldom re-
driven by the rapid developments in the field and the turned from my “adventures” without being wet and
pursuit of these developments by the authors and ed- muddy. Camping vacations to the Sierras and Califor-
itors alike. Elliott and I have engaged in countless nia coast further fed my interests in the outdoors.
meetings and phone calls, and I appreciate his hospi- I began the path to scientist with the desire to be a
tality in Redmond, where he graciously hosted me on teacher. I had a long list of teacher role models, some of
a number of occasions. Thanks to Elliott and to his whom were inspirational and others who taught lessons
lovely wife, Irene! devoid of passion, interest, or excitement. “Stubby”
It is difficult to fathom whom to acknowledge for McKaye taught the first challenging biology course I en-
a lifetime of experiences that led to this book. I think countered; he was the first of many field naturalists and
the foremost credit belongs to my father, Earl Crow- environmentalists I encountered at key stages in my ca-
der, an Iowa farm boy who, as a teenager, blew west reer. At California State University – Fresno, Richard
to California during the dust bowl in the 1930s. He Haas saved me from a passionless career in engineering
endured hardship and displaced opportunities, but and pulled me toward my first love, which was field bi-
maintained a strong “can do” attitude. Before he ology. Bert Tribbey became a real mentor and fueled my
could complete high school, Pearl Harbor intervened passion for aquatic ecology. His no nonsense approach
and like many in his generation, he went to sea as an to field biology and his demands for excellence were
enlisted man at the age of 20. His bailing wire and nothing less than inspirational. He became more than
binder twine mechanical abilities were refined as a a professor—he was a mentor and friend and is still a
machinist mate and he rose to Chief over five years of model for interactions with students. My Ph.D. advisor
cruising, fighting, and swimming (unexpectedly) in at Michigan State University, Bill Cooper, combined
the world’s oceans—an unusual experience for some- passion and energy with a broad perspective on applied
one so strongly linked to the land. His encourage- ecology and environmental management. John Mag-
ment to be independent and to work hard still res- nuson and Jim Kitchell at the University of Wisconsin
xxv
xxvi Acknowledgments
hosted my postdoctoral years, some of the most exhil- I thank the following agencies and foundations for
arating of my career. All these teachers and mentors de- their support, particularly their recent support for crit-
serve my thanks. ical research in marine conservation: the Environ-
My wife Judy has been a steadfast supporter since mental Protection Agency, the National Science Foun-
we first met 35 years ago. She and my three children, dation, the National Oceanic and Atmospheric
Emily, Sean, and Elias, have shared many of my expe- Administration, the Office of Naval Research, the Na-
riences in the field and have endured periods of sepa- tional Oceanic Partnership Program, the Great Lakes
ration as I pursued research and travel opportunities. Fisheries Commission, the Pew Charitable Trusts, the
Judy is a writer and has always pressed me hard to Alfred P. Sloan Foundation’s Census of Marine Life pro-
communicate my science clearly and effectively to a gram, the Oak Foundation, the Gordon and Betty
wide variety of people. My family has provided love Moore Foundation, and individual contributors like
and support even when I didn’t deserve it—for this I Jim Sandler and Jeff Gendell. I thank these and all oth-
am most grateful. ers who have supported my research and teaching en-
Over the years I have been blessed with opportu- deavors over the years. This book was completed dur-
nity to interact with outstanding people, including ing my sabbatical, which was supported by a Center
undergraduates, graduate students, postdoctoral asso- Fellowship at the National Center for Ecological Analy-
ciates, and research collaborators. They are too many sis and Synthesis, by the William R. and Lenore Mote
to mention here and I resist mentioning them by Eminent Scholar Chair at Florida State University and
name lest I inadvertently slight any of them. Gradu- Mote Marine Laboratory, and by the Nicholas School
ate students, in particular, are the lifeblood of many of the Environment and Earth Sciences.
academics, including me. Young, energetic minds con- Finally, I thank the authors of the chapters printed
stantly challenge us to rethink our understanding of here. As individuals, they are the foundation of the
the work and to refine our methods to temper that un- field of marine conservation. Although none of the in-
derstanding. One administrator, who shall remain dividuals writing here were trained in marine conser-
nameless, once suggested that the university could vation per se, they earned their ranks in the trenches
save money by funding fewer graduate students—to and on the rough seas of this evolving field—indeed
which I replied, “Who do you think does all the they are defining it. Today’s students and postdoctoral
teaching and research around here!” Teaching is a real associates are the first generation of marine conserva-
privilege that too few faculty seem to really enjoy—I tionists, and it is for them that this book has been
am grateful for the opportunities I have had to inter- written and edited. I hope they enjoy it and find it
act with thousands of students. Recent support from useful. In a field evolving as rapidly as marine con-
the Educational Foundation of America, the Panaphil servation, a textbook is a moving target. Still, I hope
Foundation, the Lumpkin Foundation, the Oak Foun- the foundational effort of the authors and editors set
dation, and the Munson Foundation among others al- the stage for continued exciting developments in re-
lowed the creation of the Global Fellows in Marine search, education, and policy in marine conservation.
Conservation program at Duke University Marine Lab.
This program broadened our interactions and capac- Larry B. Crowder
ity building to a global scale. Bahia Magdalena
Research runs on both financial and moral support. Baja California Sur
1 Why Marine
Conservation Biology?
Elliott A. Norse and Larry B. Crowder
Nature provides a free lunch, but only if we control our appetites.

WILLIAM D. RUCKELSHAUS,
member of the US Commission on Ocean Policy
For many people old enough to remember, 1962 was son, who had written three best-selling books about
the year of the Cuban Missile Crisis, when the world the wonders of the sea as well as Silent Spring, said al-
came very close to nuclear self-immolation. The mis- most nothing in these books about human impacts on
siles and bombers were ready to launch, but—just the sea. And despite the Santa Barbara oil spill’s role in
barely—people in positions of power made difficult mobilizing the public, decision makers, and scientists,
choices, and sanity prevailed. That same year, marine environmental consciousness focused mainly on land
biologist Rachel Carson alerted the world to another and freshwaters. Having been jolted into awareness by
crisis that we humans had made for ourselves. Before a marine biologist and mobilized by an offshore oil
then, humankind had largely overlooked our impact spill, the environmental movement nonetheless
on our environment. But Carson’s (1962) best-selling shifted its attention away from the sea.
Silent Spring demonstrated that “progress” (in this Of course, our species evolved on land, and we
case, from new synthetic chemical pesticides) is tend to focus on what we can readily see. Although
threatening the integrity of our natural world. In our Paleozoic ancestors were marine animals, they left
1969, widely televised footage of dying seabirds from the sea hundreds of millions of years ago, and hu-
an oil well blowout in California’s Santa Barbara mans’ physiological mechanisms and senses are ill-
Channel quickly catalyzed a broad-based environ- suited for acquiring knowledge beneath the sea’s
mental movement that led to passage of a flood of wavy, mirrored surface. This profoundly affects ma-
new and strengthened environmental laws in the rine conservation because, on land, people can see at
United States and beyond. A decade later, Lovejoy least some of the consequences of our activities on
(1980), Myers (1979), and Norse and McManus (1980) ecosystems and species. Such understanding forms a
showed that our planet is losing its biological diver- basis for social, economic, and legal processes that can
sity, and Soulé and Wilcox (1980) called upon the protect biodiversity. But judging the integrity of ma-
world’s scientists to join forces to work to stop the ac- rine ecosystems and their species is far more difficult
celerating erosion of life on Earth. These were wake-up because most people never go below the sea surface
calls that society could no longer afford to ignore. Yet, and cannot gauge the health of a marine ecosystem.
ironically, the visionary and courageous Rachel Car- Rather, the general public and key decision makers
1
2 Marine Conservation Biology
(including legislators, agency officials, industrialists, years, since a giant meteorite smashed into Earth,
funders, and environmental advocates) depend on causing the extinction of dinosaurs, mosasaurs, am-
those with information about what occurs beneath monites, and countless other species (Ward 1994).
the surface, especially fishermen, offshore oil drillers, Moreover, this time it is not a mindless mass of rock
and marine scientists. And because people who extract that threatens the sea’s biodiversity. It is our species.
commodities from the sea have strong economic in- Marine conservation biologists do not need to
centive to minimize public concerns about marine plumb the scholarly literature to frame the challenge
biodiversity loss, marine scientists are by far the most we face; two quotations accessible to a broad audience
credible providers of information. suffice. In the 2002 movie Spider-Man, Uncle Ben
It is difficult even for scientists to appreciate how teaches Peter Parker, “With great power comes great
the sea has changed. On that Pleistocene day when responsibility.” Armed with the power either to de-
our ancestors first stood on an African shore and stroy or to protect, recover, and sustainably use ma-
looked seaward, they must have been stunned by the rine biodiversity, people can choose, just as people in
wealth of marine life they saw. Were the mudflats 1962 chose not to make nuclear war. The other quo-
paved with mollusks? Did the sea surface boil with tation is from a 1968 speech by American black radi-
fishes? Could dugongs and green sea turtles have cal Eldridge Cleaver: “You’re either part of the solution
filled the shallows like wildebeests and zebras on the or you’re part of the problem.” Marine scientists’
plains? We cannot know for certain, but scientific in- unique combination of knowledge and credibility cre-
formation increasingly coming to light from around ates for us a unique niche in a world desperately in
the world suggests that the sea was home to an as- need of answers. Some of us might have been part of
tounding diversity and abundance of life as recently as the problem in the past. But now the question is, Are
hundreds, even tens of years ago (Crowder, Chapter we up to the challenge of being part of the solution?
2). It was only in the 1990s (Butman and Carlton Are we ready to tackle the biggest question of all;
1995; Norse 1993; Thorne-Miller and Catena 1991) namely, How can humankind live on Earth without
that scientists assembled compelling information ruining its living systems, including the largest com-
showing that biological diversity in the sea is imper- ponent of the biosphere, the sea?
iled worldwide. And not until the International Year The marine conservation challenge shares most as-
of the Ocean did 1,605 the world’s scientists join pects of the terrestrial conservation challenge but also
forces to publicly voice their concern about marine has some distinctive features. There are many impor-
biodiversity loss in a statement called Troubled Waters: tant physicochemical and biological differences, but
A Call for Action (MCBI 1998) (Box 1.1). these pale in comparison to the human dimensions,
The land and freshwaters are anything but safe, how people treat estuaries, coastal waters, and the
and no knowledgeable person would suggest that we open oceans. Even more than poverty, affluence, tech-
can afford to reduce our commitment to them, but it nology, and greed, it is ignorance and indifference
is also time to focus much more attention on the sea. that are the enemies of marine biodiversity. Our not
The sea’s vital signs are disquieting: most everywhere knowing the sea, not living in it, and not having a
scientists look—in tropical and polar waters, urban es- sense of responsibility for it have led to a “frontier
tuaries and remote oceans, the sunlit epipelagic zone mentality” that has governed our social contract with
and seamounts in the black depths—we are seeing the sea. Signs that the end of the frontier is rapidly
once-abundant species disappearing, noxious species approaching indicate the need for a new system based
proliferating, ecosystem functions changing, and fish- on the idea that marine ecosystems are heterogeneous
eries collapsing. And although nature is always chang- and have many legitimate human interests. The last
ing, these changes are without precedent in 65 million chapter in this book (Norse, Chapter 25) offers ocean
B O X 1.1. Troubled Waters: A Call for Action
We, the undersigned marine scientists and conservation to diseases or undergoing mass bleaching in many places.
biologists, call upon the world’s citizens and governments There is no doubt that the sea’s biological diversity and
to recognize that the living sea is in trouble and to take integrity are in trouble.
decisive action. We must act quickly to stop further To reverse this trend and avert even more widespread
severe, irreversible damage to the sea’s biological diversity harm to marine species and ecosystems, we urge citizens
and integrity. and governments worldwide to take the following five
Marine ecosystems are home to many phyla that live steps:
nowhere else. As vital components of our planet’s life sup-
1. Identify and provide effective protection to all popula-
port systems, they protect shorelines from flooding, break
tions of marine species that are significantly depleted
down wastes, moderate climate and maintain a breath-
or declining, take all measures necessary to allow their
able atmosphere. Marine species provide a livelihood for
recovery, minimize bycatch, end all subsidies that
millions of people, food, medicines, raw materials and
encourage overfishing and ensure that use of marine
recreation for billions, and are intrinsically important.
species is sustainable in perpetuity.
Life in the world’s estuaries, coastal waters, enclosed
seas and oceans is increasingly threatened by: (1) over- 2. Increase the number and effectiveness of marine pro-
exploitation of species, (2) physical alteration of eco- tected areas so that 20% of Exclusive Economic Zones
systems, (3) pollution, (4) introduction of alien species, and the High Seas are protected from threats by the
and (5) global atmospheric change. Scientists have docu- Year 2020.
mented the extinction of marine species, disappearance 3. Ameliorate or stop fishing methods that undermine
of ecosystems and loss of resources worth billions of dol- sustainability by harming the habitats of economically
lars. Overfishing has eliminated all but a handful of Cali- valuable marine species and the species they use for
fornia’s white abalones. Swordfish fisheries have collapsed food and shelter.
as more boats armed with better technology chase ever 4. Stop physical alteration of terrestrial, freshwater and
fewer fish. Northern right whales have not recovered six marine ecosystems that harms the sea, minimize pol-
decades after their exploitation supposedly ceased. Steller lution discharged at sea or entering the sea from the
sea lion populations have dwindled as fishing for their land, curtail introduction of alien marine species and
food has intensified. Cyanide and dynamite fishing are prevent further atmospheric changes that threaten
destroying the world’s richest coral reefs. Bottom trawling marine species and ecosystems.
is scouring continental shelf seabeds from the poles to the 5. Provide sufficient resources to encourage natural and
tropics. Mangrove forests are vanishing. Logging and social scientists to undertake marine conservation biol-
farming on hillsides are exposing soils to rains that wash ogy research needed to protect, restore and sustain-
silt into the sea, killing kelps and reef corals. Nutrients ably use life in the sea.
from sewage and toxic chemicals from industry are
overnourishing and poisoning estuaries, coastal waters Nothing happening on Earth threatens our security
and enclosed seas. Millions of seabirds have been oiled, more than the destruction of our living systems. The situ-
drowned by longlines, and deprived of nesting beaches ation is so serious that leaders and citizens cannot afford to
by development and nest-robbing cats and rats. Alien wait even a decade to make major progress toward these
species introduced intentionally or as stowaways in ships’ goals. To maintain, restore and sustainably use the sea’s
ballast tanks have become dominant species in marine biological diversity and the essential products and services
ecosystems around the world. Reef corals are succumbing that it provides, we must act now.
zoning as an alternative to what Hardin (1968) called vation biology training, and the extreme scarcity of
the “tragedy of the commons.” But to do this suc- funding for research, with the net result being that
cessfully, we must know more than we do now. And if considerable uncertainty (Botsford and Parma, Chap-
unfamiliarity with the sea and its rapidly increasing ter 22) always impedes informed decision making.
conservation needs is the problem, then appropri- The problem, however, goes beyond gaps in data
ately swift development of a vibrant interdisciplinary and theory to a pernicious asymmetry in standards for
science of marine conservation biology must be an in- taking action. While lawmakers and officials have
tegral part of the solution. often accepted “management” plans to exploit species
based on very thin evidence, they often demand sci-
entific “proof” that human activities are harmful,
A Long-standing Problem
and, in its absence, allow harmful activities to con-
Marine scientists have come a long way since the tinue. So long as the burden of proof falls on scien-
Challenger expedition (1872–76) and the founding of tists, losses will accelerate except in those rare cases
the Stazione Zoologica Anton Dohrn (Italy) in 1872 where we can show persuasively—not just to one an-
and the Marine Biological Association (UK) in 1883. other, but to the public, agency officials, and political
We have reason to be proud of what we have learned leaders—that decisive action is essential. The “Pre-
and for the increasing use of scientific information as cautionary Principle” that activities cannot proceed
a basis for conservation decision making. But large unless they clearly pose acceptable risk does not yet
knowledge gaps are still numerous, and scarcity of fact govern most human activities that affect the sea.
and theory bedevils marine conservation decision Moreover, although awareness that there is a prob-
making. A telling sign is that, for some six decades, lem is very recent, impoverishment of the sea has
marine scientists failed to notice the extinction of a been a longstanding problem ( Jackson et al. 2001;
once-abundant nearshore limpet, Lottia alveus (Carl- Crowder, Chapter 2). The gigantic Steller’s sea cow
ton et al. 1991) along a coastline studded with as (Hydrodamalis gigas), ranging from Japan to California,
many marine labs as any comparable stretch in the disappeared almost everywhere as humans spread
world. Similarly, nobody seemed to notice for five along the North Pacific coast. The last ones survived
decades while once-abundant populations of oceanic only 27 years after Western civilization discovered
whitetip sharks (Carcharhinus longimanus) in the Gulf their final island redoubt in 1741 (Scheffer 1973; Ste-
of Mexico were being reduced more than 99 percent jneger 1887). Gray whales (Eschrichtius robustus) were
(Baum and Myers 2004). When marine scientists do eliminated from the Atlantic Ocean in the same cen-
see worrisome phenomena, such as the toxic phyto- tury (Mead and Mitchell 1984), and a number of
plankton bloom or viral disease that devastated Medi- mammals (Day 1981), seabirds (Fuller 1987), and in-
terranean monk seal (Monachus monachus) populations vertebrates (Carlton et al. 1999; Roberts and Hawkins
in Mauritania in 1997 or jellyfish population explo- 1999) followed. Yet despite these extinctions, there is
sions in the Bering Sea and Gulf of Mexico in the late reason for tempered optimism: the number of docu-
1990s, we are often unable to ascertain definitively mented extinctions in marine systems is much lower
why they are occurring. Scientists have not con- than in terrestrial systems. Undoubtedly this is, in
vincingly determined why Atlantic cod (Gadus morhua) part, an artifact of our having overlooked organisms’
and North Atlantic right whales (Eubalaena glacialis) final disappearance, but if it is nonetheless true that
have not rebounded after their exploitation in the far fewer marine species have completely disappeared,
Northwest Atlantic declined sharply or ceased. The rea- we still have important opportunities to recover pop-
sons for this include the youth of our science, the lack ulations and restore ecosystem structure and function
of an institutional basis for supporting marine conser- in the sea.
Why Marine Conservation Biology? 5
However, the loss of marine biodiversity is not

The Need for Marine Conservation Biology
merely the extinction of taxa but the loss of functions
(Soulé et al. 2003). Marine species have also become Before maintaining biodiversity was a widely accepted
economically and ecologically extinct even when conservation goal and conservation biology coalesced
remnant individuals survived. Fisheries for Atlantic as a science, conservation in terrestrial ecosystems
halibut (Hippoglossus hippoglossus), Atlantic cod, white was largely the province of wildlife biologists and
abalone (Haliotis sorenseni), and wool sponges (Hip- foresters whose focus was maximizing populations or
piospongia lachne) arose and then crashed when their biomass of species preferred by hunters and loggers.
populations became commercially extinct (Cargnelli The birth of conservation biology brought two dra-
et al. 1999; Davis et al. 1992; Hutchings and Myers matic changes: (1) it had a broader focus on main-
1994; Witzell 1998). The ecological extinction of sea taining biological diversity as a whole, including bio-
otters (Enhydra lutris), California spiny lobsters (Pan- diversity elements that had previously been of little
ulirus interruptus) and California sheephead (Semi- concern, such as nongame species; and (2) it exam-
cossyphus pulcher) led to widespread loss of kelp forests ined questions relevant to management using insights
in southern California (Dayton et al. 1998). Human from a much broader range of disciplines, including
impact on the sea has a long history, but biotic im- biogeography, landscape ecology, evolutionary biol-
poverishment has accelerated sharply in recent ogy, molecular genetics, genecology, biogeochemistry,
decades, and the hand of humankind is now visible ethnobotany, ecological economics, and environ-
everywhere scientists look. mental ethics. By posing new questions and offering
It is difficult for people to grasp how quickly things useful answers, conservation biology has been sup-
have changed. Gilbert and Sullivan’s 1885 proclama- planting narrower traditional wildlife biology and
tion in The Mikado that “there’s lots of good fish in the forestry approaches. Conservation biologists are now
sea” was largely correct then, and as late as the last providing answers to previously unasked questions,
decade some scientists (e.g., Jamieson 1993) believed such as, How much of Oregon’s ancient coniferous
it is difficult to drive marine invertebrates to extinc- forest is needed to have a 95 percent probability of
tion. Even the wisest people have underestimated hu- sustaining spotted owl populations beyond 2100? and
mankind’s unique power to change the biosphere’s Why do songbird populations decline when coyotes
parts and processes. Our species is now the sea’s dom- disappear from southern California’s urban “islands”
inant predator (Pauly et al. 1998), leading source of of chaparral? Armed with such insights, managers,
seabed disturbance (Watling and Norse 1998), and pri- legislators, and voters can make much better decisions
mary agent of biogeographic (Carlton and Geller 1993) about land use. Conservation biology is increasingly
and geochemical (Smith and Kaufmann 1999; Vi- affecting decision making on issues affecting species
tousek et. al 1997) change. So one of the biggest chal- and ecosystems.
lenges marine conservation biologists face is chang- But because estuaries, coastal waters, and oceans
ing people’s notion that the sea is an inexhaustible weren’t on conservation biologists’ “radar screens,”
cornucopia. nor was conservation on marine scientists’ agendas,
People have shown that we have great power, and the biodiversity-focused, multidisciplinary approach
intemperate use of our power has made us the prob- has, until very recently, been neglected in the marine
lem. Can we accept our responsibility for being part of realm. Most attention has focused on managing
the solution and help our species develop the knowl- fishes, crustaceans, and mollusks that people eat.
edge, ethical values, and institutions needed to reverse Some researchers have paid attention to protecting
these trends? That is the hope that motivates this imperiled megafauna, particularly whales, seabirds,
book. and sea turtles. But marine conservation lags its ter-
restrial counterpart by decades. To illustrate, when the Kuhn (1970) called a “paradigm shift,” a fundamen-
1990s began, there was a substantial scientific litera- tally different way of thinking.
ture on modern extinctions of terrestrial species (e.g.,
Ehrlich and Ehrlich 1981; Wilson 1988) yet none for
Encouraging the Growth of
modern marine species other than mammals or birds.
Marine Conservation Biology
At that time the US Endangered Species Act—which
inspired similar laws in other countries—provided A new science needs a worthy subject, stimulating
protection for hundreds of terrestrial and freshwater ideas, journals, and meetings to discuss them, institu-
plant, invertebrate, and fish species, but for no marine tions that employ and train the senior and young in-
plant or invertebrate species and for only one truly vestigators who will test ideas and discover new ones,
marine fish species (totoaba, Totoaba macdonaldi). Ex- and one more thing: a critical mass of public under-
cept for a minority of commercially fished species and standing and acceptance that can translate into sub-
some “charismatic megavertebrates,” the status of stantial support for research and training. Most fund-
most marine organisms was (and continues to be) ing for this field will ultimately come from public
unknown. sources, and decision makers will not fund marine
The science with the largest effect on management conservation biologists until the public feels this is a
of the sea, fisheries biology, is still concerned mainly high priority. No matter how deserving we are, re-
with assessing “stocks” of commercially “harvested” search grants will not fall into our hands; we must de-
species to maintain biomass production, rather than vote substantial energy to making our case to the
maintaining and restoring biological integrity: species nonscientists whose support will fuel the growth of
composition, habitat structure, and ecosystem func- marine conservation biology.
tioning. It generally measures them in tons, as com- Because building a multidisciplinary science of pro-
modities, not in numbers of individuals (consider tecting, recovering, and sustainably using the living
this: hardly anyone discusses tons of jaguars or sun- sea cannot be done at an unhurried pace appropriate
birds). The utilitarian focus of fisheries biology is anal- for some other scholarly pursuits, marine conserva-
ogous to wildlife biology and forestry before the rise of tion biology needs to avoid some of the digressions
conservation biology. Only recently (e.g., Benaka and other growing pains that are common (and often
1999) has fisheries biology begun considering habitat welcome) in less crisis-oriented disciplines. An essen-
needs of commercially important species in offshore tial shortcut, especially for people trained in the ma-
waters, a concept that leading wildlife biologists such rine sciences, is learning what we can from conserva-
as Aldo Leopold (1933) espoused for terrestrial species tion biology in terrestrial and freshwater realms. Of
long ago. Ideas that became commonplace in terres- course, the marine realm and people’s relations with
trial conservation biology in the 1980s and ’90s, in- it are not the same as with the land and freshwaters,
cluding food web dynamics, metapopulations, pro- so conservation approaches that work elsewhere will
tected areas as islands, connectivity, minimum viable not always work in the sea. Determining which prin-
populations, and restoration ecology, are still at or be- ciples and strategies marine conservation biologists
yond the intellectual horizon of the two fields—fish- can and cannot borrow is one of the greatest strategic
eries biology and oceanography — that have domi- challenges for our infant science. Our work in both
nated marine sciences. The multidecadal gap between nonmarine (Crowder et al. 1992, 1998; Letcher et al.
both the science and the practice of terrestrial and ma- 1998; Norse 1990; Norse et al. 1986) and marine (Carl-
rine conservation at a time of accelerating loss of ma- ton et al. 1999; Crowder et al. 1994, 1995, 1997;
rine biodiversity has created an urgent need for what Crowder and Werner 1999; Norse 1993, 2005, in press;
Norse and Watling 1999) realms has afforded the au- ploit marine populations while few nations exercise
thors useful opportunities to observe the evolution of effective responsibility for them.
nonmarine conservation biology and its relevance to
conservation biology in the sea. Indeed, it embold- Seawater Is Less Transparent than Air
ened us to take on the task of producing this book. Visible radiation and other wavelengths, including
radio waves, penetrate far less through seawater than
through air. On land, aerial and satellite observers can
Conservation-Related Differences between
see through the fluid medium (air) to the biota on the
Nonmarine and Marine Realms
land surface, and radio signals from the land surface
Substantial differences between terrestrial and marine can be picked up by orbiting satellites. But in the sea,
ecosystems (Steele 1985), species, and, most impor- although 98 percent of species live in, on, or just
tant, the ways in which humans think about and above the seafloor (Thurman and Burton 2001), over-
deal with them (Carr et al. 2003; Dallmeyer, Chapter flying aircraft and satellites cannot see the seafloor
24), have important implications for strategies to pro- below a depth of a few tens of meters at most, and
tect, recover, and sustainably use marine biodiversity. they cannot receive signals from radio tags on sub-
Some key differences and implications for conserva- merged animals. Furthermore, at 100 meters’ eleva-
tion follow. tion on land, light intensity is scarcely brighter than
at sea level, but at 100 meters’ depth in the ocean,
The Sea Is Much Larger only 1 percent of the light striking the surface remains
The marine realm is much, much larger than the ter- (Garrison 1999). Photosynthesis is not known to
restrial realm; the area of the Pacific Ocean alone occur deeper than 268 meters (Littler et al. 1985) even
would be great enough to accommodate all of the in the clearest oceanic waters, and no deeper than 70
continents even if there were two Australias. More- meters in clear coastal waters (often much shallower),
over, the sea averages more than 3,700 meters deep, preventing growth of benthic seaweeds, seagrasses,
while multicellular life on land and in freshwaters and photosynthesizing animals such as hermatypic
permanently lives in a thin film that averages a few reef corals. Except in the chemoautotrophic eco-
tens of meters in thickness. Hence, the sea comprises systems of hydrothermal vents and cold seeps, essen-
more than 99 percent of the known biosphere. Many tially all production in the sea depends on nearshore
marine animals deal with the very large scale of ma- plants, benthic algae, and (on a worldwide basis, to a
rine systems by having one or other life history stage far greater degree), on epipelagic phytoplankton.
capable of actively or passively moving over large
distances. CONSERVATION IMPLICATION
It is much more difficult and expensive to do remote

CONSERVATION IMPLICATION observations of species and ecosystems in the depths of
Nations and their marine jurisdictions are small rela- the sea than on land. And anything that affects primary
tive to the ambits of many marine species and human producers and higher trophic levels in the shallows,
activities. To a greater degree than in the terrestrial from nutrient addition to elimination of apex preda-
realm, key ecological processes of marine species and tors, affects nearly all biological activity below them.
ecosystems routinely violate the boundaries among
and within nations that are the spatial basis of gover- The Sea Is More Three-Dimensional
nance. This mismatch of scale leads to numerous Air is a much thinner soup than seawater. Its low buoy-
problems, including the ability of many nations to ex- ancy severely limits the number of species that can fly
or drift in the wind, so nearly all terrestrial life is ben- the sea more complex and renders two-dimensional
thic. Functional groups that are scarcer and much less maps much less useful.
important on land include suspension-feeders, plank-
ton, and nekton. Innovative research tools, such as use Dispersal Stages Are Usually Smaller
of climbing gear to study life in forest canopies (Pike et A majority of marine species whose reproductive
al. 1977), have put nearly all of the terrestrial realm per- modes are known produce very small (less than a
manently inhabited by multicellular life—a layer that millimeter to a few centimeters) gametes, spores, or
is usually a few meters and almost never more than 150 larvae that are dispersed by currents. Many terrestrial
meters in depth—within scientists’ reach. But multi- plants, fungi, and spiders disperse small propagules as
cellular marine life occurs from the sea surface to the well, but most terrestrial animals disperse as subadults
maximum ocean depth of about 11,000 meters. More- or adults, some of them being large and strong
over, the water column is almost always stratified into enough to be tagged, which makes their movements
distinct density layers determined by temperature and easier to track. In contrast, the small size and fragility
salinity, so the sea has far more three-dimensional of marine propagules makes tracking their trajectories
structure than the land. Because of its greater stratifica- very difficult.
tion, biological communities and biogeographic pat-
terns have greater differences at different depths. For CONSERVATION IMPLICATIONS
example, tropical deep-sea ecosystems more closely re- It is difficult for scientists to learn the source of ben-
semble polar deep-sea ecosystems thousands of kilo- thos and nekton in a particular location, a serious
meters distant than the shallow tropical ecosystems just problem for those interested in place-based conserva-
a few kilometers above them. Except in intertidal zones, tion tools such as fishery closures and marine reserves.
direct observation and sampling biota in marine eco- Genetic tags and otolith microchemical analyses could
systems are much more difficult. Less than 2 percent of help scientists understand crucial population source–
the ocean’s average depth is accessible to scientists sink dynamics.
using scuba, and research submarines and remotely op-
erated vehicles are few; limited in depth, range, and du- Marine Species Have Longer Potential
ration; and far more expensive to operate than tools for Dispersal Distances
studying terrestrial life. Indeed, it is much easier to ex- Many terrestrial species have local recruitment and
ploit the sea’s biodiversity than to study it; trawling, for can be conserved within protected areas. When indi-
example, routinely occurs far below depths accessible vidual protected areas are not large enough to support
with scuba. viable populations, corridors of suitable habitat be-
tween protected areas can help populations with
CONSERVATION IMPLICATIONS larger area requirements (Beier and Noss 1998). But, as
Scientists, the public, and decision makers know Grantham et al. (2003) note, a majority of marine spe-
much less about biodiversity patterns and threats in cies whose reproductive modes are known produce
the sea, and the fact that the precautionary principle meroplanktonic early life history stages (gametes,
seldom drives marine management puts the burden of spores, or larvae) that drift in the water column for
proof on scientists to demonstrate that human activ- anywhere from a few minutes to >12 months (typi-
ities harm biodiversity. Many marine plankton and cally days to weeks). At current velocities of 1 to 5
nekton essentially never encounter solid objects, so km/hr, their maximum theoretical dispersal distances
they don’t withstand walls of aquaria or other ex situ range from <10 – 1 to >4 × 104 km. There is increasing
holding facilities. And three-dimensionality makes evidence that actual distances are almost always a very
mapping distributions and biogeographic patterns in small fraction of the theoretical maximum because
currents produce eddies that entrain and slow delivery plankton and nekton, can shift up to tens of kilome-
of propagules ( Jones et al. 1999; Swearer et al. 1999), ters in a day, or in the case of tidally induced island-
and some propagules have behaviors that favor re- wake ecosystems, minutes to hours. Dramatic shifts in
tention near their site of release (Cowen et al. 2000). oceanic pelagic ecosystems are visible in images of
Nonetheless, the potential for long-distance dispersal chlorophyll a concentrations (reflecting phytoplank-
suggests that marine metapopulation dynamics (Lip- ton abundance) or sea surface temperature images
cius et al., Chapter 19) — even with infrequent re- from consecutive days.
cruitment episodes—can operate at a much larger
spatial scale than in terrestrial systems. Furthermore, CONSERVATION IMPLICATION
their dependence on the vagaries of currents also Efforts to classify and manage pelagic ecosystems need
makes recruitment in any one spot far more variable to incorporate scientists’ understanding of these rap-
than on land; indeed, a canonical belief in fisheries bi- idly shifting ecosystem boundaries. Spatial ecology in
ology is that there is almost no correlation between marine systems must become dynamic and three-di-
the number of young produced and the number that mensional and not rely on two-dimensional “snap-
recruit to the population. shots” that are used in landscape ecology.
CONSERVATION IMPLICATIONS Primary Producer and Consumer Biomass Are

Connectivity works differently in the sea. Corridors of Much Patchier in Time
suitable benthic habitat between protected areas are On land, the dominant primary producers are sessile
not needed for dispersal of planktonic larvae, al- and large and live years to centuries. In the sea, their
though they may be valuable for species that migrate counterparts are planktonic, microscopic, and live
as postsettlement juveniles or adults. Single protected days to months. Phytoplankton respond to favorable
areas will not always be large enough to conserve ma- environmental changes with increases in numbers
rine populations except those with direct develop- and changes in species composition much faster than
ment or short-lived planktonic larvae. Recruitment land plants, so the distribution of the sea’s producers
from the plankton depends so much on variable cur- is much patchier in space and time. Blooms of certain
rents that networks of protected areas arrayed at a phytoplankters stimulate population growth of zoo-
wide range of distances will best accommodate con- plankters, whose dense aggregations attract predators.
servation needs of different species by allowing pop- Many species of fishes, seabirds, sea turtles, and ma-
ulation sources to repopulate population sinks (Crow- rine mammals in neritic and oceanic pelagic eco-
der et al. 2000). systems range hundreds to thousands of kilometers,
crossing desertlike oligotrophic waters to locate
Pelagic Ecosystems Undergo Rapid ephemeral food-rich, oasislike patches.
Spatial Shifts
Benthic marine ecosystems, like their terrestrial coun- CONSERVATION IMPLICATIONS
terparts, usually take years, decades, or more to The dense but often short-lived concentrations of
change position in response to changes in overlying large, valuable species make locating and killing them
fluids; neither cerianthid anemones nor beech trees very lucrative (in the extreme cases of bluefin tunas or
shift quickly in response to transitory cooling episodes baleen whales, a single individual can bring tens of
or nutrient pulses. But pelagic ecosystems are differ- thousands of dollars). Large-scale movements increase
ent. Identifiable physical ecosystem features, such as organisms’ vulnerability to ambush predators, such as
currents, water masses, convergence zones, eddies, pelagic longliners and driftnetters, but make them dif-
and upwellings, with their distinctive assemblages of ficult to protect because they cross through the juris-
dictions of diverse nations with diverse laws and en-

forcement capabilities. Thus marine conservation de- CONSERVATION IMPLICATION
cision making needs to occur at interstate and Fisheries managers and laws have largely overlooked
international levels to a greater degree than terres- the importance of seabed structure-formers as habitat
trial decision making. Also, marine species at higher elements because they are more difficult to see (hence
trophic levels are more vulnerable to starvation- less studied) in many benthic communities than in
caused population declines and mass mortalities due terrestrial communities.
to changes (natural and anthropogenic) in weather
and climate (Glantz 1996; Mathews-Amos and Bernt- There Are Steeper Gradients with Distance
son 1999). from Shore
In general, temperature and salinity fluctuations, nu-
Structure-Formers Are Smaller trient runoff, sediment resuspension, productivity,
Like the land surface, the seafloor is structurally com- seabed disturbance, species’ growth and reproduc-
plex due to geological processes and structure-forming tion, fishing pressure, recreation, and pollutant levels
living things. Although giant kelp fronds can be are highest in shallower waters near shore. Deeper wa-
longer than the tallest trees, few marine structure- ters farther from shore are much quieter. The largest
formers are as large as shrubs, let alone trees. Marine storm waves can disturb the seabed only as deep as 70
ecosystems dominated by large (>50 cm), living three- meters (Hall 1994), although severely disturbing re-
dimensional structures, including kelp forests, man- suspension events can occur in some places in the
grove forests, seagrass beds, gorgonian-sponge forests, deep sea (Gage and Tyler 1991). Because primary pro-
and coral reefs, actually constitute a very small (albeit duction is limited to the upper layers of the water col-
very important) part of the marine realm (for exam- umn except in hydrothermal vents and cold seeps,
6
ple, shallow-water coral reefs occupy only 0.6 × 10 waters are progressively more food limited (and
km2 or 0.1 percent of the Earth’s surface; Reaka-Kudla colder) with increasing depth, so growth and repro-
1997). The benthic ecosystems that people see most— duction are slower. The scarcity of large-scale severe
the narrow bands of sandy beach—appear almost fea- disturbance has exerted far less selection pressure on
tureless because pounding waves prevent persistence deep-sea species to resist or recover from disturbance.
of most biogenic structures (e.g., sponges, corals, am-
phipod tubes, polychaete worm tunnels, sea cucum- CONSERVATION IMPLICATIONS
ber fecal deposits). But many sandy bottoms and, to Nearshore species and ecosystems are both most heav-
an even greater extent, mudflats and the vast ex- ily impacted by humans and fastest to recover from
panses of the deeper muddy seafloor, are riddled and pulsed seabed disturbance (such as trawling impacts)
covered with structures, albeit ones often too small to and exploitation. Species in ecosystems that are
be resolved by cameras towed meters above the deeper and farther from shore may have high bio-
seabed. Yet, no less than on land, seafloor structures mass, but they recover more slowly, making them and
are crucial habitat features for most of the world’s ma- their habitats less suitable for exploitation (Merrett
rine species. They provide habitat for organisms that and Haedrich 1997).
raise their feeding and respiratory structures above the
slow-moving, often hypoxic bottom boundary layer, The Sea Is Geochemically Downhill from
and provide hiding places from predators. Coral reefs the Land
alone host 25 percent of the world’s marine fish spe- Rainfall washes materials deposited on the land, in-
cies (McAllister 1991). cluding nutrients and toxic materials, into streams or
storm drains and eventually into estuaries and coastal pend on our willingness to burden future generations
waters. Other chemicals manufactured on land are de- with the irreversible effects of our wastes.
posited at sea through aerial deposition. Any sub-
stance manufactured on land thereby finds it way into Opportunities for Ex-Situ Conservation
the sea. Because there are few pathways by which such Are Fewer
materials are returned from the sea to the land (per- Although some marine species can be spawned and
haps the most prominent being spawning migrations reared in laboratories, aquaria, and aquaculture facil-
of anadromous fishes), what happens on land has far ities, that number is far smaller than for terrestrial spe-
more influence on the sea than vice versa. cies and is likely to remain so for a long time. Among
the reasons: (1) seawater chemistry is difficult to man-
CONSERVATION IMPLICATION age, (2) the young of many species are small and dif-
Except in riparian systems driven by anadromous ficult to feed, and (3) oceanic plankton and nekton
fishes, terrestrial conservation is little affected by any have not evolved in a world with solid barriers.
but the most profound changes in adjacent marine eco-
systems, but marine conservation, especially in estuar- CONSERVATION IMPLICATION
ies and coastal waters, is critically affected by human ac- Even more than on land, marine conservation must
tivities on land. Conservation in marine systems can rely mainly on in situ methods for the foreseeable
necessitate the ability to modify activities on land. future.
Nutrients and Pollutants Become Unavailable Humans Depend Far More on Consuming
Until They Are Returned by Circulation Marine Wildlife
On land, nutrients and many pollutants deposited on Most land animal protein that humans consume is
the ground are quickly decomposed and become avail- from species that are domesticated and farmed; in the
able for uptake by plants. But in oceanic ecosystems, sea, most is from wild species. Unlike livestock farming,
a log, whale carcass, dead fish, or copepod fecal pellet humans generally do not control breeding, feeding,
can survive intact until it sinks below the euphotic waste disposal, or disease in marine species (which is
zone into waters where light levels are too low for why the term harvesting is inappropriate and misleading
photosynthesis. Decomposition in the black water and is not used here except in reference to aquaculture).
column and on the seabed releases nutrients and car-
bon dioxide that become available to photosynthe- CONSERVATION IMPLICATION
sizers only when upwelling brings them into the As capture fisheries decline due to overfishing, aqua-
oceanic or neritic euphotic zone, which takes an av- culture is increasing substantially and is likely to ex-
erage of hundreds of years. pand from estuarine (e.g., oysters) and anadromous
(e.g., salmon) species to truly marine species such as
CONSERVATION IMPLICATIONS moi or six-fingered threadfin (Polydactylus sexfilis) in
The delay between the time when human activities Hawaii. But for now, the sea is in the late stages of
alter the flow of detritus to the deep sea (e.g., Smith and what Safina (1997) calls the “last buffalo hunt.”
Kaufmann 1999) and when they begin to affect pro-
ductivity in the shallows is so long that humans could Technologies for Killing Wildlife Are Less
be committing surface waters to essentially irrevocable Selective and Evolving Faster
alteration in nutrient cycling and productivity. Schemes On land, technological improvements have made it
to dump pollutants (including CO2) in the deep sea de- easier for hunters and more difficult for their targets,
but excesses from more powerful technologies have tions’ exclusive economic zones, or EEZs), comprising
been countered with changes in both cultural conven- much of the oceanic realm, are not owned by anyone,
tions and game laws (Posewitz 1994). The commercial and EEZs are controlled (but not technically owned)
hunting in the 1800s that caused the extinction of pas- by coastal nations. There are strong disincentives for
senger pigeons (Ectopistes migratorius) and almost elim- individuals to conserve publicly controlled or owned
inated American bison (Bison bison) is banned in the resources that they can exploit freely, a situation
United States, and weapon technologies for subsis- called the “tragedy of the commons” (Hardin 1968).
tence and sport hunting weapons have not greatly im- On land, ownership confers the right to destroy bio-
proved the hunter’s advantage in decades. Moreover, diversity, but our “social contract” with the sea is
hunters cannot legally use nonselective methods such even more harmful to the sea’s biodiversity.
as poison gas, explosives, or bulldozers to kill wildlife.
In the sea, however, technologies to transport fisher- CONSERVATION IMPLICATION
men and to find and catch fish have improved con- The tragedy of the commons is manifested as wide-
tinuously, while selectivity has decreased. Steel hulls spread overcapitalization of the fishing industry,
have replaced wood, diesel engines have replaced sail, which means “too many boats chasing too few fish.”
freezers have replaced salt curing and crushed ice, Even in countries where the rule of law generally pre-
nylon has replaced hemp, global position systems have vails on land, the fishing industry often succeeds in
replaced sextants, and satellite oceanography feeds, pressuring governments to allow overfishing and use
precision depth finders, and fish finders have replaced of fishing gear whose wholesale killing of nontarget
guessing where fish are. Larger and more powerful species and damage to seafloor habitat would be less
oceangoing vessels are deploying gear such as 60-mile likely on land. Some nations that show signs of good
pelagic longlines armed with thousands of hooks and stewardship within their EEZs lose all their inhibitions
rockhopper and roller trawls that can fish even on on the high seas and in the waters of other nations. As
mile-deep boulder and reef bottoms. a result, achieving international cooperation for man-
aging the commons will be an exceptional challenge.
CONSERVATION IMPLICATION
Improving technologies have turned the seas trans- Species and Ecosystems Have Far Less
parent and allowed people to fish anywhere in the Cultural and Legal Protection
world. Fishes that once escaped in the vastness of an Traditional societies that depend on the land and sea
opaque ocean or on bottoms too rough to trawl no have survived by evolving customs, often institution-
longer have a chance. Customs and laws to curtail our alized in religion and law, that encourage sustainabil-
increasing advantage over marine wildlife have not ity. In some modern societies, scarce large carnivores
kept apace. such as eagles, tigers, bears, and wolves are increas-
ingly prized for their roles in ecosystems and for rep-
Lack of Ownership and Responsibility Are resenting wildness, and enjoy special protection. In
Even Less Favorable to Conservation many countries, the largest and oldest structure-
Although humankind generally does not create land, forming species (e.g., trees) are protected from all pre-
an indication of our sense of entitlement is that peo- ventable anthropogenic disturbance in parks. In the
ple almost everywhere have the legal right to own it. sea, de jure and de facto protections are generally
On land there are some circumstances that do not en- much weaker. Scarce large carnivores such as sharks,
courage people to harm biodiversity or pollute land tunas, billfishes, groupers, and (in some countries)
they own. But under the UN Law of the Sea, the high dolphins are prized mainly for their meat, and man-
seas (the 64 percent of the marine realm outside na- agement agencies typically deal with them in terms of
tonnage, not numbers of individuals. Agencies differ- private funders. In view of the fact that the sea occu-
entiate protections of “fish” from those of “wildlife,” pies 71 percent of the Earth’s surface and constitutes
coastal national parks that prohibit hunting on land >99 percent of the biosphere, our inattention to it is
allow fishing in the sea, and even many people who striking.
consider themselves vegetarians eat fish. The largest
structure-formers in benthic ecosystems — corals,
Principles That Transcend Land–
sponges, and seaweeds—are commonly demolished
Freshwater–Marine Differences
by dynamite fishing, muro ami (fishing done by
pounding and crushing corals underwater to scare As challenging as maintaining terrestrial or freshwater
the fishes toward the nets), trawling, and dredging, biodiversity is, the preceding factors combine to make
and receive effective protection in relatively few un- marine conservation even more so. But in fashioning
dersea areas, especially in the deep sea. Furthermore, the framework for marine conservation biology, it is
responsibility for managing marine life is often held important to remember the fundamental similarities
by agencies different than those that manage activities between sea and land:
on land, so marine management agencies generally
• Biodiversity is threatened by the same proximate
have no authority to modify terrestrial activities, such
factors: overexploitation, physical alteration of
as logging, mining, construction, and agriculture, that
ecosystems, pollution, alien species, and global
harm the sea’s ecosystems.
climate change.
CONSERVATION IMPLICATIONS • Biodiversity is threatened by the same ultimate

factors (driving forces): overpopulation, excessive
The prevailing lack of strong marine environmental
consumption, insufficient understanding, under-
ethics is reflected by the scarcity of customs and weak-
valuing nature, and inadequate institutions.
ness of laws to protect marine biodiversity. Laws and
policies reflect people’s values. Underdeveloped pub- • Ignorance and indifference are the greatest
lic awareness and concern about the oceans and the enemies of biodiversity.
absence of an Aldo Leopold who can articulate a “sea • Protection is essential but attempts to preserve
ethic” for our time have led to insufficient protection the status quo may fail because we have already
for marine biodiversity. reduced biodiversity so much; active efforts for
population recovery and ecosystem restoration
Much Less Is Spent on Conservation
are increasingly necessary.
Because conservation in the sea is seen as a lower pri- • The sea and land are heterogeneous physico-
ority than terrestrial conservation, it receives even chemical, biological, cultural, economic, and
less funding. For example, in fiscal year 1999, the se- legal mosaics, so management solutions need
riously underfunded US National Park Service had a to reflect this diversity.
budget of $1,700 million, whereas the National Ma-
• There are several distinct kinds of rarity (Rabino-
rine Sanctuary Program had a budget of $14.3 million
witz et al. 1986), ranging from local endemism
(Hixon et al. 2001), a 119-fold difference. Despite
to quasi-cosmopolitan rarity, with very different
some improvement in this ratio, differences are still
implications for conservation.
unjustifiably large.
• Small populations are at special risk.
CONSERVATION IMPLICATION • Top carnivores, other keystone species, and struc-
Spending reflects society’s perceptions. Marine con- ture-forming species are crucial conservation pri-
servation is not yet a priority for many nations and orities because of their exceptionally important
interactions with other species and on ecosystem mer are given overriding legal, budgetary, and po-
processes. litical authority.
• Even species with huge populations and seemingly • Managing biodiversity is primarily about manag-
robust reproductive modes can be eliminated by ing humans.
technological “advances” and by failures to diag- • Resources are scarce.
nose declines and to act effectively and quickly.
• Time is short.
• In situ conservation is preferable to ex situ meth-
ods whenever possible.
• Conserving species one by one is essential in Conclusions
some cases, but places an enormous drain on In the book that introduced conservation biology to
scarce intellectual and financial resources; most the world, Soulé and Wilcox (1980) issued “[a]n emo-
species and ecosystem processes are best con- tional call to arms”:
served in the context of their habitats.
The green mantle of Earth is now being ravaged and
• Conservation of charismatic species in their habi- pillaged in a frenzy of exploitation by a mushroom-
tats can provide an “umbrella” for less favored ing mass of humans and bulldozers. Never in the 500
species. million years of terrestrial evolution has this mantle
• Protected areas are essential but not sufficient; we call the biosphere been under such a savage at-
tack. Certainly there have been so-called “crises” of
the effectiveness of management outside
extinction in the past, but the rate of decay of bio-
protected areas often determines the success of
logical diversity during these crises was sluggish
parks and reserves.
compared to the galloping pace of habitat destruc-
• There is enormous, pernicious disparity between tion today. . . . This is the challenge of the millen-
the conservation capabilities of wealthier and nium. For centuries to come, our descendants will
poorer nations; the countries poorest in financial damn us or eulogize us, depending on our integrity
resources generally have the most biodiversity and the integrity of the green mantle they inherit.
to lose.
Now it is early in another millennium. Armed with
• There are strong vested interests who oppose the new understanding that our species is ravaging the
conservation. Earth’s blue mantle as well as its green one, marine
• Laws to protect, recover, and sustainably use spe- conservation biologists are about to become key play-
cies and ecosystems are essential but not suffi- ers in determining the future of marine life. Having
cient; public support to ensure compliance and begun this work mostly because of our fascination
enforcement is essential. with living things, the current generation of marine
conservation biologists faces the appalling certainty
• Technological change can work for conservation
that we will be the last one able to study many marine
as well as against it.
species and ecosystems unless we succeed in asking
• Vigilance must be unending due to a fundamen-
the key questions and conveying our insights and val-
tal asymmetry: species and ecosystems must be
ues to people whose decisions are shaping their fate.
conserved forever, but even the briefest failure to
Some decisionmakers pass legislation, set bound-
conserve can be irreversible.
aries on maps, determine quotas, command fleets, or
• Ministries and agencies charged with protecting initiate lawsuits. Some write news stories or checks.
the environment are almost always weaker than Some catch fish. Some teach. And some merely con-
advocates for exploiting resources unless the for- sume, produce waste, reproduce, and vote. Most of
these six (soon to be seven, then eight . . .) billion de- of a marine invertebrate in an ocean basin: The de-
cisionmakers (for it is not only political leaders who mise of the eelgrass limpet Lottia alveus. Biological
make decisions that affect the sea) won’t seek our wis- Bulletin 180(1): 72–80
dom by reading our learned papers or taking our Carr, M.H., J.E. Neigel, J.A. Estes, S. Andelman, R.R.
classes. To reach them with a message so compelling Warner, and J.L. Largier (2003). Comparing marine
that they act quickly and effectively, we must tran- and terrestrial ecosystems: Implications for the de-
scend ourselves and use our intellectual and other tal- sign of coastal marine reserves. Ecological Applica-
ents to do what few scientists have done before. Un- tions 13(1) Supplement: S90–S107
doubtedly reaching out to nonscientists will take Carson, R. (1962). Silent Spring. Houghton Mifflin
many of us beyond our “comfort zones.” But our Company, Boston, Massachusetts (USA)
cause is great, and the prospect of succeeding makes Cowen, R.K., K.M.M. Lwiza, S. Sponaugle, C.B. Paris,
the effort worthwhile. and D.B. Olson (2000). Connectivity of marine
populations: open or closed? Science 287: 857–859
Crowder, L.B. and F.E. Werner (1999). Fisheries
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2 Back to the Future in
Marine Conservation
Larry B. Crowder
Oh the sea is so full of a number of fish,

if a fellow is patient he might get his wish.
D R . S E U S S , McElligott’s Pool (1947)
Nearly 60 years ago when Dr. Seuss penned McElligott’s This year, with the sardine population perhaps at a
Pool, the world was beginning to recover from a dev- record low, we have the greatest number of plants in
astating global economic depression followed by a war the history of the industry, with a greater number of
that was fought at an unprecedented scale both on larger and more efficient boats than ever before,
land and at sea. The world seemed full of possibilities scouring the ocean more intensely than any time in
the past . . . we can answer for ourselves the question
and the sea’s resources inexhaustible. We approached
“What became of the fish”: they’re in cans.
the postwar era with enthusiasm to rebuild our
economies and infrastructure; we also took advantage
of new technologies developed in the time of war to After the collapse of the California sardine fishery,
enhance our ability to exploit fishes in a time of Monterey’s Cannery Row rusted and eventually tran-
peace. Dr. Seuss promoted the myth that the sea con- sitioned to fishing for tourists rather than sardines.
tained abundant, diverse fish populations in 1947, de- Many of the fishermen and processors moved on to
spite data from the North Sea that shows unequivo- Peru to fish Peruvian anchoveta (Engraulis ringens),
cally that fish populations were suppressed by fishing which by the late 1960s supported the largest fishery
well before both world wars, recovered during the in the world. And the fantasy of inexhaustibility con-
wars due to the fishing hiatus, and were subsequently tinued along with the elaboration of the theory of spe-
suppressed when fisheries were reinitiated. On the US cies-based fisheries management. Managers argued
Pacific Coast, California sardine (Sardinops sagax) pop- that fisheries could now be exploited sustainably
ulations that once seemed inexhaustible were begin- using state-of-the-art stock assessment techniques.
ning to collapse. While scientists and managers de- But the anchoveta fishery collapsed in 1972 despite
bated the cause of this decline, they did little to international efforts to provide sound management
prevent it from occurring. Ed Ricketts (scientist, advice. We later learned that climate-driven variation
philosopher, and inspiration for “Doc” in John Stein- in ocean productivity drives production of anchoveta
beck’s Cannery Row) wrote in December 1946: and the food web of organisms (and fishermen) that
19
depend upon them. Ah, the kneebone connected to ing baseline syndrome was introduced a decade ago by
the thighbone . . . sounds like Dr. Seuss. Daniel Pauly (1995). The shifting baseline syndrome
About the same time, the good doctor wrote the arises because scientists and managers of each genera-
environmental touchstone, The Lorax (1971) con- tion accept as a baseline the conditions in marine sys-
cerning the effects of clearcutting the forest on the as- tems and of fisheries populations they observed early
sociated wildlife, the habitat, and ultimately the econ- in their careers and then document changes over the
omy. “I am the Lorax, I speak for the trees,” still next 30 years or so. The next generation does the
resonates for many children and for many adults same without reference to what previous generations
whose attention has been drawn to the devastating ef- have documented. The result is a gradual shift of the
fects of human activity on the biodiversity of terres- baseline in terms of what species were caught and at
trial systems. In the past decade, we have become in- what levels in the past. Fisheries managers often lack a
creasingly aware of the impact of human activity on sense of history of periods longer than a career and
the biodiversity of marine systems (Vitousek et al. routinely ignore the evolutionary history that drives
1997; Botsford et al. 1997). Fished species are in de- the life history of the organisms whose populations
cline—some, including Pacific salmon and rockfishes, they manage (Law and Stokes, Chapter 14).
have been petitioned for or placed on the endan- As Jackson (1997) points out, this temporal myopia
gered species list. Other species, including sea turtles, also afflicts scientists. Most of modern reef ecology fo-
seabirds, marine mammals, and sharks taken unin- cuses on the last 50 years, but reef systems in the
tentionally (as bycatch) in fisheries are also in steep Caribbean had undergone several hundred years of
decline. Coastal zones and other critical marine habi- change and loss of their megafauna before scientists
tats are degraded, polluted (often anoxic), and overrun began careful observations or experiments.
with invasive species. If Dr. Seuss were still with us, it
would be timely to hear from the Lorax’s salty Studying grazing and predation on reefs today is like
cousin—“I am the Meernet, I speak for the seas.” The trying to understand the ecology of the Serengeti by
studying the termites and the locusts while ignoring
seas are no longer full of a number of fish and even a
the elephants and the wildebeeste. Green turtles,
(very) patient fellow may not get his wish.
hawksbill turtles and manatees were almost certainly
comparably important keystone species on reefs and
Marine Ecosystems: Extending the History seagrass beds. . . . Loss of megavertebrates dramati-
cally reduced and qualitatively changed grazing and
[H]e departed, and went to his own boat again,
excavation of seagrasses, predation on sponges, loss
which he had left in a little cove or creek adjoining:
of production to adjacent ecosystems, and the struc-
as soon as he was two bow shot into the water he
ture of food webs. ( Jackson 1997).
fell to fishing, and in less than half an hour, he had
laden his boat as deep, as it could swim.
Arthur Barlowe (1584) on an encounter Jackson (1997) urges us to take the long view with
with a Native American fisherman (Stick 1998) respect to baselines for conservation. Marine systems
have a history, often better documented than most
Although scientists have been drawing attention to people assume, about what was, what was removed,
overfishing of target populations for over a century, and what might be recovered. Although the estimates
publications in the last decade have shown this impact are speculative, the Caribbean is thought to have con-
to be much more extensive and widespread than pre- tained tens to hundreds of millions of green turtles be-
viously understood. Attention to the historical condi- fore exploitation by humans (Bjorndal and Jackson
tions in marine ecosystems and of fished populations 2003; Jackson 1997). These numbers are difficult for
is gathering increased attention. The idea of the shift- contemporary sea turtle biologists and marine ecolo-
Back to the Future in Marine Conservation 21
gists to fathom. But the impact of grazing turtles at Maclean 2003; Pauly et al. 1998). Fisheries initially ex-
these densities would cause Caribbean reef/seagrass ploited large, long-lived, iteroparous predators. Their
mosaics to look much different than they do today. As long reproductive life history buffered their popula-
Jackson rightly notes, few species of marine mega- tions against recruitment variation—they were also at
fauna are globally extinct, though many are now eco- the top of the food web, so their populations were
logically extinct. The fact that these species still exist probably food, rather than predation, limited. They
means that the potential for restoration of ecosystem could be managed initially using single-species popu-
structure and function remains in marine systems lation dynamics models and concepts like maximum
and provides opportunities forgone in many terrestrial sustainable yield (MSY). But as these “old growth”
systems where the megafauna were forever lost or re- fishes were driven to low levels, fisheries moved on to
duced to an irrecoverable remnant. shorter-lived, lower trophic level fishes and inverte-
Dayton et al. (1998) make a similar case for reduced brates. This concept is termed “fishing down marine
abundance and ecological extinction of many large food webs” and is a well-documented transition in
consumers in the kelp forests of the Northeast Pacific. fisheries exploitation since 1950 (Pauly and Maclean
In this case, the best baseline data extend back only 30 2003; Pauly et al. 1998). This shift to short-lived, lower
years and characterize the abundance of the major trophic level fishes also redirected fisheries to formerly
structure-forming species, the giant kelp (Macrocystis predation-limited populations, which are more vul-
pyrifera). Kelp distribution and abundance are highly nerable to recruitment variation and to subsequent
influenced by changes in ocean productivity at a va- variation in population size. Because we are now fish-
riety of scales from storm events, to El Niño/Southern ing from the middle (rather than the top) of the food
Oscillation fluctuations, to decades-long regimes of web, single-species stock assessment is no longer ef-
the Pacific Decadal Oscillation. The kelp “baseline” is fective (if indeed it ever was). Pauly et al. (1998) note
thus a very dynamic one. We have insufficient his- that fishing down marine food webs initially increased
torical data for the major consumers (e.g., sea otters, total biomass yields, but later led to declining catches.
large fishes, urchins, abalones) before exploitation to We’ve long known that fisheries reduce the bio-
understand what role they played in these food webs. mass of exploited populations, particularly in inten-
But their reductions in biomass were undoubtedly sively fished coastal regions like the North Sea, the
substantial and exploitation by humans began in this Gulf of Thailand, and the North Atlantic (Gulland
system some 10,000 years ago. 1988; Pauly and Maclean 2003). Researchers have re-
cently raised concerns about the multiple impacts of
In terms of the megafauna of kelp forests, the base-
industrialized fishing on target stocks, bycatch, and
line had shifted considerably before the ecological
habitat (Chuenpagdee et al. 2003; Dayton et al. 1995;
roles of most of these species could be studied and,
Pauly and Christensen 1995; Pauly and Maclean 2003;
in many cases, even before fishery-dependent data
Pauly et al. 2002). These concerns led to a United Na-
were collected. Available data on ghosts of once
abundant species are scarce, of limited value, or an- tions resolution on restoring fisheries and marine eco-
ecdotal. (Dayton et al. 1998). systems (UN 2002). But the impact on large predatory
fishes is global—a recent estimate suggests that the
As industrialized fishing expanded after World War biomass of large predatory fishes now is at least 90
II, our technological abilities also increased fishing percent lower than preindustrial levels (Myers and
power of individual vessels. This led to a rapid increase Worm 2003, 2004). Nearly 80 percent of the observed
in global fisheries yields. These yields (taking into ac- declines occurred in the first 15 years of exploitation,
count discarded bycatch) now exceed the productive often before stock assessments could be completed or
capacity of the sea (Dayton et al. 2003; Pauly and fishery-independent surveys begun. Myers and Worm
(2003) point out that “management based on recent whether additional effects of species removals cas-
data alone could be misleading.” Right now we are cade to the structure and function of the entire marine
trying to establish “sustainable fisheries” on the rem- food web. The notion that the removal of top preda-
nant of the former populations. As in many cases in tors could have dramatic effects on food web structure
marine conservation, we often lack reliable baselines and function actually began with Bob Paine’s (1966,
upon which to base restoration efforts (Dayton et al. 1969) classic species removal experiments in the rocky
1998; Myers and Worm 2003; Pauly 1995). But recent intertidal of the Northeast Pacific. But increasing data
publications suggest populations of marine organ- point to dramatic impacts of removals of large preda-
isms were once much larger than we currently observe tors by fisheries on marine ecosystems.
or even imagine ( Jackson et al. 2001). The cascading relationship between sea otters (En-
Whaling targeted the great whales around the hydra lutris), kelps, and sea urchins is a textbook ex-
world, many of which are now extinct, endangered, or ample in marine ecology. Sea urchins can eliminate
threatened. We know from catch records on depleted kelp populations from particular habitats in the ab-
whale populations that a large proportion of whale sence of predation by otters on urchins. The alternate
biomass was removed as directed takes throughout the states of this community, kelps or urchin barrens, can
world’s oceans. Using genetic analysis of neutral ge- persist for long periods of time (as determined from
netic variation, Roman and Palumbi (2003) estimated the exploration of Indian middens in Alaska; Simen-
that the number of whales in the North Atlantic prior stad et al. 1978). Because kelps are structure-forming
to whaling was an order of magnitude higher than species they create habitat for a vast number of fishes
historical estimates–based whalers’ log book data. In and invertebrates; when kelps are lost, their associates
the paper, they explore various assumptions that are lost as well. The state of this system changes
might cause the genetic analysis to overestimate the spatially as well as temporally. After being protected
baseline numbers of whales in the North Atlantic. from overhunting, recovering populations of otters
Very conservative estimates are still three to five times changed nearshore reefs from two- to three-trophic-
higher than historical estimates from reported ex- level systems by reducing the abundance of urchins
ploitation rates. A recent analysis comes to similar and so promoting kelp forest expansion (Estes and
conclusions about the pre-European densities of green Duggins 1995). However, in the late 1990s, sea otter
(Chelonia mydas) and hawksbill (Eretmochelys imbri- populations declined precipitously over large regions
cata) sea turtles in the Caribbean relative to current of western Alaska. The single best explanation for
population sizes (Bjorndal and Jackson 2003). these declines is increased predation by killer whales
(Orcinus orca) (Estes et al. 1998). In an orca-domi-
nated system, otters are suppressed, urchins recover,
Ghosts and Ecosystem Function
and kelp forests decline, exactly what one might ex-
It seems apparent that species are only commas in a pect from theory given a four-trophic-level system.
sentence, that each species is at once the point and
Other large pinnipeds including Steller sea lions
the base of a pyramid, that all life is relational. . . .
And the units nestle into the whole and are insepa- (Eumetopias jubatus), northern fur seals (Callorhinus
rable from it. ursinus), and harbor seals (Phoca vitulina), also col-
John Steinbeck, Log from the Sea of Cortez (1941) lapsed in the western North Pacific beginning in the
late 1970s. By the late 1990s, when Estes et al. (1998)
Intensive fishing leads to declines in both target wrote their paper, the conventional wisdom was that
and nontarget (or bycatch) species. One would expect these declines were due to food limitation. But recent
this in any fishery. The question is whether the effect analyses suggest that the decline of Steller sea lions is
is limited to the population of the exploited species or much more consistent with hypotheses related to in-
Back to the Future in Marine Conservation 23
creased mortality than to reduced food or any other In Caribbean coral reef–seagrass systems, increased
bottom-up effect (Paine et al. 2003). fishing leads to reductions in sharks and crocodiles
Springer et al. (2003) attribute the sequential declines and the extinction of Caribbean monk seals (Monachus
in this suite of marine mammals to whaling in the tropicalis). Predatory fishes and invertebrates decline,
North Pacific ecosystem. Although whaling had sup- as do grazers, including manatees (Trichecus manatus)
pressed the abundance of a number of great whales spe- and sea turtles. The cascading effects propagate to the
cies prior to World War II, the rapid expansion of whal- corals (which decline) and to the seagrass, sponges,
ing fleets by Japan and the Soviet Union after the war and macroalgae (which increase). The current ecolog-
led to the reported removal of over half a million great ical role of grazing sea urchins, fishes, sea turtles, and
whales from 1946 to the mid-1970s, when whaling was manatees is severely diminished and the primary pro-
severely reduced. Killer whales likely consumed great ducers are flourishing.
whales (in fact they were first dubbed “whale killers” by Finally, heavy exploitation in estuaries has reduced
the early whalers; Scammon 1874) and when the great the abundance of nearly everything with the exception
whales were suppressed, killer whales moved on to har- of jellyfish, worms, phytoplankton, and microbes. This
bor seals, fur seals, sea lions, and finally sea otters. Al- is what Jeremy Jackson refers to as the “rise of the
though there are limited data to support their hypoth- slime.” A key loss in estuaries is the filter-feeding ca-
esis, Springer et al. (2003) have made a compelling case pacity of oysters. In Chesapeake Bay and other East
for potential cascading effects that begin with the re- Coast estuaries, oysters at historical densities filtered
moval of great whales and cascade to killer whales, large the entire volume of the estuary every few days; now
pinnipeds, and ultimately sea otters, which provide a it takes months to years. The accumulating organic
very small energetic reward to a consumer as large as a material not only contributes to water quality prob-
killer whale. Of course, otters and their abundance also lems but leads to substantial problems with hypoxia
influence the rest of the system down to the kelp and and anoxia in the formerly productive estuarine habi-
their fish and invertebrate associates. tat. Of the five threats to marine biodiversity, Jackson
Historical perspectives are critical to setting base- et al. (2001) place fishing effects first in both magni-
lines for the restoration of marine ecosystems ( Jack- tude and time. In most coastal ecosystems, fishing im-
son et al. 2001; www.shiftingbaselines.org). We are pacts were followed in time by pollution, habitat de-
now recognizing that documentation of the structure struction, invasive species, and finally climate change.
and function of our highly altered coastal ecosystems
The historical magnitudes of losses of large animals
provides little insight into the past. An exploration of and oysters were so great as to seem unbelievable
likely changes due to fishing in kelp forest ecosystems, based on modern observations alone. Even seem-
coral reef–seagrass ecosystems, and estuaries all show ingly gloomy estimates of the global percentage of
a dramatic simplification of food webs with cascading fish stocks that are overfished are almost certainly
effects ( Jackson et al. 2001). People dominate current too low. The shifting baseline syndrome is thus even
food webs and top predators disappear or decline to more insidious and ecologically widespread than is
ecological extinction. In kelp forest habitats in Alaska, commonly realized. ( Jackson et al. 2001).
Steller’s sea cows (Hydrodamalis gigas) are extinct, as
are sea mink (Mustela macrodon) along the Gulf of
What the Ocean Was,
Maine. Sea otters, California sheephead (Semicossyphus
What the Ocean Could Be
pulcher), and Atlantic cod (Gadus morhua) in these sys-
tems are reduced to extremely low levels, releasing Our understanding of the magnitude of the human
lower trophic levels (e.g., lobsters, sea urchins) from footprint on coastal ecosystems is just now coming
predation and ultimately reducing kelp abundance. into focus (Botsford et al. 1997; Jackson et al. 2001;
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"Have you the courage?"
He could scarcely falter out,—
"I have!"
Her point was gained. She no more meant her horrible suggestion to be
realized, than he meant to realize it. They were both trying each other. She,
to see the extent of her power; he, to ascertain the truth of his suspicions.
Imagining her power great enough to lead him into any crime, she burst
out laughing.
"And do you mean to say you thought me serious, Marmaduke?"
"I did."
"Then what a villain you must be!"
"I am only your slave."
She shook her head at him, and said,—
"Slaves should not listen to such thoughts. If I thought you were serious,
I should loathe you."
"And I should loathe myself," he said, coldly.
It so happened that both believed the other guilty of the serious

intention, and attributed the disavowal to fear of having been understood.
Marmaduke had noticed the affected tone of her laughter; it was affected,
but not from the cause he imagined: it arose from a sense of uneasiness at
having pushed the experiment too far, and from a dread of his really
believing her to be serious.
On the other hand, she noticed the faltering hesitation and coldness of
his tone, which she interpreted into the uneasiness of guilt, but which really
arose from the intense loathing he felt for her. It only seemed a
confirmation of her power.
Nothing could ever have persuaded Marmaduke that Mrs. Vyner was
innocent of the thoughts he attributed to her; and his loathing was so great,
that it not only completely crushed the sort of love he had felt for her, but
revived his desire for vengeance, which he thought could not be made
terrible enough to fitly punish such a wretch.
He dissembled his disgust, and only more urgently pleaded her to elope
with him. At the conclusion of one of his speeches to that effect, he noticed
that she seemed not to attend to him, but to be eagerly listening. Presently
she put her finger upon her lips by way of caution, and then, in a voice she
strove to make calm and distinct, said,—
"Marmaduke, I do not doubt your love, but I must not, will not listen to
it. I am married. I never can forget that; do not you! If a sisterly regard will
suffice you, that will I give; but you must here engage to think of me as a
brother, and, above all, never again to let me hear from your lips the
language I have heard to-day. Will you promise me?"
She nodded significantly to him to reply in the affirmative, and he said,

—
"Will you, then, give me no hope?"
"None. You have heard my conditions. Do you accept them?"
"If I must."
"You must."
"Then I do."
"That's right. Now go home."
She put her finger again upon her lips, and motioned him to listen.
The gentle creak of retiring footsteps stealing away was then distinctly
heard. As they ceased, she said,—
"That was my husband. He has overheard us. But fortunately he heard
nothing which I cannot explain. Leave him to me."
Marmaduke went home in a state of fever, torn by the most vehement

emotions, and seeing all darkly before him.
CHAPTER VII.
FATHER AND CHILD.
Meredith Vyner stole back to his study, after having overheard a portion
of the foregoing scene, like one who has just received a sentence of death.
He loved his wife with the unreasoning idolatry of one who has centred all
his affections on a single object. His children had been gradually estranged
from him, his wife had taken their place in his heart, and now she was
listening to the vows of another!
What he had heard was enough to make him fear the worst. Her refusal
to listen to Marmaduke, and her offer of a purely sisterly regard, although it
assured him that at present she was resolved not to forget her duty, gave
him no assurance that such prudence would long continue. Could she
restrict herself to that sisterly love? Could she know that one so young, so
handsome, so imposing, loved her, and not at last yield to his love?
He would snatch her from the danger by taking her at once from
London. Away from her lover, she might forget him, or he might seek
another. It was necessary to take a decided step.
When Mrs. Vyner came into his study, he at once assumed an unusual
tone of command, and informed her that it was his pleasure they should at
once return to the country.
"My dear Meredith, what are you thinking of? The country! We cannot
leave town in the height of the season."
"I have my reasons," he said, with as much dignity as he could assume.
"And I have mine for not going."
"But I insist upon it."
She seated herself in one of the easy chairs, and said, quietly,—
"You will not insist when you have heard me. This very morning, Mr.
Ashley has made a foolish declaration of love to me."
He was thunderstruck. The quiet matter-of-fact style in which she

communicated this intelligence, was indeed a masterpiece of adroitness.
There are moments in our lives when audacity is prudence; and this was
one, when nothing but an audacious avowal could, by anticipating, defeat
the accusation she knew he would bring forward. She lost nothing by
avowing it, as she was certain he already knew it; but, on the other hand, by
anticipating him, she was enabled to give her own colouring to the
appearances which condemned her.
"I see your surprise," she added; "you little expected it, nor did I. You
thought he was attached to Violet; I thought so too; and as I am sure Violet
is attached to him, I have set my mind upon the match. But now, look here:
I received his declaration without anger and without encouragement. I told
him I would love him as a sister, and made him promise, on pain of
instantly refusing to see him, to cease all such language, and to crush all
such hopes. Did I act rightly?"
"Yes—very—very."
"But, suppose I run away into the country, what will he imagine? That I
am afraid of him, afraid of myself; that I love him, and avoid him. Do you
wish him to think that? You do not. Then we remain."
"But ... and you ... will you continue to see him?"
"Why not? If I am to avoid him let it be done at once. If not, let us treat
him as if he had never made that silly declaration. He will soon get over
this. It is only a passing fancy. He saw me a mere girl, wedded to one old
enough to be my father, and imagined, as all men would imagine, that I
should be easily persuaded to forget what was due to my husband, and to
myself. I have undeceived him. My coldness and firmness will soon cure
him. He will then think of Violet."
She ceased. He took vast pinches of snuff in an agitated absent manner,

but made no remark. She perceived that she had gained the day, and left
him to his reflections.
Bitter enough those reflections were. The explicit avowal had staggered
him—had taken from him the very weapon he was to use; but it had in no
way alleviated his jealous anguish. He could not answer her—yet could not
satisfy himself. The reference she had made to his age still rung in his ears,
and told him plainly that his rival would one day be happy.
That afternoon Violet and Rose returned. He received them with

unwonted tenderness, for his heart ever yearned to those whom he had
excluded from it, and he felt bitter remorse for having sacrificed them to his
wife. Violet was peculiarly dear to him at this moment. He felt for her
misplaced attachment, and remembered how ill she had been treated at
home. He folded her to his breast, with a lovingness which brought the tears
into her eyes, and as she sat down on his knee, one arm around his neck,
delighted with this change in his manner, she divined at once the real cause
of the change. As it was Mrs. Vyner who had estranged him from her, so
must it be Mrs. Vyner who had brought back his love.
It was a touching sight to see this parent and child united by a common
sorrow, mutually pitying and mutually comforting each other, having in one
embrace forgotten all that had once been distrust and coldness, and now
possessed by that overflowing love which, in its exaggeration, desires to
atone for past coldness.
It was not what they said; for few words passed between them; it was
their eloquent looks, significant pressure of hands, convulsive embraces,
and tones pregnant with meaning. The father mutely demanded forgiveness,
and the child demanded a continuance of love.
After an hour of this intense emotion they grew calmer, and began to
talk of indifferent things. From time to time they hovered about the name of
Marmaduke, and betrayed, in their very recurrence to the subject, and
hesitation in speaking openly of it, how predominant it was in their minds.
At last they ventured on the name. It is impossible to convey an idea of the
conversation which ensued, because it was conducted in phrases of the most
guarded vagueness, but made full of meaning by the looks which
accompanied them. Slowly, but irresistibly, the conviction came upon her,
that her father had discovered his wife's guilty passion; or, at least,
suspected it. Her object, therefore, was, if possible, to persuade him that
Marmaduke came there for herself; and she even went so far as to laugh
faintly at his efforts to make himself agreeable to Mrs. Vyner, by way of
using a stepmother's influence in his favour.
Her voice shook, as she uttered this heroic falsehood.
He gazed at her with mournfulness; a tear rolled down his cheek; his
heart swelled as he sobbed out,—
"My poor child! my poor child!"
He dared not undeceive her, dared not tell her what he knew.
She saw that she was not believed, but little did she know the mournful
pity with which her supposed credulity filled him.
It was a relief to her when the dinner-bell rang, and put an end to their
interview.
He saw her depart, and sat sighing deeply, wholly bewildered at the
inextricable difficulties of his position; and when Mrs. Vyner came in, and
chatted away about the opera, to which they were going that night, as if
nothing whatever had occurred, he almost felt as if he had just awakened
from a dream-troubled sleep.
CHAPTER VIII.
THE CRISIS.
Quelle nouvelle a frappé mon oreille!
Quel feu mal étouffé dans mon cœur se réveille!
Quel coup de foudre ô ciel! et quel funeste avis!
RACINE.—Phèdre.
False! I defy you both:

I have endured you with an ear of fire;
Your tongues have struck hot irons on my face.
CYRIL TOURNEUR.—The Revenger's Tragedy.
There was an appalling struggle in Marmaduke's breast that day as he

reached home. The terrible scene which had passed between him and Mrs.
Vyner, with its plainly expressed hint at assassination, made him shudder as
he again and again went over it in memory. False and heartless he had
known her; but for this he had not been prepared. And he felt a sort of
sickness come over him as he reflected on the peril which he had escaped,
in Vyner's not coming to listen at the very moment when she had proposed,
and he had affected to accept the proposition! He would then have been
accused of having really meant to perpetrate the crime; for who would have
credited his disavowal? who would have credited his assurance that he was
but feigning what his soul abhorred?
By a retrospective glance at his own conduct, and at the peril he had

escaped, he was led to meditate on the nature of this woman; and by a
reflection on her criminal thoughts, he was shown the criminality of his
own.
This revenge which he had planned so remorselessly, what was it but a
crime? If he had been wronged by a heartless woman, was it for him thus to
measure out the punishment? and did her jilting him deserve so terrible a
retribution?
After all, was not vengeance a "wild justice," but only the justice of
savages? Was it worthy of civilized, christianized man? And for a man to
wreak it on a woman, was not that petty, ignoble, more like spite than
retribution?
Such were the thoughts which possessed him. That they never suggested
themselves before, arises from the fact of his never having before been cool
enough to question the legitimacy of his feelings. But now they staggered
him; now they came upon him like a remorse; and he relinquished his
scheme of vengeance!
The next day, impelled by some strange impulse which he could not
explain, he went to the Vyners. Violet observed the agitation of his manner,
and attributed it to meeting her again, after what had evidently transpired
during her absence. She was, therefore, considerably surprised when he
begged for a few moments' private conversation with her, at the same time
entreating Rose to leave them alone. She had intended to refuse the request,
but Rose had departed before she could open her lips. Rose too well
understood the purport of that interview, not to be anxious to forward it by
her absence.
"Mr. Ashley," said Violet, coldly, "there is no subject upon which I can
hear you alone; you will oblige me, therefore, by suffering me to follow my
sister."
"Violet!"
"Mr. Ashley, by whose authority do you address me in that manner?"
"The authority of my love. Violet, I love you ... you know it ... but I
must tell you so ... I must..."
She moved towards the door; he intercepted her, and put his back
against it. Drawing herself up to her full height, with a haughty gesture she
motioned him to let her pass.
"I did not expect this," he said, without moving; "I thought I should, at
least, be heard. Miss Vyner had given me reason to hope that she would at
least suffer me to tell her.... Violet, I cannot control myself. You must know,
you must long have known I loved you, you must have seen it in every...."
"Mr. Ashley I request to be allowed to leave the room."
"Do you refuse to listen to me?"
"I do."
He stared at her bewildered; there was something so calm and collected

in her manner, yet the manner was so incomprehensible, that he was
speechless for a few moments.
"You cannot ... cannot have mistaken ... for so many months ... do you
mean that you mistook my looks ... my words ... my actions? ... Did you?"
"I did not."
"Good God! have you then been playing with me?"
"Playing!" she repeated scornfully, yet sadly. "I play!"
"Then what can all this mean? There is some delusion ... a word may set
it right.... You knew I loved you—did you not? You hear it now. Violet, I
love you—love you as man never loved before. Will you accept that love?"
"Dare you ask me?" she said, fixing her large eyes on his with searching
keenness.
"Violet ... what is it that you doubt?"
"Your purpose."
"My purpose is, to tell you that my heart is yours.... That I live but in the
hope of calling you mine."
Her bosom heaved—her nostrils dilated, and with flashing eyes she
proudly, almost fiercely, exclaimed,—
"Let me pass!"
"What is the meaning of this?"
"Let me pass, sir!"
"Violet! are you mad, or do you think me so? Is my love an insult,

that...."
"It is an insult—a deep insult. Now, sir, will you let me pass?"
"I will know what is at the bottom of all this. You may reject me, but
you shall explain. It is so utterly inconceivable that, after the
encouragement you have given me, you should pretend to regard my
avowal as an insult, that I demand an explanation."
In spite of the rising passion in his breast, he uttered this so collectedly

and so earnestly, that Violet was somewhat perplexed, and began almost to
doubt her own conclusions.
"Mr. Ashley," she said, "a short while ago, such an avowal could only
have been felt by me as an honour; but since that, your own conscience will
tell you why I reject, and reject with deep scorn, the offer of your hand.
Pray let me say no more."
His conscience did tell him, at least it suggested what the cause most
probably was; but wishing to come to an explanation, he said,—
"My conscience tells me that I love you—only you; will you tell me
wherein lies the insult?"
A long struggle ensued in her mind; she could not give him the
explanation he demanded, because unable to bring herself to mention her
stepmother.
"If you persist," she said at last, "I must persist also. I tell you again, the
offer of your love to me—here, in this house, is an outrage, and scorn is my
only answer. Does that suffice? Would you have me add more bitterness to
my refusal?"
"Violet, I cannot quit you without..... Tell me, is there not that in your
mind which you shrink from uttering, and which has reference to some one
in this house?"
"You understand me, then?"
"I do."
"Then let me pass at once."
"Not until you have heard me. Will you hear me.... Will you, in this
solemn moment, let me lay before you the whole history of my heart? You
think me a villain, will you listen before you condemn?"
"I know not what plausible excuses..."
"Truth—the simplest truth shall be my defence. If that condemns me, I

will submit in patience. Will you hear me?"
There was something so solemn and so touching in his tone, that Violet
was deeply affected by it; the sad earnestness of his voice pleaded
eloquently in his favour.
He approached, and took her hand; she withdrew it hastily, and moved
towards the mantelpiece, against which she leaned in an attitude of
exquisite dignity, turning her face towards him, prepared to listen. After
gazing stedfastly at her for a few seconds, while he collected his thoughts,
he thus spoke,—
"Violet, I am about to make a most painful avowal; one that will startle
you; one that will seem wholly inexplicable. When but a boy, I loved—
loved as boys love, unreasoningly and ardently. I have tropical blood in my
veins, Violet, and all passions become intense with me. The girl I loved
returned my affection. We were to have been married. I was called away
from England. I returned to my father in Brazil. My father gave his consent
to our marriage. I wrote to inform her of it: she was overjoyed. Her letters
were as ardent as even I could wish. Suddenly they ceased. My father died.
I was settling his affairs, and preparing to quit Brazil for England, when I
learned from a newspaper that my affianced wife had married another."
He paused: a choking sensation in his throat impeded utterance. Violet

had listened eagerly, and still kept her eyes fixed upon him.
"I cannot tell you," he resumed, "what I suffered on awakening from the
sort of stupor in which this intelligence threw me. You have never known—
may God preserve you from ever knowing it!—what that desolation is,
when those we love are found unworthy of our love! The anguish and
despair which then tore my soul to pieces, I shudder to look back upon. It
was not that my love had been destroyed—it was not that which made the
pang; it was the horrible, heartless cruelty with which I had been deceived. I
had been sacrificed to wealth. That I might have forgiven; but it was done
so cruelly! Until she had accepted her husband, her letters were as
affectionate and hopeful as ever. The blow was unbroken in its fall—no
wonder that it nearly crushed me!"
He paused again; and saw tears glisten in the earnest eyes of his listener.
She, too, had known what it was to suffer from hopeless love!
"Violet, I am fierce and brutal in my instincts, and my education had

been but indifferent—on one point especially it had been deficient;—in the
Christian spirit of forgiveness. Vengeance—the justice of the savage—was
what I had never learned to disown. Writhing under the torture which had
been inflicted, I took comfort solely in the hope of vengeance. I came to
England with that one absorbing object. Now comes the painful part of iny
disclosure—if indeed you have not already guessed it—the girl who had ...
in a word, it was Mrs. Vyner!"
He expected to see her vehemently startled, but she only whispered, in a

hoarse and broken voice,—
"I knew it."
"You knew it! Then have you understood me?"
"Not quite."
"I must explain, then, my conduct further. I was here in England,

resolved on reparation of the wrong I had suffered. I knew not what shape
my vengeance would take, but I was resolved to have it in some shape or
other. I saw you. To know you, was to love you—and I loved. In my love, I
forgot my misery, and ceased to think of revenge. You were sometimes cold
and haughty to me, Violet; sometimes kind and encouraging. Do you
remember when we rode to the sands that afternoon, and sat upon the rock
together listening to the sea? I could have told you then how much I loved
you, had not your coldness chilled me. Well, on that very day, while I was
suffering from your indifference; she, jealous of you, chose to recall me
back again to my schemes, by pretending that her marriage had been an act
of jealous despair; she roused the demon in me by her infernal arts, and
once more I resolved to wreak upon her the vengeance you had made me
forget. From that moment I have pursued a scheme which involved her ruin.
Many times have I been vacillating, many times has a kind word or look
from you brought me back again to a purer atmosphere; but the devil would
have it! and a haughty gesture from you has thrown me back again. Hurried
onwards by the irresistible course of events, I was nearly losing myself for
ever, when last night I had my eyes opened. I saw that the only vengeance
worthy of a man was contempt. At once, I resolved to cease feigning love
for the miserable being whom I had marked as my victim; resolved to break
away from the net in which I was entangled, by quitting England.
"Before I left England, I had only to learn my fate: if you refused me, I
should carry my despair into distant lands; if you accepted the offer of a
heart, I thought you would not refuse to quit England with me. You have
now heard all. I have told you of my crime: if repentance will not clear me
from the stain...."
The door was thrown violently open before he could conclude the
sentence, and Mrs. Vyner stood before them.
They started as at an apparition.
Fearful indeed was the aspect of the little fury, as with bloodshot eyes,
quivering lips, and spasm-contracted face, she trembled before them. All
that was diabolical in her nature seemed roused, and looking from her eyes:
passion made her hideous.
"Your little history is incomplete," she said in a hissing tone; her voice
lowered by the intensity of her feeling; "there is a chapter to be added,
which you will allow me to add. Miss Vyner is so excellent a listener that
she will not refuse to hear it."
Violet looked haughtily down upon her, and said,—
"I desire to hear no more."
"But you must hear this; it concerns you. You cannot be indifferent to
anything which relates to your honourable lover; you cannot be unwilling to
know that he who offers you his hand is vain fool enough to be the dupe of
any woman, as he has been mine. He has told you, and how prettily he told
it! what pathos! what romance! he told you how I played with him. That is
true. He was such a vain silly creature that no one could resist the
temptation. Not only did I make a fool of him as a girl. I have done so as a
married woman. I persuaded him that even respect for my husband, respect
for the world could not withstand the all-conquering beauty of his lumpish
person, and he believed it! believed that his face was a charm no woman
could resist. This besotted vanity brought him to my feet; yes, even at the
time you were sighing for him, he was at my feet, ardent, submissive, a
plaything for my caprice!"
She saw Violet writhing, and her savage heart exulted in the pain she
was inflicting; she saw Marmaduke's calm contempt, and her exasperation
deepened at the unavailingness of her sarcasms to wound him.
Turning from her, as from one unworthy of notice, he said to Violet,—
"I repeat, my fate is in your hands. I love you, love you as I never loved
before—with my whole soul: love you with deep reverence for all that is so
great and noble in you, and to that generous and exalted mind I leave my
errors to be judged."
The sarcasm implied in this avowal almost maddened Mrs. Vyner.
"Accept him, Miss Vyner," she said with a short, hollow, and hysterical
laugh; "pray put him out of his misery; accept the offering of his deep
reverence, for that offering is my leavings!"
Marmaduke and Violet both started as this poisoned sarcasm, issued

from her lips, and their faces told her plainly she had struck deep.
"A reformed rake, you know, makes the best husband," she pursued; "so
that one so inflammable as he is, will be sure to make a constant and
adoring husband. You will be so happy with him! Whenever conversation
grows dull, he can amuse you with narrating little episodes of his love for
me, and my cruelty that will be so pleasant! you will never tire of that!
Accept him: you will be sure never to repent it!"
Marmaduke could have strangled her.
Violet, seeing clearly the purpose of these horrible phrases, cut them
short by saying,—
"Mr. Ashley, on some better occasion we will speak again of this; do not
let the present ignoble scene continue."
She held out her hand to him. He pressed it to his lips. Mrs. Vyner
nearly shrieked with mad jealousy; but suppressed the explosion of her
agony; while Violet swept out of the room, disdaining to give even a
passing glance at her.
Mrs. Vyner sank exhausted into a chair. Her brain was as if on fire, and
her whole frame shook violently with the unutterable rage, jealousy, and
hate which stormed within her heart.
Marmaduke could not in his fiercest moments have desired a more

terrible retribution than that which now had fallen on the miserable woman;
and he gazed upon her with a pity which astonished himself. To this he had
brought her; unwittingly it is true, but he felt it was he who had moved the
stone which had fallen and crushed her; and now that she lay there suffering
before him, his anger had gone, and pity filled its place.
She expected him to speak; she saw his fixed gaze and endeavoured to
interpret it; but he spoke not. Before she was aware of his intention, he had
left the room.
Five minutes afterwards, Meredith Vyner found her apparently lifeless

on the floor: she had swooned.
END OF VOL. II.
London: Printed by STEWART and MURRAY, Old Bailey.

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CONSERVATION IMPLICATIONS Primary Producer and Consumer Biomass Are

dictions of diverse nations with diverse laws and en-

CONSERVATION IMPLICATIONS • Biodiversity is threatened by the same ultimate

Oh the sea is so full of a number of fish,

He could scarcely falter out,—

"And do you mean to say you thought me serious, Marmaduke?"

"Then what a villain you must be!"

"I am only your slave."

She shook her head at him, and said,—

"And I should loathe myself," he said, coldly.

It so happened that both believed the other guilty of the serious

She nodded significantly to him to reply in the affirmative, and he said,

"Will you, then, give me no hope?"

"None. You have heard my conditions. Do you accept them?"