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Living Marine

Menhaden purse-seiner set. The net is hauled aboard each seine boat with power-driven blocks until
the fish are closely confined in the bunt. The catch is then pumped from the purse seine into the hold
of the carrier vessel for transport to the reduction plant. Photo courtesy U.S. Bureau of Commercial
Edwin S. Iversen
Rosenstiel School of Marine
and Atmospheric Science
University of Mjami
Miami, Florida

Living Mart
Their Utilization
and ManaQetnent

New York· Alb:lny • Bonn· Boston· Cincinnati · Detroit

London • ~'iadrid • Melbourne· Mexico City · Pacific Grove
Paris • San Francisco· Singapore · Tokyo • Toronto • WaShington
Cover photo: A salmon purse seine is retrieved using a power block.
Courtesy Alaska Seafood Marketing Institute.
Cover design: Curtis Tow Graphics

Copyright © 1996 by Chapman & Hall

Softcover reprint of the hardcover 1st edition 1996

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used in any form or by any means-graphic, electronic, or mechanical, including photocopying,
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1 2 3 4 5 6 7 8 9 10 XXX 01 00 99 98 97 96

Library of Congress Cataloging-in-Publication Data

Iversen, Edwin S.
Living marine resources: their utilization and management / Edwin
S. Iversen
p. cm.
ISBN-13: 978-1-4612-8513-7 e-ISBN-13: 978-1-4613-1211-6
001: 10.1007/978-1-4613-1211-6
1. Fisheries. 2. Marine resources. 3. Fishery management.
I. Title.
SH331.194 1996
338.3'727- - dc20 96-16566

To order this or any other Chapman & Hall book, please contact International Thomson
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This book is dedicated
to the memory of Donald L. McKernan, friend and colleague,
in recognition of his enormous contribution
to national and international fisheries

Preface ix
Introduction xi










viii Contents








Appendix A Selected General References 389

Appendix B The Magnuson Fishery Conservation and Management Act 393

Subject Index 397


Although there is voluminous literature on the subjects covered in this book, I have tried to
summarize the main points of each subject that will helpful to the reader. This book is di-
vided into five major parts. Part One, Ocean Environment and Resource Species, is organ-
ized so that the reader will understand enough about the ocean environment and kinds of
renewable resources to appreciate fishing methods and fisheries management as described in
subsequent chapters. Space does not permit them to be covered in detail in this volume. A
basic knowledge of the ocean environment is presupposed, and here only those aspects of the
oceans most important to fisheries are reviewed. The uneven distribution of living resources
in the sea is discussed together with reasons for differences in ocean productivity in different
regions, and the importance of seafood in the world's economy.
Humankind can reap the greatest return from the sea harvest without exceeding its carrying
capacity only if we wisely manage the stocks of fish and shellfish in the sea. Many aspects of
the material in Part Two, Fisheries Biology, age, growth, fecundity, mortality, etc., must be un-
derstood to assist in the efficient harvesting of seafood and to serve as background information
for stock management. With this knowledge in hand the student can appreciate yield models
designed to give estimates of maximum sustainable yield and maximum economic yield from
the sea. Therefore, in Part Two, biological aspects of fish and shellfish populations necessary
for management are described as to how they are studied and applied to management.
Part Three, Fisheries: Gear, Methods, and Landings, covers the history of fishing and vari-
ous kinds of vessels and gear. The special requirements of handling and processing seafood,
and the problems with improper handing and naturally toxic seafood are discussed. The large
nonfood (industrial or reduction) fisheries are documented here. Major fishing nations of the
world based on weights of commercial landings are reviewed together with the characteristics
that contribute to their emphasis on fishing. The potential production of world fisheries is
reviewed. Sea farming makes a contribution to the world's food supply; although presently
small in comparison to that of the capture fisheries, it is predicted to continue to grow. The
reasons for this, typical species reared, and problems in the industry are discussed in this part.
In Part Four, Fisheries Management and Regulations, basic yield models are given to in-
troduce the student to theory on population dynamics, stock assessment, and management.
Advanced models can be found in the references at the end of the book. Management by en-
vironmental manipulations and regulations are discussed, and examples of management
plans for shellfish, finfish, mammals, and reptiles are given. While the book is worldwide in
scope, emphasis is on U.S. fisheries. Management problems and techniques, however, are
similar in other fisheries around the world. Objectives of biological and economic fisheries

x Preface

management are discussed together with forecasting abundance as a service to industry and
to prevent overfishing. The last chapter on commercial fisheries discusses the future of com-
mercial fisheries of the world.
Part Five, Recreational Fisheries, profiles recreational fishermen and gives a history of this
type of fishing, the living resources they seek, gear used, and value of recreational fishing.
Management of recreational fishermen is more varied than commercial fishermen because of
the different objectives of the two groups. Yet both groups have to be considered in many
management plans where both fisheries are actively harvesting from the same stocks of fish.
The future of their sport is reviewed. The recreational section is shorter than the commercial
fisheries section because more historical data and records are available for commercial fisher-
ies worldwide. Also, many aspects of both kinds of fisheries are similar: species fished, re-
search methods, and aspects of management. Therefore, once the general aspects of
commercial fishing have been discussed, including oceanography and ocean productivity,
only distinctive differences are covered in the recreational part.
I have included in this broad subject called "fisheries," substantial reference lists to direct
the readers to additional information. Instructors using this book as a course text can pro-
vide updating of important current problems as information becomes available. It is, after
all, a broad field of knowledge with many important and rapid changes taking place in to-
day's fisheries.
This book is being published during a time of very considerable turmoil in the history of
fisheries management. The decimated condition of numerous fisheries worldwide is ex-
tremely serious, unprecedented, and threatens the economy of many coastal states and the
livelihood of their inhabitants. Developed countries are in the worst position, due for the most
part to overfishing and coastal pollution. In the late 1970s and the 1980s numerous symptoms
of fishing failures became much more evident and the scientific and popular literature warned
of the dangers ahead. Extended jurisdiction seaward to 200 miles in U.s. fisheries began in
1977, but unfortunately did little to reduce overfishing. Complete moratoriums on fishing
have been a necessary recourse in fisheries where catches have been dropping sharply for
years and there is no hope in sight. The current management thinking incorporates the con-
cept of dropping open access fisheries and replacing them with ownership or property rights
such as individual transferable quotas. This is much more complicated than it sounds, but
fisheries selling shares or certificates to fishermen in some countries suggest the concept may
be a good replacement for fishery scientists and administrators with an "open access fisheries"
mind set. There is an urgent need to learn from past mistakes and take action now!
To my wife, Jane, I owe a debt of gratitude for the arduous task of editing very rough
drafts, consistency of style, proofreading, and other numerous tasks in preparing drafts of the
manuscript. My son, E. S. Iversen, Jr., with great patience and care and numerous lessons,
convinced me that computers are a valuable tool in today's world. Kay Hale, Helen Albert-
son, and other library staff members of the University of Miami, Rosenstiel School of Marine
and Atmospheric Science were most helpful in assisting me in locating and obtaining
references. Many of my colleagues at the University contributed advice and information for
this volume. Carl Sindermann provided valuable suggestions on the layout of the book. I
would be remiss if I did not acknowledge the comments and suggestions from the many stu-
dents who used early typed versions of sections of this book as a study guide in my course
Marine Science 310, Living Marine Resources, Their Exploitation and Management, and my
Marine Biology and Fisheries course 509, Introduction to Fisheries.

Edwin S. Iversen

The bulk of the world's fish and shellfish is caught in marine waters. For example, about
90% of the world's catch of ca. 90 million (mt) came from marine waters in 1991. The great
variety of fish and shellfish species in the sea with their numerous shapes, large sizes at-
tained, and frequently bright colors are attractive to consumers and recreational fishermen.
Because of this variety of species and habitats available in the sea and consumer preferences,
many kinds of fishing gear and management plans are required.
The contribution fishing makes to a nation's economy varies, but in most countries it is
small compared to the total national economy. In large developed countries, the value of
landing represents no more than a fraction of a percent of national income. Naturally, in
small developed countries with long coast lines and well-established fisheries-for example,
Norway-fishing is of great importance. On the other hand, in developing countries, as in
the Far East where there is little heavy industry, fishing can generate considerable income,
and their fisheries can provide a cushion against famine when land crops fail.
The moral and social aspects of the challenge of hunger in the world today require the
need for resources of all nations to be effectively managed to reduce or avoid waste and
thereby help to reduce world hunger and starvation. The many living resources of the sea
are one source that can assist in this most worthy cause if we understand them and treat
them properly. Unfortunately we have failed miserably in many management attempts.
There is no doubt that hunger abounds in the world. Modern medicine has lengthened
human life the world over, but especially in the developing countries. This fact, together
with limits on food production and distribution, and natural disasters that reduce food pro-
duction, such as droughts, freezes, long-term climatic changes, and floods cause grave con-
cern that mass starvation in some parts of the world is imminent. Doomsayers and
forecasters state that an acute food shortage exists already and that it will worsen. Twenty
percent of the world faces actual starvation, beyond hunger or malnutrition. The global food
reserve 20 years ago, in 1974, was said to be a meager 30 day supply. To underscore the se-
verity of this problem, in November 1974, the United Nations deemed the world food situa-
tion so critical that it met in Rome in a World Food Conference to consider ways to increase
food production and improve distribution.
One must understand at the start that the problems in feeding the world will not be re-
solved by increasing the production of food from the sea alone. The sea can produce a share
of the food for the world and can be a godsend, especially when land crops fail for one rea-
son or another. But the fisheries are absolutely limited in production, as will be explained
later. Sea farming also must be viewed in the proper light, not as an answer to world food

xii Introduction

shortages (as has been expressed by some sea farming proponents), but as another source of
food production. The time has long since passed when unrestricted harvest from the sea was
strongly encouraged, yes, even allowed.
It is important to learn some definitions and to keep them in mind as we go through the
text. Terms on the first few pages are very important to the readers so they are described in
some detail.
A renewable resource pertains to living sources of supply able to reproduce and hence
perpetuate themselves.
Conservation is the wise utilization, intelligent management, or use of any resource; for
example, a portion of the animals (renewable resources) can be taken from a population with-
out harming it. This is usually referred to as surplus production. Conservation does not
mean simply not using a resource.
Common property resources are resources held jointly by a community. During the Mid-
dle Ages village commons were well-established. They consisted of a tract of land that could
be used by every member of the village to graze their horses or cows. As the need for keep-
ing these animals declined due to delivery of milk and better forms of transport, the com-
mons fell into disuse. Without management they became run down and were an eyesore in
the village. This is called the ''Tragedy of the Commons," and is exemplified by the state-
ment, "that which belongs to everyone is taken care of by no one."
Res nullis is a term from Roman law used to describe fish in the sea. It means things that
belong to no one. The concept that fish in freshwater and the sea belong to no one until
caught also extends to land animals as well. There is a famous law case about two hunters
where the first hunter wounds a fox that is later shot and killed by a second hunter. Owner-
ship was contested, but the fox went to the second hunter. The notion of common property
is much more involved than this example shows, but the principle has a highly important
impact on fishing rights and the regulations made by world nations.
Many people consider fishing to be a God-given right or privilege that no one can take
away from them. This is a false notion because equal rights or common property are not true
in fact; only certain people are allowed to fish in many fisheries (license restrictions, limited
entry). The terms "community property," or "public owned," have been suggested to avoid
these misconceptions resulting from the use of the term "common property." As a result, the
200 mile limit permits individual nations to restrict any other nations from fishing in their
territorial waters.
A stock is usually considered to be a manageable unit of an exploited population of fish
or shellfish, sometimes a single interbreeding population or group of fishes.
A population is difficult to define, but in this book it means an actual biological unit, and
includes terms such as races, subpopulations, etc., that will be dealt with in more detail in
later sections.
Fish and shellfish are both edible resources, but fish means finfish (vertebrates), and shell-
fish means aquatic invertebrates, not just having shells like clams, but sea urchins, shrimp,
crabs, etc.
Fisheries scientists are people who study living resources of freshwater, brackish water,
and the oceans with the objective of obtaining the greatest sustained return or harvest, either
from a commercial or recreational standpoint. Marine fishery biologists differ from marine
biologists in that marine biologists study any marine organism for its own sake, and their
findings may have absolutely no application to man's quest for food from the sea. On the
other hand, the fishery biologist studies aspects of the biology of marine organisms that are
exploited (harvested) by man for commercial value or recreational purposes. When we discuss
Introduction xiii

sampling by fishery biologists in

subsequent chapters, we will see that
their expertise goes beyond that of
the marine biologist in that they put
their knowledge to practical use
within a fishery that provides large
samples from which they can obtain
data. These data are costly and time
consuming to obtain by one's own
effort; research vessels and funds are
usually underwritten by govern-
ments. Sampling by fishery biolo-
gists carries with it a strict
responsibility to understand how,
where, and when the fishermen ob-
tain the catches the scientists will
Fishery biologists sort a bottom trawl catch to collect
use. Although some authors are now biological data on crab and fish populations during an
using "fishers" in lieu of fishermen Alaska Fisheries Science Center's eastern Bering Sea crab-
to include women who fish, I use the groundfish assessment survey. Photo Terry Sample, Alaska
time-honored term "fishermen" with Fisheries Science Center, NMFS, Resource Assessment and
the understanding it includes both Conservation Engineering division (RACE).
Fishery biology is defined by the Food and Agriculture Organization of the United Na-
tions (FAa) as follows:
A specialization of biology (more especially ecology) applied to groups of aquatic organisms
which are of actual or potential economic importance to man, and modified (from pure bi-
ology) by its employment of certain biological data. It is concerned with the identification
of natural units of stock of these organisms, with the elucidation of migratory, feeding and
reproductive habits and of growth and mortality rates, and with the measurement of popu-
lation levels including measurement and analysis of fluctuations of these levels and the ef-
fect upon them of fishing operations; it aims at the formulation of programs for the effective
utilization of these resources.

There are many other fishery professionals: several examples are fishery engineers, gear
technologists, fishery economists, fishery managers, and fishery lawyers.

Fisheries: Education and Employment

Fisheries science, as a profession, is relatively young. Fish and wildlife education programs
began in the United States during the 1930s. Over the years, numerous educational programs
at academic institutions have been designed and implemented to satisfy the increasing need
for trained fisheries scientists. Some programs are found in a department of fisheries; other
fisheries programs, or courses, come under a broader field, such as wildlife management, ma-
rine biology and fisheries, agriculture, forestry, or zoology. Some educational programs offer
an Associate Degree, others the B.S., the M.S., and the Ph.D. Recently, certain schools initiated
B.A. degree programs in "marine affairs," a survey of the science that provides a limited and
less scientifically oriented practical approach that fits into nonscience fields such as law, etc.,
where some basic fisheries knowledge is essential.
xiv Introduction

Fisheries graduates participate in research at governmental agencies, or academic institu-

tions, and/or teach at the latter. Private consulting has provided additional openings for
graduates in recent years. In some cases, they enter the fishing industry or related fields.
Educational training for fisheries has been reviewed and examined to determine if stu-
dents are being adequately prepared for their career goals. In the United States, leaders in
the fisheries field and professional fisheries organizations, through committee reports such as
The American Fisheries Society Professional Educational Standards Committee Report (Au-
gust 1977), have evaluated educational programs designed to train fisheries scientists. The
conclusion of many fishery scientists was that the education system at that time was not sen-
sitive to the needs of their graduates and was becoming inadequate. While each report eval-
uating fisheries training programs differed slightly, some opinions are common to many of
the more recent (1980s) programs.
The first criticism leveled at current programs is
the length of the curriculum. Most authorities point
out that in order to properly train a fishery scientist,
4 years of university / college training is inadequate,
and suggest that a 5 year undergraduate curriculum
may be necessary. Opponents of a 5 year program
say that the high costs of college education, rela-
tively few scholarships and fellowships available,
and jobs for fisheries graduates are often limited to
governmental agencies, mostly state conservation
agencies. Private industry (engineering firms, busi-
ness corporations, etc.) offers higher pay to gradu-
ates from 4 year programs than governmental
agencies do. So, while trying to strengthen fisheries
educational programs by stricter requirements or
additional courses, prospective students question
whether the effort to struggle through school is
worth the employment opportunities that may
await them upon graduation.
Along this same line, some faculty members be-
lieve, because of the increased complexity of the
profession, that 5 year undergraduate fisheries
training is not a rational option. At the undergrad-
Size composition and biological data are uate level students should receive a broad founda-
collected from tanner crabs during the tion in the sciences and humanities with only an
Alaska Fisheries Center's eastern Bering Sea introduction to fisheries. Specialization in fisheries
crab-groundfish survey. Photo Terry
science and management should take place at the
Sample, Alaska Fisheries Science Center,
NMFS, Resource Assessment and
graduate level.
Conservation Engineering division (RACE). A second criticism of fisheries educational pro-
grams is that they are too heavy on biology and
fisheries courses, being designed to manage the resource and overlooking political aspects.
The suggested remedy for this deficiency is for students to take social science or humanities
courses that unfortunately lengthen their college program. In addition, students should ob-
tain on the job training prior to graduation.
A third problem area results when students doing graduate research normally aggressively
compete with fellow students. Also, graduates' research projects tend to isolate them from
Introduction xv

their colleagues, so that upon graduation, when they become part of a team requiring close
group participation and cooperation, they may be ill prepared for important interaction.
Changes in technology, greater complexity in fisheries sciences, and exponential increases
in available information (e.g., mathematics and computer applications) require improvements
in training of fisheries scientists, meanwhile adhering to traditional studies and classic re-
search of pioneers in the field. Educators must be able to anticipate new requirements and
modify their courses and programs accordingly to meet new objectives.

Fisheries Literature
There is an obvious need for a book with a broad overview on fisheries subjects and special-
ized up to date information, to avoid repeating earlier studies and maximize research find-
ings by introducing the latest techniques. Today, fisheries students, scientists, and
administrators may take for granted the wealth of information on many fish species and fish-
eries available to them. But it has taken many years, conscientious effort, and many expen-
sive research programs to reach where we are today.
Virtually no reliable information sources on fisheries biology and management were avail-
able in scientific journals a few decades ago. This was partly due to the lack of scientific
research. There was such a paucity in the body of fisheries information, in so many aspects,
that no specialized journals for publication were available.
The formation of the Marine Biological As-
sociation at Plymouth, England in 1884 pro- The logarithmic increase in
vided encouragement to biologists to study number of scientific and
fish as populations rather than as individual 5,000 technical journals in the
fish. The Association's Journal provided impe- USA, doubling every
4,000 15-20 years
tus for research results. The International II)

Council for the Exploration of the Sea began in c:
Sweden in 1902 and encouraged the study of ~ 3,000
fish populations and the publication of results.
The U.S. Fish Commission, formed in 1871, be- 2,000
gan publishing results of federally sponsored 1,000
studies. The development of fisheries sciences
as a discipline is discussed fully in Chapter 4.
Since 1956, when Graham's book Sea Fisher- 1849 69 '89 1909 '29 '49 '69
ies was published in England, many good qual- Year
ity fisheries books have followed. However,
Scientific literature, including marine biology
most of these, like Graham's, were based on and fisheries, has grown at a logarithmic rate.
data on fish and fisheries in the North Sea. Numbers of scientific and technical journals in
Many of them are now out of print. While the the United States from 1849 to 1969. From
management principles espoused in these Maclean (1988). The ICLARM Quarterly,
books have rather wide application in other January.
temperate water fisheries, their contents tend
to have relatively localized interest. Many fisheries books subsequently published are di-
rected at mathematics of population management and do not provide adequate introductory
descriptive material on fisheries biology, ocean environment, and methods of exploitation of
fish stocks. Therefore, the beginning student cannot appreciate the need for management of
heavily fished stocks and methods used to attempt to maximize production from these stocks
on a sustained basis.
xvi Introduction

A wide range of quality exists in the literature today because of the policy of the journals
(peer-reviewed/non-peer-reviewed), or the ability and experience of editors (volunteer assist-
ant editors), and reviewers. Other information sources below the peer reviewed journals such
as popular periodicals, progress reports, summaries of symposia, and meetings can be found.
These sources sometimes called "gray literature," should be used with care because some of
the papers included in them may be preliminary or prepared hurriedly for a deadline.
Bibliographic access information systems available through computer systems are to varied
and technical to be explained here. One must learn by doing. If you are specific in what you are
searching for (key words) and have access to a quality library, the process is very useful. Most
librarians today can save researchers very considerable time and effort in computer searches by
suggesting the most useful data base for their needs. Some are only "in house," and other give
national or international coverage of the literature. Charges are made for this service by some
systems, but considering the time saved in literature searches, the money is well spent.
Many of the measurement conversions (U.S. to metric) used in this book are approximate
or rounded to whole numbers, for example, habitat depths, weights and lengths of organ-
isms, and fishery landings. Measurements at sea are given in nautical miles.

Fisheries Education and Employment

The appendix contains a list of selected General References together with journals and peri-
odicals useful to the study of fisheries.
Chapman, D. G. 1979. Fisheries education as viewed from inside. Fisheries 4(2):18-21.
Lackey, R. T. 1979. Fisheries education in the 1980s: The issues. Fisheries 4(2):16-17.
Oglesby, R. T. and C. C. Krueger. 1989. Undergraduate fisheries education: Technical specialization or
broad foundation? Fisheries 14(5):17-21.
Paulik, G. J. 1968. Fisheries education: A critical review, and a look at future programs. Univ. of Wash-
ington Publ. in Fisheries. New Series 4:295-299.
Royce, W. F. 1972. Undergraduate education of fishery scientists. Fish. Bull., 70:681-69l.
Royce, W. F. 1984. A professional education for fishery scientists. Fisheries 9(3):12-17.
Royce, W. F. 1984. Introduction to the practice of fisheries science. Academic Press, New York.
Sea Technology. 1992. Buyers guide directory, 1994-95. Compass Publications, Inc., Arlington, VA. (Pag-
ination not continuous). Section E-Educational Institutions.
UNESCO (United Nations Educational, Scientific and Cultural Organization), 1981. Fishery science
teaching at the university level. Report of a workshop on university curricula in fishery science.
May 5-8, 1980, Paris, France. UNESCO Reports in Marine Sciences 15. 75 pp.
U.S. Navy. 1984. University curricula in oceanography and related fields. National Oceanographic Of-
fice, Bay St. Louis, MO, NSTL, MI. 45 pp.

Fisheries Literature

Carlander, K. D. Fisheries education and training. Benson, N. G. (ed.). A century of fisheries in North
America. Special Publication No.7. Am. Fish. Soc. pp. 57-69.
Maclean, J. L. 1988. The growth of fisheries literature. NAGA. The ICLARM Quarterly. 11(1): 3--4.
McHugh, J.L. 1970. Trends in fishery research. Benson, N. G. (ed.). A century of fisheries in North Amer-
ica. Special Publication No.7. Am. Fish. Soc. pp. 25-26.

Living Marine

Their Habitats and Fisheries

Chapter 1

Ocean Environment

Ocean conditions control both the occurrence and abundance of commercial and recreation
fish and shellfish. To understand and follow these conditions, fishery scientists access ocea-
nographic studies to more efficiently harvest the sea's living resources.
About 70% of earth's area is covered by water. This can be divided roughly into three ar-
eas: lagoons, bays, sounds, and estuaries along the shore; continental shelves, offshore and
adjacent to land masses; shelves that drop sharply to a depth of 2 mi (1,733 fathoms), deep
sea basins that constitute the remaining area.


Estuaries are prominent and important features. They are bodies of water bordered by,
and partly isolated from, the ocean processes by land masses that were shaped by nonmarine
forces. An important characteristic of estuaries is that they are usually perpendicular to the
coast line. Many occupy drowned river mouths and stream valleys.
Estuaries are hard to completely define. If 20 biologists were asked to define an estuary,
there would be 20 different responses. Each biologist, however, would stress the fact that an
estuary is more complex than either a freshwater or a seawater environment because it is a
transition zone between the two. One simple definition states says that estuaries are places
where rivers flow into the sea and measurably dilute seawater. Generally, estuaries are in
bays or semienclosed areas. Many situations do not fit this definition, but for our purposes
this simplification should serve.
One of the most prominent characteristics of the estuary is the dynamic nature of the proc-
esses taking place. For example, the temperature, salinity, and pH (acid-alkalinity balance)
change markedly and quickly. In a long, narrow estuary significant differences are found in
the physical and chemical characteristics of the water from one end to the other. At the end
fed by freshwater, salinity is low and the temperature reflects its freshwater supply; on the
seaward end, the physical and chemical characteristics of the estuary more nearly resemble
the sea. The fauna living in the long, narrow estuary will vary throughout, and the differ-
ence can be quite pronounced, especially if it receives a high out-flow of freshwater. Con-
versely, estuaries that receive limited freshwater may have salinities so high that only a few
freshwater species can survive.
An important estuarine characteristic is the extremely high biological productivity. The
water in estuaries is brackish; that is, it is a mix of freshwater and saltwater having salinity

4 Part One Living Resources

values ranging from approximately 0.50 to 17.00 parts/thousand (ppt). Just as it is difficult
to obtain a clear-cut definition of estuaries, it is difficult to establish accurate figures on
whether or how much more productive estuaries are than the coastal sea, the upland areas,
or the entering freshwater. Many attempts have been made to measure estuarine productiv-
ity, but the measurement methods were not standardized or applicable in all areas. Notwith-
standing these obstacles, estuaries and tidal marshes are regarded as among the most fertile
natural areas in the world. Rates as high as 20 times as productive as the open sea, and
seven times as productive as the "average" wheat field have been estimated. However, high
fertility of brackish water does not mean that everything produced is usable by humans, but
rather that the weight of all organisms produced per unit area per year is greater than that
produced from other aquatic locations.
Life in estuaries is tenuous. Their inhabitants do not get added benefits of abundant food,
phosphorus, and nitrogen without sacrifice. They must be uniquely adapted to withstand the
wide and rapid fluctuations of physical and chemical factors mentioned above.
When floods occur in rivers feeding into estuaries, the salinity can drop sharply and stay
low for a sufficiently long time to cause a high mortality of many animals, especially the ses-
sile ones. An example of flood damage occurred in the Gulf of Mexico near the mouth of the
Mississippi River, where salinity was so reduced by freshwater that disastrous mortality to
oysters resulted.
Estuaries can be beneficial to fish and shellfish because they allow them to escape from
predators or parasites. Certain fish and shellfish can survive in lower salinities than can their
enemies. Continued low salinity prevailing for long periods will kill starfish, notorious pred-
ators on shellfish, and may prevent the spread of some shellfish diseases, such as fungus in
oysters, because the fungus cannot survive in low salinities.
Estuarine biology is also controlled to a great extent by bottom configuration and substrate
materials. Whereas in the open sea, water characteristics are important in determining pro-
ductivity and species present, the bottom is relatively unimportant by reason of depth. In es-
tuaries, however, fish live in close contact with the bottom or are bottom dwellers. The
shallower the estuary, the more important the role of the bottom becomes. Muddy bottoms
are alive with microbes, algae, and higher animals, that, by complex reactions, release nitro-
gen, phosphorus, and other vital plant foods. These vital chemicals permit high production
not possible on sandy bottoms.
The mechanism of enrichment in estuaries by freshwater differs among areas. In most re-
gions, enriched drainage runoff from the land flows into the estuaries and causes high
productivity. Generally, out-flOwing freshwater helps to stir up or "cycle" nutrients on estu-
ary bottoms, thereby causing high production (Fig. 1.1).
The principal elements that photosynthetic plants, such as algae and sea grasses, need for
growth are carbon, hydrogen, sulfur, nitrogen, oxygen, and phosphorus. Hydrogen and oxy-
gen are widespread, and carbon is derived from carbon dioxide. Sulfur dissolved in seawater
in its oxidized form, is subject to rapid oxidation and is dominant in the environment be-
cause of sulfur bacteria in the mud. Nitrogen is another controlling factor in estuarine pro-
ductivity, but we do not know how much is needed to keep a good chemical balance, or how
much bacteria and algae need.

Lagoons generally have poorer exchange with the sea than estuaries because they are in-
termittently cut off by bars or barrier islands. As a rule, they are parallel to the coast. Be-
Chapter 1 Ocean Environment 5

.. .
Wg ... ~. . . . ..


. ... . . . :'" ..•

Figure 1.1 Two-layered estuarine circulation pattern in Chesapeake Bay. Zone of maximum turbidity
is on the left. From Boicourt (1992). Influences of circulation processes on dissolved oxygen in the
Chesapeake Bay. In Oxygen dynamics in the Chesapeake Bay: A synthesis of recent research, eds. D. E.
Smith, M. Leffler and G. Mackiernan, 7-59. Maryland Sea Grant Book, College Park, MD.

cause of poor water exchange, the physical and chemical parameters in lagoons are generally
more stable and extreme than in estuaries. They are commonly shallower than estuaries and
may lie across the mouths of one or more streams. Because lagoons grow along the long axis
they may receive sand from over the bar by storm waves, or through an inlet. Fine sedi-
ments that occur in many lagoons may be stream derived. As is apparent from comparing
the definitions of estuaries and lagoons, there is considerable overlap. Few cases of clear-cut
examples of each can be found. Extremes of salinity are characteristic of certain lagoons.
Few species are adapted to hypersalinity. The more common species are copepods, polycha-
etes, mollusks, brine shrimp, and usually a few finfish species. Mullets are frequently abun-
dant enough in a lagoon for harvest by humans. Lagoons with more moderate salinity
support a wider variety of fish and crustaceans and therefore are fished heavily by subsist-
ence fishermen, particularly in developing countries.


Bays are defined as that part of a sea that indents the shoreline, usually having a wide in-
let, but not as large as a gulf. They are important features of the edge of the sea that support
a variety of fish and shellfish of commercial and recreational value. Because bays are often
protected from heavy weather, they provide a livelihood for commercial fishermen who use
small vessels. Unfortunately, these areas are often convenient dumping grounds for pollut-
ants that are not only harmful to aquatic organisms, but also can make edible fish harmful to
In North America one does not have to look far to see bays that are important to the fish-
ing industry. Chesapeake Bay is an excellent example of a water habitat with special features
that support many species in abundance. This drowned river (it can also classify as an estu-
ary) cuts into the states of Virginia and Maryland for about 200 mi (322 km) and is between
4 and 40 mi (6.4-64 km) wide. In good years millions of pounds of shellfish, especially oys-
ters, and a wide variety of finfish, such as the Atlantic salmon, are landed along with millions
6 Part One Living Resources

of tons of fish for reduction purposes. Many bays around the world may be estuaries, yet in
popular terms they are called bays just as Chesapeake Bay is. The Bay of Fundy in Canada,
well-known for its spectacular tide range, supports a wide variety of valuable cold water
species. Some bays are important to fisheries because they provide both near shore fishing
and harbors for fishing vessels. Noteworthy bays are the Bay of Biscay in southwest Europe,
and the Bay of Bengal between India and Burma.


A sound is defined as a wide channel or strait that links two large bodies of water, some-
times separating an island from the sea, or as a long inlet or arm of the sea. Puget Sound in
northwestern Washington State is large, about 125 rni (201 km) long and between 5 and 25 mi
(8-40 km) wide with large estuarine flats and subtidal basins that support a wide variety of
essentially nonmigratory but commercially important shellfish like oysters and clams, and
finfish. Species of Pacific salmon support important commercial and recreational fisheries.


Continental Shelves

The continental shelf slopes gently away from the shore to a depth of about 500 ft (83
fathoms). Due to the gentle slope, the outer margins may extend seaward to about 800 mi
(1.482 km) from shore, as is the case off Siberia. Shelves are not simply flat plains, but are of-
ten cut by ancient river beds, lagoons, and beaches. As a general rule, off young mountain-
ous shores, for example the Pacific coast of North America, the continental shelf is usually
narrow, not much more than 20 mi (37 km) wide, or nonexistent. Conversely, off land masses
with flat plains, the continental shelf is usually wide (Fig. 1.2). Siberia's wide continental
shelf is of little value to fisheries because it is ice covered year-around. On the other hand,
the shelf on the east coast of North America north of Cape Hatteras extending seaward for
about 150 mi (278 km) is highly productive. The shelves of greatest importance to fisheries
are off northwest Europe, eastern Asia, Argentina, Alaska, and the east coast of North
America. In Chapter 10, the paramount importance of continental shelves to total world fish
production is explained.

Coral Reefs

Another feature of the oceans important to fishery scientists and commercial and recrea-
tional fishermen alike are coral reefs. These marvels of nature occur in shallow depths hav-
ing warm temperatures of 77-86°F (25 to 30°C) and lie in a 3,000 mi (5556 km) wide belt
around the Equator at latitudes seldom more than 22°. Many tropical areas seemingly suita-
ble for reef formation do not support coral reefs because rigid conditions necessary for reef
formation are not met. Reefs are composed of consolidated limestone debris that provides
support for corals, mollusks, and algae. The death of these organisms provides additional
support for successive generations and increases the reef's size. Reef growth is generally up-
ward until it reaches the surface, when growth proceeds laterally.
Most coral reefs fall into three general categories:

1. Barrier reefs occur offshore with a channel behind them. The Great Barrier Reef of Australia,
about 1,200 mi (2,222 km) long, is an outstanding example of a barrier reef.
Chapter 1 Ocean Environment 7


Shelf edge
70 '" (13' m)
Continental slope

~ 1."'-'.""'.)
2,1!Z fin(4,OOOm)

Figure 1.2 Schematic profile of the continental - Continental terrace~ I'-- Continental rise - _ Deep sea bed _
margin. Modified from Berryhill (1974). The Continental margin
world wide search for petroleum offshore-
a status report for the quarter century, 1947
-1972. U.S. Department of the Interior.

2. Fringing reefs normally occur in shallow water and tend to increase toward the open ocean
where favorable water conditions prevail. Examples are along the Florida Keys.
3. Atolls are circular or horseshoe-shaped reefs, rising from the bottom of the sea generally miles
from any continent. Christmas Island in the Line Islands is one of the largest atolls in the world
and has a shallow interior lagoon, a characteristic of atolls.

Fish living on and around coral reefs are numerous and represent the greatest variety of
species found in a single marine habitat. Small scale fisheries exist on reefs but many of the
fish are too small for use as human food. Butterfly fish, damsels, surgeons, and wrasses, ex-
ceptionally colorful finfish species, are the main groups on coral reefs. These are often seen
in aquaria. All find food in the plant materials on the reefs. A few large predatory species
such as groupers, amberjacks, and barracudas are exceptions to the rule that most reef fish
are small and delicate. Some reef-dwelling species leave the reef to feed in seagrass beds or
mangrove swamps, and others live as juveniles off reefs and later enter the reef community
as adults.
Estimates of maximum sustained yields from coral reefs have been made for finfishes liv-
ing on or near the bottom (demersal) in flowing waters (neritic) and range from 2.5 to 5
tons/O.4 mi 2 (km 2)/yr, less than 10% of the total annual fish catch from all oceans. In areas
having a dense cover of actively growing corals, large fish harvests can be expected. Inverte-
brates including spiny lobster, and conch are often present in sufficient quantities to support
small fisheries. Due to the nature of coral reefs, hook and line, traps, and spear fishing are
mostly used to capture fish. Occasionally seines and gill nets are employed.

Continental Slopes

Upon leaving the continental shelf and moving down the continental slope, pressure and
darkness increase, forming an environment unable to support plant life. The average height
of a slope is 12,000 ft (2,000 fathoms), but 30,000 ft (5,000 fathoms) has been measured, far
greater than land escarpments. In addition to their towering height, they exhibit dramatic
geomorphology in the form of submarine canyons with extensive valleys whose origin is not
8 Part One Living Resources

well established. One theory suggests mud flows could have carved out these canyons.
Spectacular terraces also exist on some slopes. Due to the absence of plants, slope animals
are carnivorous and make their living by preying on other slope animals. The continental
slope is of little importance to most fisheries because conventional fishing gear cannot be op-
erated there, and fish and shellfish populations are substantially smaller than those on the

Deep Ocean
It is estimated that the deep ocean floor that begins at the base of the continental slope has
an area of about one-half of the earth's surface. The use of the term "ocean floor" conjures
up an image of a flat level plain. True, some of the more spectacular topographic features
were known to exist many years ago, such as the Mid-Atlantic Ridge and the exceedingly
deep Mindanao Trench (Philippines). But the uneven, rugged character of the bottom came
to light through the results of the 1947 Swedish Deep Sea Expedition. Using fathometers, the
scientists when covering only a few miles, found the ocean bottom to rise and fall. In con-
trast, the Indian Ocean was found to be quite level over distances covering several hundred
Because of its great depth, averaging about 12,500 ft (3,800 m), the majority of the slope
and the ocean bottom are of little importance to fishermen because they are out of reach of
fishing gear. Also the populations of potentially commercial or recreational fish are very
Early attempts to map the seafloor required tedious soundings using a large lead and a
long sounding line. At each station the vessel stopped, the crew located its geographic posi-
tion as best it could, then lowered the lead to the bottom and measured the depth at that
point on a calibrated line. At many stations the vessel could not anchor, and moderate to
heavy seas made accurate measurements difficult to obtain. Since then, electronics (echo
soundings) provided a much better view of the seafloor, but vessels needed to be fitted with
expensive sounding equipment and to travel endless miles to obtain depth measurements at
countless locations to provide the data for nautical maps.
A new technique, SEASAT satellite data (Fig. 1.3), has revolutionized not only mapping
the sea bottom but shows features important to fish distribution as well as areas where fish-
ing cannot be carried out, such as seamounts, ridges, rifts, and fracture zones. In one study,
over 1,000 orbits around the earth were used. Rather than actually mapping the sea floor,
SEASAT measures variations in the height of the ocean surface reflecting variations in the
earth's gravity field, that in turn shows the variable heights of the seafloor. A rise in sea sur-
face can be as much as 2-10 ft (0.6-3 m) in the vicinity of a mile high seamount.

Seamounts, also called guyots, are former islands that have lost their summits. Once high
islands that were large submarine volcanoes, they rise 3,280 ft (1,000 m) or more from the
seafloor, and are flat with limited area on top. Some appear as low, sandy atolls; others have
subsided well below the sea surface. Their summits lack coral caps. Many of those with sub-
merged summits were discovered after electronic bottom sounders came into wider use on
ocean vessels.
Seamounts are numerous in the central and western North Pacific Ocean. They are of in-
terest to fisheries scientists because commercial fish frequently accumulate on their summits.
The Hawaiian Archipelago and numerous seamounts to the north and northwest of the chain
Chapter 1 Ocean Environment 9




Figure 1.3 Configuration of SEASAT-A. From
Australian Fisheries, October 1978. LINK ANTENNA

have been explored so far (Fig. 1.4). To the north, the Musicians Seamounts have summits of
mostly well over 5,414 ft (1,650 m) below the sea surface.
Not until 1967 did fishery scientists realize the potential fishery resources associated with
seamounts. The Russians were the first to commercially exploit the seamount groundfish
resources. The 200 mi Fishery Conservation Zone (FCZ) includes many submerged banks
and seamounts, including Hancock Seamounts (Northwestern Hawaiian Islands). Japanese
groundfish trawlers fished this area under quota from the United States. Total catch from
these seamounts has remained level despite increasing fishing effort, which suggests that the
rate of fishing may have surpassed the level of optimum fishing effort.


As mentioned earlier it is important for the fishery scientist to understand the physical and
chemical conditions for life in the oceans and how these are measured. The well-being of the
biota in the oceans depends on many parameters; however, we will discuss only a few of the
more important ones. Texts of general descriptive oceanography should be consulted for a
complete understanding of conditions of life in the various habitats in the sea.

The basic physical properties of seawater are primarily dependent upon salinity, tempera-
ture, and pressure (i.e. depth level). Salinity is required in computing density (weight per

, C'Jlngu Smt.
';OOjln Smt.
, q.
.. ~
?\,," c.~
.~>_,~" Klmmei Smt. :~)
... -
GMelliIh BcnI

:~"(:yurJGkU I :") ,.:, Ber,:~:2\ ;L I" olI9t Smt.
MILWAUKEE (.., • r·· ... '.' ."
SMT. GROUI'.j Kammu Smt.
I CoIahan Smt Kern Smt'.......
"" 1!,.\1\ r:::\ ':RaIeini
....... ;.. :.:'
Smt , lDonizetti Smt
,::~ ~~Smt.1 MahierSmt. :.~ 'j. ,::'.. ·~tSmt .
,.~Smt.14 Goddard Smt. . :::- "::.:::',.. --Brahms Smt. .
"-s.: ...... ·.
'-"""-'''f~.. .. o-ak Smt.
''...... "De Smt.

Smt.11,·ci··;=,''''Smt.10 RachmaninOff Smt. '. ':. .., 3o--J

oSlilt 15 ·········:;;~... L .•Ladd Sml I ''Tchailcavsky Smt.
. "~' .. \;,-,,~yl, I ." POQaninlSmt.:,::yszt Smt.
Nero Smt..-- :.,"-,:.PeilrI6 Her_ ,:.... .i " . --Mozart Smt.
Salman :2.";i';:~··':~l.~;
I ,'.smt 8
-.;.:-:'" ".' . .:,
Grl=~~>::::::.::: . . ; .' .,HandeI Smt. 114~~:;'\PIoneer Smt. ' .Voladar Smt. H Smt.:.,: :.'"., Glucli Smt,

I . ';j Smt.! ,. ::;:\,.. ~' ... ttiCfltftf. . .' ,.,
I. . RC!i.~o I!an!'. Chapin ~'.::;.., ..:'
r .: :. .: : =r(·r7:.:!Bank
. Banks B.~'. ..'.., : ,":',
~ _ _~
.:: Paul Smt. FrlS:";'~ >"" ;. ,,*ker I
Fr.Shoo , .,::.-< . ", j ':"'" . .,: .Nihaa I•
.::": t.::\ .,< :;.~::.Nec:ker -.Cha~ IOKauai Tuscolaaa
>. Ridc.Ie. Smt.· 'Nllhau Smt.
. . Wisconsin S m t . ' r'l.
() . .... '. "\...;)~alakOI.
. ... Tlfu Smt. lanai'\) <;c:>Maui

175°E ItIo- 1150 110" I~oW

Figure 1.4 Hawaiian Archipelago showing the numerous seamount chains and groups (SMT). From
Uchida and Tagami (1984). Mar. Fish. Rev. 46(2).
Chapter 1 Ocean Environment 11

unit volume), dynamic currents, and sound velocity. Density is important in its relationship
to salinity and ocean currents. The properties of temperature and dissolved oxygen, along
with salinity, enable physical oceanographers to plot water masses and to chart their
movements. Each specific area in the ocean has its own salinity range, and variations occur
depending on the locations in the world where the water mass is sampled. The salinity value
will thus give a clue to the age of the water in a given area: the higher the salinity, the
"older" the water.
Salinity is often referred to as the "saltiness" of the oceans, because the value is roughly
proportional to the amount of dissolved salts (solid material) present in a given volume of
seawater. This is not the same as "chlorinity," which is equal approximately to the amount
of chlorine in the water (or the amount of silver required to precipitate out all chlorides in a
sample of sea water). An empirical relationship exists between salinity and chlorinity that is
basic to chemical oceanography:

salinity = 0.03 + 1.805 x chlorinity

Both salinity and chlorinity are expressed in grams per kilogram of seawater, or parts of
dissolved solid material per 1000 parts of water (parts per thousand, ppt, symbolized %0).
The salinity range in the ocean is generally 33-37%0, and averages about 35%0 (or 3.5%) for
open sea areas. When seawater has been diluted, such as near the mouth of a river or after
a heavy rainfall and in colder climates where little evaporation occurs, the salinity may be
much less (down to 5%0 and lower). In hotter areas of the world, where excessive surface
evaporation takes place, and in special circumstances of confined saltwater bodies, as in la-
goons, the salinity may reach 40%0 and occasionally even higher. Chlorinity accounts for an
average of 19%0 in seawater.
One of the simplest devices for measuring the density and/or salinity of coastal seawater
is a calibrated, stem-type hydrometer. Hydrometers are especially designed floats that meas-
ure the density (specific gravity) of seawater, based on the principle that a floating body rides
higher in a heavy liquid than in a light one. The specific gravity is determined by noting the
depth to which the vertical scale on the stem of the hydrometer sinks when it floats free in a
water sample.
The most common type of hydrometer is made of glass, with a graduated stem attached
to a hollow bulb section. Below the flotation bulb is a smaller glass bulb filled with mercury
or lead shot weight to balance the instrument and make it float upright. The graduations
marked on the stem of the hydrometer indicate the desired measuring range and type of
scale used. One or more hydrometers may be needed to cover the normal seawater density
range. Because density varies with temperature and pressure, the standard practice is to
make corrections for these variables by taking the sample temperature and indicated specific
gravity readings, and then standardizing density values to 5g e F (15 e C). Once the standard
density is obtained this value may be converted directly into corresponding salinity and/or
chlorinity values by utilizing appropriate tables.
The classic method for determining salinity (amount of chloride ions present) is the Knud-
sen titration method. It gives the high accuracy normally required in the deep and midlayers
of the ocean where small salinity fluctuations are encountered. The method consists of add-
ing silver nitrate and potassium chromate as a color indicator to a seawater sample under
test; the amount of silver nitrate required to precipitate out all of the dissolved salts present
is converted into chlorinity units, and then salinity is computed by using the standard for-
mula relationship between salinity and chlorinity. It is tedious and impractical under certain
12 Part One Living Resources

field conditions (aboard vessels in heavy weather) where detailed profile measurements are
Because salinity is a measure of the ion content of saltwater, it may be determined directly
if the solution electrolytic conductivity (reciprocal of electrical resistance) and temperature are
both known; this variation of salinity with conductivity and temperature forms the basis of
measurements made by the modern-day electronic salinometer. Continuous measurements
are transmitted instantaneously to the surface operating station by a sensing mechanism as it
is raised and lowered, or towed, through the water. In some cases the instrumentation pro-
vides salinity as a direct, computed output and, in other cases conductivity and temperature
are supplied as separate outputs.

Seawater Temperatures

Throughout history, the most fundamental recorded characteristic of the ocean has been its
temperature, probably because this variable has been the easiest for people to record and un-
derstand (Fig. 1.5). And, even during the early studies of the oceans, biologists knew that
temperature greatly affected where aquatic animals could live and the rate at which they
grew. As human's sophistication in understanding the dynamics of the sea has grown, we
have learned to combine temperature data with salinity, pressure, and oxygen measurements
to provide a more accurate concept of the marine environment and the role it plays in the oc-
currence of marine organisms and their abundance.
There are generally three characteristic layers in any water mass profile. The first charac-
teristic layer is the surface layer, which reflects the ambient temperature and may fluctuate
dramatically. The second layer is referred to as the thermocline. This layer includes the zone
where temperature changes rapidly with depth, reflecting a transfer of heat both vertically
and horizontally from surface to deep water. The constant fluctuations indicate that mixing
of the water mass occurs regularly. The third characteristic layer is the deep water layer, or the
bottom water. The polar region origins of the water mass can be traced to this zone.
The centigrade or Celsius temperature scale, °C, is the standard used for scientific investi-
gations of the sea. Accuracy within 0.05°-O.01°C is usually required for adequate description
of middle and deep water layers because small changes in temperature have significant ef-
fects on salinity as well as density and other physical properties, and because extremely
small variations in temperatures are found at great depths. The degree of precision required
can be obtained only from well-constructed and carefully calibrated thermometers that are re-
checked periodically. Temperature conditions in the surface water layers vary over a wider
range than at intermediate and deep zones, and somewhat lower accuracy standards (0.5°-
O.I°C) for measurements are permissible.
Sea temperature is measured by a variety of methods, and accuracy has been improved by
the sophistication of the equipment used. Early data was collected by means of bucket ther-
mometers, or water drawn from below the surface by sensors pulled behind a moving
vessel. The Nansen reversing thermometer introduced over a century ago remains the most
reliable and accurate means of measuring sea temperature, particularly at levels below 900 ft
(274 m), and is used extensively (Fig. 1.6). The thermistor temperature probe, unlike the re-
versing thermometer, provides an instant, continuous record and does not require interpola-
tion or elaborate calibration equipment. Basically a semiconductor resistor, the probe
resistance changes as a function of temperature.
Chapter 1 Ocean Environment 13

W 'lO
,so_ 40
..r:: 100 _r-

200- x 4·59
Cl 3oOf- r- 87 BT 87
r- S
14OCl~ 40 -'0 eo 10

Modern temperature profiling system (expendable bathythermograph)


J l
An expendable temperature probe is Next, the probe is deployed from the When the probe reaches maximum
inserted into a launcher which is hard launcher over the side of the underway recording depth, the wire is broken and
wired to the displayed unit on the vessel's vessel, and the temperature depth profile the probe sinks to the bottom.
bridge. is relayed to the display on the bridge.

Figure 1.5 An old fashioned bathythermograph, grid for reading temperature tracing on smoked
slides, and a recording. From Holden and Raitt (1974). FAO Fisheries Technical Paper No. 115 Revision
1. The device measured about 32 in. (81 cm), weighed more than 20 lb (9 kg), and was awkward to use
in heavy seas. It has been replaced by a temperature probe that is easy to use, inexpensive, and
provides a temperature depth profile on the bridge. It is popular with fisherman and oceanographers
as well. Illustrations courtesy of Sparton Marine Products, Bainbridge Island, WA.
14 Part One Living Resources

11+-_ _ Auxiliary thermometer

iM---- Main thermometer

Figure 1.6 Early pressure protected reversing

thermometer. From Holden and Raitt (1974).
FAO Fisheries Technical Paper No. 115
Revision 1.

Dissolved Oxygen (DO)

In seawater, the amount and type of gaseous components are related to the solubility of
the particular gases in water. All are relevant to a number of oceanographic phenomena,
such as the accumulation of gases in the swim bladders of deep sea fish, the metabolic proc-
esses of marine organisms, deep sea currents, water turbulence, air-water-surface exchange,
etc. Knowledge of the gaseous components in water can be used in identifying water masses
or understanding the environment where chemical and biochemical reactions occur. DO in
water is the most often-measured dissolved gas. Levels of DO in sea farming facilities are
critical to successful operations.
Dissolved oxygen is of fundamental importance because it plays an active part in the me-
tabolism of organisms as well as in the formation and solution of lime, and in the rotting of
organic matter. DO is added to the sea in two basic ways: by absorption of air in the upper
layers and by photosynthesis of plants in a layer strictly limited by the depth of light
penetration. Conversely, DO can be lost from the sea at the surface interface by exchange
with the atmosphere, and it is consumed at all depths by the respiration of plants and ani-
mals plus the decomposition of organic material by bacteria.
The percentage of DO varies considerably from one body of water to another. Surface wa-
ters are normally saturated with DO, the content varying between 4.5 and 9.0 mg/L (depend-
ing on the salinity and temperature). The higher values correspond to the lower
Owing to convection, wave action, etc., the water layer held in equilibrium with the at-
mosphere is of variable but considerable thickness. Alteration in the amount of dissolved ox-
ygen (DO) below the surface layer is caused in part by currents. Photosynthetic activity of
Chapter 1 Ocean Environment 15

plants under the influence of light causes the extraction of carbonic acid and the delivery of
oxygen, a process that may ultimately lead to oxygen oversaturation. Because sunlight pen-
etrates only to a few dozen meters with sufficient intensity to permit photosynthesis, the "eu-
photic zone," where oxygen can increase by photosynthesis, forms a thin layer across the
ocean. At lower levels, two processes tend to reduce (rather than increase) the amount of free
oxygen: animals extract oxygen from the water in the process of respiration, giving off carbon
dioxide (C02 ); a similar exchange is produced by rotting organic matter. Also, because bio-
logical processes are different throughout the ocean due to unequal dispersement of plants
and animals, no accurate indicator for the age of water masses can be based on the amount
of DO present. Generally speaking, the longer a body of water has been withdrawn from
contact with the surface, the lower its DO content will be. Only the diffusion of a water mass
by turbulent motion (eddies, internal waves, and movements of swimming animals) can
bring in a supply of outside oxygen and preclude entire depletion. Because oxygen con-
sumption by organisms appears to be independent of the oxygen content, the slower a mass
of water moves over a certain distance, the more oxygen will be withdrawn from it, other
factors remaining equal.
The classical technique used to measure DO is the Winkler method. This wet chemical
technique employs a titrimetric procedure to derive either milligrams/liter or parts/million
(ppm) by weight DO. The method depends upon oxidation of manganese hydroxide by the
DO. When acid is added, the oxidized manganese reacts with potassium iodide and releases
iodine in amounts equivalent to the original DO content, determined by titration with so-
dium thiosulfate. Several modifications of the standard Winkler titration method have been
developed to alleviate the influence of various ions and compounds such as nitrite oxygen.
While this method is simple and inexpensive to use, it has major disadvantages. Wet
chemical procedures are tedious and time consuming to perform when there are many
samples. The accuracy of the readings is further affected by the subjective interpretation and
handling by each operator, commonly referred to as "human error."
Other problems associated with the measurement are that the DO concentration in seawa-
ter is influenced by the oxygen reacting with the brass linings of Nansen bottles during the
long time needed to haul the bottles from deep casts to the surface (Fig. 1.7). Plastic liners or
even all-plastic bottles are often used to overcome this problem. In addition, bacteria and
other biological organisms in the water sample can easily alter the DO content of the sample
if it is stored too long before analysis.
With the development of the polarographic oxygen sensor, all of the shortcomings of the
Winkler method are eliminated. The polarographic probe contains two electrodes connected
by an electrolyte, between which voltage is applied. When the sensor is in contact with the
sample, oxygen diffuses through a gas-permeable membrane and is reduced at the cathode
where a small current is generated proportional to the oxygen pressure. In short, it provides
greater accuracy, quicker readings, and permanent records not provided by the Winkler
method (Fig. 1.8).

Visibility in the Sea

Light is the basis for almost all plant activity in water. The distribution and production of
phytoplankton, minute rooted and suspended aquatic plants, are directly related to the quan-
tity of radiant energy penetrating the water. The abundance of phytoplankton in turn affects
the distribution and production of zooplankton and ultimately the production of fish. Light
also directly influences the distribution of these organisms.
16 Part One Living Resources

Figure 1.7 Reversing water bottle for use in series. The messenger trips the first bottle collecting a
water sample at that depth and releases the next messenger below to trip the next bottle attached at the
desired depth. Photo from the author.

The principal factors affecting the depth of light penetration in natural waters include sus-
pended microscopic plants and animals, suspended mineral particles such as mineral silt,
stains that impart a color, detergent foams, dense mats of floating and suspended debris, or
a combination of these. The region in which light intensity is adequate for photosynthesis is
often referred to as the trophogenic zone, the layer that encompasses 90% of the incident
light. The depth of the trophogenic (nutrition + suitable) zone may vary from less than 5 ft
to greater than 90 ft 0.52-27.4 m).
The physical relationships governing the penetration and absorption of light, color, and
transparency of natural water bodies are of prime importance to physical and biological stud-
ies in oceanography. Transparency or "clarity" of natural water bodies may be defined by
any of the following terms:

1. the percent of surface light present at a specified depth;

2. the percent of absorption of light per unit length of path;
3. transmission of light as a percentage of transmission through distilled water;
4. Secchi disc measure of water transparency: a flat disc marked with black and white paint that
is lowered with a calibrated line into a water body providing a reading of the depth as it dis-
appears from view, and, hence, the water transparency.

The degree of transparency, or "transparency index," varies with the number, size, and na-
ture of particles suspended in the water that, along with the water surface and the water
molecules themselves, affect the absorption and "scattering" of light. Scattering is the redi-
rection of light by suspended particles, usually smaller in size than most plankton. In dis-
tilled water, scattering is related to the molecular structure of the water. When a train of light
encounters particles (including water molecules) where dimensions are comparable to the
wavelength of the light transmitted, scattering occurs. Minute particles scatter "blue light"
most effectively; larger particles (size of chalk dust or larger) scatter all wavelengths equally
Chapter 1 Ocean Environment 17

Figure 1.8 Modern glass instrument housing with "hard hat" shell used to enclose an automatic radio
or radar transceiver (transponder) that transmits signals when triggered by an interrogating signal to
determine locations of underwater objects. Among other uses, transponders can also be used to study
ocean currents. Photo courtesy of Ferranti o.R.E., Inc., Falmouth, MA.
18 Part One Living Resources

"Absorption" is the process of reduction of light through its conversion into another form
of energy, heat. This is accomplished by suspended particles and/or water molecules. Ap-
proximately 50% of the total radiant energy is absorbed in the first 2 in (50 mm) layer of the
sea; about 60% is absorbed in the first 3 ft (1 m) of clean oceanic water, and much more in
coastal or turbid areas.
Light absorption in normal seawater (35%0 salinity) as well as coastal and inland water
bodies, does not significantly differ from that in distilled water. As far as true absorption is
concerned, most natural water bodies are identical with distilled water. In clear oceanic wa-
ter and offshore areas of large deep lakes, absorption predominates; but in turbid near-shore
waters, scattering is the dominating factor with absorption playing a somewhat less, but still
important, role in the process of light attenuation.
"Transmittance" of light through water is closely related to its color. The Forel/Ule scale
is used to obtain an approximate measurement of surface water color; it gives an indication
of the transparency of the water and also assists in classifying water masses as to gross bio-
logical activity. The Forel portion of the color-comparison scale consists of a series of 11 nu-
merically designated vials, each containing a fixed color solution, and the scale graduated in
shades from deep blue to intense green. The color graduations of the Forel scale correspond
to open seawater colors as they appear to an observer ashore or onboard a vesseL Blue wa-
ters are typical of the offshore zone, particularly in middle and lower latitudes, whereas the
green water is more common near the coastline. Turbid coastal and inland water bodies,
such as harbors, may vary in color from greenish yellow to brown; these color graduations
are covered by the Ule scale, also consisting of 11 color standards and similar in construction
to the Forel Scale.

The U.S. Naval Oceanographic Office defines turbidity as a condition of water clarity re-
sulting from the presence of suspended matter. It further states that water actually is consid-
ered turbid when its load of suspended matter is visibly conspicuous. Because all waters
contain some suspended matter they are all turbid by this definition.
Fish culturists have noted evidence of gill irritation in trout and salmon fingerlings held in
turbid water that can cause infection by fungi and pathogenic bacteria. Turbidity can reduce
the oxygen-holding capacity of water. In coastal waters suspended matter may settle out and
smother bottom-dwelling species. For the most part turbidity is not a problem in most oce-
anic areas.
Instruments or devices used to measure underwater visibility are the Secchi disc; the trans-
missometer (hydrophotometer), and the submarine photometer (irradiance meter). The most
common method of reporting turbidity is in Jackson Turbidity Units (JTU's) that are equal to
1 mg/L of turbidity as measured on a silica scale. Silica is used as a means to express tur-
bidity with 1 mg/L of Si02 equivalent to one unit of turbidity (1 JTV).

Marine Sedimentation
Marine sedimentation relates to the deposition, composition, classification, and structure of
organic and inorganic material from the ocean floor. Sediment analysis usually includes deter-
mination of size, shape, and percentage of component particles therein; identification of miner-
als and ratio of light to heavy minerals; wet density; pH; and calcium carbonate content.
Biological and ecological studies emphasize the animal population as well as the environmental
factors determined by temperature, depth, type of sediment, and geographic location.
Chapter 1 Ocean Environment 19

Almost every type of marine deposit, with the exception of those exposed to extreme dis-
turbance by waves or currents, is inhabited by an assortment of animals; some, the epifauna,
live on the surface; others, the infauna, burrow or lie buried beneath the surface where they
maintain contact with the overlying water by means of a tube or siphon. The depth to which
such animals burrow is usually within 12 in (30 cm) of the surface. Epifauna, which tend to
be active predators, are themselves protected from attack by means of their rapid movement,
their mechanical means of protection, or their inaccessibility. Infauna are often soft bodied,
relying on their burrowing habits to protect themselves from predators; nevertheless, they
form the chief food of many bottom-feeding fish.
Classification of bottom sediment is based on a combination of grain size and origin.
Therefore, it is possible to have samples composed of terrigenous material, subaerial or sub-
marine volcanic material, organic matter, inorganic material, and extraterrestrial matter. Size
of these component materials varies and is used as a further classification criterion. Oceanog-
raphers collect marine sediments using three basic types of samplers: dredges (drag buckets),
snappers, and coring tubes. When collecting rock samples from outcrops of solid rock,
dredges with chain mesh bags are used. Smaller cylindrical dredges with solid bottoms are
sometimes used to collect nonconsolidated material in relatively shallow water. Dredging
operations for bottom sediments usually are conducted where coring or grab sampling de-
vices have failed to obtain a sample. Snapper samplers (clamshell type) have been widely
used to obtain samples of superficial sediment layers. A disadvantage of this sampler is that
the contents tend to wash out while being hauled to the surface. This is particularly true
when the bottom is sandy or contains coarse fragments, because fragments caught in the jaws
may prevent them from closing completely.
Coring tubes are long tubes held vertically in weighted samplers that are driven into the
sediment by their own momentum (Fig. 1.9). Depth of penetration is determined by the
weight of the instrument (over 500 lb or 225 kg), character of the sediment, diameter of the
tube, and type of cutting nose. In areas of predominantly rock or coral bottoms, it may be
impossible to obtain a core.
Large areas of the ocean floor are covered by uncompacted deposits of mud, sand, or
gravel. The proportion occupied by rocky outcrops is relatively small and confined to limited
coastal areas, parts of the continental slope, submarine ridges, and canyons.

Density of Seawater
The density of seawater depends upon salinity, temperature, and pressure. At constant tem-
perature and pressure, density varies with salinity or, because of the relationship between salin-
ity and chlorinity, with the chlorinity. Salinity variation essentially involves a change in mass of
a given volume, thus increasing the density. Pressure increases in magnitude with depth (Fig.
1.10), and a unit volume of seawater carried from the surface down to some point beneath will
be compressed and occupy a slightly smaller volume; this results in an increase in density (if
the mass and temperature of the same remain unchanged). Unlike pressure, temperature
changes cause a direct variation in volume of a water sample; an increase in temperature results
in an increase in volume and a decrease in density, if other factors remain constant.
Normally, waters of different densities occur in nearly horizontal layers, with the lightest
water at the surface and the heaviest (densest) water at the bottom. This difference in density
of water masses provides a built-in resistance to vertical mixing of the various water layers,
and maintains stability in the water column; the greater the vertical increase (gradient) of
density, the greater the vertical stability of the water column. The greatest changes in density
20 Part One Living Resources

Figure 1.9 A piston coring sampler used to

obtain bottom sediment samples and core
samples. Photo courtesy of U.S. Navy.

Figure 1.10 Example of enormous pressure exerted by seawater at great depths. In the background
are iron buoys 2 ft (0.6 m) in diameter constructed of 1/4 in (2.5 mm) plate. The crushed buoy in the
foreground was lowered to 1,200 ft (400 m). Organisms living at great depths must contend with this
extreme pressure. Photo by the author.
Chapter 1 Ocean Environment 21

of seawater occur at the surface, where the water is subject to influences not present at
depths. Here density is increased by cooling, evaporation, and formation of ice; or decreased
by heating, precipitation, run-off from land, and melting of ice. The density of coastal water
is usually less than at corresponding depths offshore because of land drainage. Where there
is a significant amount of freshening due to land drainage, a tendency is for freshwater to
move offshore on the surface and denser (more saline) water to move in beneath it. Thus the
salinity of the surface water increases from the surface downward. This is an unstable con-
dition where the density surfaces slope downward in an inshore direction, and the resulting
unequal pressure causes currents.
Density determinations are rarely obtained by direct methods in the open ocean, because
it is difficult and time consuming to approach an accuracy comparable to converting from sa-
linity-temperature-depth profile data obtained via analyses of Nansen bottle casts and elec-
tronic salinometers. These indirect methods can provide density values accurate within
±0.0000l g/cm 3. In coastal waters, however, where great differences in density are found in
short distances, it is generally both practical and sufficiently accurate to determine density di-
rectly using stem hydrometers.

pH of Seawater
The term pH is used as a measure of the relative acidity or basicity of a solution. It is clas-
sically defined as the negative logarithm of the hydrogen ion concentration and can range
from 1 to 14: the lower the number the greater the acidity.
The pH of seawater varies within remarkably small limits all over the world (except in the
Dead Sea) and is usually between 8.0 and 8.4 at the surface. There are slight additional var-
iations (down to 7.5) with increase in depth and at the estuaries of rivers. The pH range of
natural freshwater bodies is usually 5.3-8.0. Most natural waters thus have a pH range that
varies from a little above neutral, pH 7.0 or basic, to a little below neutral or acidic. Ocea-
nographers are interested in pH information because it is directly related to processes of pho-

Figure 1.11 Large well-equipped oceanographic vessels are required platforms for gathering data on
the world's oceans. Photo courtesy of SOEST University of Hawaii School of Ocean & Science
22 Part One Living Resources

tosynthesis and other chemical changes. Different concentrations of hydrogen ions, like
salinity levels, will affect the physiology of marine organisms. Most marine animals have a
very narrow range of tolerance to change in pH.
To determine pH, colorimetric or potentiometric methods are used. The colorimetric
method is less precise, but is much simpler to conduct in the laboratory than the potentio-
metric, and uses pH sensitive dyes (indicators), either on paper strips or in solution. New
techniques include conductivity meters that use glass sensors (electrodes). pH sensors meas-
ure free hydrogen ions, a measure of water's acidity (Fig. 1.11).


Ocean waters are always in motion; some currents are very subtle, others are easily noticed.
The most obvious currents are those of a tidal nature, especially where they pass through re-
stricted areas such as narrow openings of bays between land masses. The primary driving
force for ocean currents is the wind, although the sun's heat and the Coriolis effect playa part.

Bottom Currents
Bottom currents in shallow water along the continental shelves carry plankton organisms
that sessile mollusks, such as oysters and clams, gather as food. Their oxygen needs and
waste product removal are met by currents. In the deep ocean, currents are difficult to
measure. They are maintained by density changes. Cold surface water in northern latitudes,
because it is denser than the underlying water, sinks and moves toward the Equator. Cur-
rents that bring nutrient-rich waters from the bottom (called upwelling) nourish the plankton
that supports vast fish populations, especially herringlike fishes.

Surface (Wind-Driven) Currents

Students should be familiar with the five great ocean gyres (movement in a circular
motion). Certain characteristics are well-known; gyres are clockwise in the northern hemi-
sphere and counterclockwise in the southern hemisphere. Winds from the west, flowing be-
tween 40° and 50° latitude north and south, produce the easterly flowing currents. Trade
winds drive the western currents near the Equator (Fig. 1.12). From a fisheries standpoint,
the Bengula current (Southwest Africa), and the Peru or Humboldt currents (western South
America) are extremely important. Both carry nutrient-rich cold water toward the Equator.
The Gulf Stream and the Kuroshio Current support large herring and tuna fisheries.
Strong trade winds move warm surface water into the western Pacific. If trade winds di-
minish, warm surface water flows back to the west coast of South America and causes a phe-
nomenon called El Nino, which generally develops just after Christmas concurrently with a
southerly shift in the tropical rain belt. In exceptional years the current may extend along the
coast of Peru to 12°5. When this occurs, sometimes very heavy rains fall on the coasts of
Peru and Ecuador. Furthermore, due to the lack of upwelled nutrient-rich cold water, plank-
ton and fish are killed in the coastal waters and a phenomenon somewhat like the red tide
results. During this time discolored water and intense displays of bioluminescence are
common. El Nino is much more widespread and destructive than the more local natural phe-
nomenon that occur every year, and affects the climates of the Americas.
The Sargasso Sea is near the center of a barren area in the North Atlantic gyre. This is an
area of clear, warm water where floating algae, called sargassum or gulfweed, accumulates
and European and American eels spawn.
Chapter 1 Ocean Environment 23

Figure 1.12 Surface ocean circulation. From Johnson (1968). Comm. Fish. Rev. 30(3): 27-38.

Ocean currents are measured by current meters, geotropic techniques (based on pressure),
and a device called a geomagnetic-electrokinetograph or GEK. The latter estimates the speed
of currents by a seawater conductor that moves through the earth's magnetic field and pro-
duces a gradient of electric potential in relation to the velocity of the current.

Tides cause water in the ocean and near-shore ocean features such as bays, gulfs, etc. to
periodically rise and fall. Were it not for this change in water height and the resulting tidal
currents, many organisms of value to humans could not live in near-shore habitats: Sessile
mollusks, for example, that live On tidal flats would not receive sufficient dissolved oxygen,
food would not be brought to them in sufficient quantities, and their wastes would not be
removed. Furthermore, when they spawn, the planktonic young would settle in close prox-
imity to their parents and increase the already strong competition for the necessities of life.
Fishermen who fish on tidal flats, such as clam fishermen, would find it difficult to harvest
clams if it were not for the fall of the water level. In many areas, where strong tidal currents
exist, fishermen take advantage of currents to set nets that capture finfish and shellfish (pe-
naeid shrimp) being carried by tidal currents as they leave lagoons, estuaries, or bays enroute
to deep offshore waters.
Tidal range results from the gravitational force of the mOOn and is aided or diminished by
that of the SUn. When the two bodies are in alignment with the earth they join forces to pro-
duce extremely high tides called "spring tides"; when the Sun is perpendicular to the mOOn
in relation to the position of the earth, extremely low tides called "neap tides" result. At the
moon quarters there is partial cancellation of the tidal forces, resulting in moderate tides
(Fig. 1.13).
24 Part One Living Resources


Figure 1.13 Tidal movements as they relate to the position of the sun, moon, and earth. Diagram
courtesy of the Alabama Department of Conservation.

Figure 1.14 Tides regulate the activities of many near-shore fishermen. Photo courtesy of Ivan FIye.
Chapter 1 Ocean Environment 25

Two high tides and two low tides every lunar day is the most common tide cycle through-
out the world. When depth increases as water moves toward the shore, the tide is said to be
"flooding"; when it reaches its highest or lowest level for an individual tide there is virtually
no water movement and it is said to be "slack water." After high slack and the water is run-
ning away from land, it is said to be "ebbing" (Fig. 1.14).
In this chapter I have not attempted to present a complete overview of the ocean environ-
ment, nor even all those aspects important to fishery scientists. That would be highly im-
practical and overambitious considering the broad coverage of this volume. Instead, I hope
that from the above text the reader will benefit from the possible useful insights discussed
and refer to suitable references for further information.


Anon. 1988. Seawater: its composition, properties & behaviour. Pergamon Press, Elmsford, NY.
Bacus, R. H. 1987. Georges Bank. MIT Press, Cambridge, MA.
Beer, T. 1983. Environmental oceanography: An introduction to the behaviour of coastal waters. Perga-
mon Press College Department, Elmsford, NY.
Black, J. A. 1986. Oceans and coasts: An introduction to oceanography. William C. Brown Publishers,
Dubuque, IA.
Bryant, T. and J. L. Pennock. 1988. The Delaware estuary. University of Delaware Sea Grant College Pro-
gram, Newark, DE.
Cushing, D. H. 1983. Climate and fisheries. Academic Press, Orlando, FL.
Cushing, D. H. and J. J. Walsh (ed.) 1976. The ecology of the seas. W. B. Saunders, Philadelphia, PA.
Day, J. w., Jr., c. A. S. Hall, W. M. Kemp, and A. Yanaz-Arancibia. 1989. Estuarine ecology. John Wiley
& Sons, Inc., Somerset, NJ.
Dugan, P. (ed) 1993. Wetlands in danger: A world conservation Atlas. Oxford University Press, New
Duxbury, A. C. and A. Duxbury. 1989. An introduction to the world's oceans. Wm. C. Brown Publish-
ers, Dubuque, IA. Second Edition.
Edwards, S. F. 1988. An introduction to coastal zone economics: Concepts, methods and case studies.
Taylor & Francis, Philadelphia, PA.
Ingmanson, D. and W. Wallace. 1989. Oceanography: An introduction. Wadsworth Publishing Co., Bel-
mont, CA. 4th Edition.
Johnson, F. G. and R. R. Stickney (eds.). 1989. Fisheries: Harvesting life from water. Kendall/Hunt Pub-
lishing Co., Dubuque, IA.
Kennedy, V. S. (ed.). 1984. The estuary as a filter. Academic Press, Orlando, FL.
Kennish, M. J. 1990. Ecology of estuaries. Volume II. Biological aspects. CRC Press, Boca Raton, FL.
Kaufmann, R. S. and R. R. Wilson, Jr. 1991. A summary and bibliography of seamount biota. SIO Ref-
erence Series. University of California, Scripps Institution of Oceanography, No. 91-8, 79 pp.
McLeod, G. C. and J. H. Prescott (eds). Georges Bank: Past, present and future of a marine
environment. Westview Press, Inc., Boulder, CO.
McLusky, D. S. 1981. The estuarine ecosystem. John Wiley & Sons, Inc., Somerset, N.J.
Negedly R. 1990. Elsevier's dictionary of fishery, processing, fish and shellfish names of the world. El-
sevier, Amsterdam.
Postma, H. and J. J. Zijlstra (eds.) 1988. Ecosystems of the World 27. Continental Shelves. Elsevier, Am-
26 Part One Living Resources

Richards, W. J. and J. A. Bohnsack. 1990. The Caribbean Sea: a large marine ecosystem in crisis. In
Large marine ecosystems: patterns, processes, and yields. Sherman, K., L. Alexander, and B. Gold
(eds.). 44-53. American Association for the Advancement of Science. Washington, D.C.
Walsh, J. J. 1987. On the nature of continental shelves. Academic Press, San Diego, CA.
Weatherly, A. H. 1972. Growth and ecology of fish populations. Academic Press, London.
Wilson, J. G. 1988. The biology of estuarine management. Croom Helm, Ltd., London.
Wilson, R R, Jr. and R S. Kaufman. 1987. Seamount biota and biogeography. In Keating, B. H. P.
Fryer, R Batiza and G. W. Boehlert (eds.). Seamounts, islands, and atolls. 355-377. Geophysical
Monograph # 43, American Geophysical Union, Washington, D.C.
Wolfe, D. A. (ed.). 1986. Estuarine variability. Academic Press, San Diego, CA.
Chapter 2

Major Resource Organisms

Plants and Invertebrates

This chapter summarizes biological and fishery information on example species, or groups of
species, of macroalgae and shellfish of commercial and/or recreational value. Productivity of
the oceans, including methods of estimating total resource production, is discussed in Chap-
ter to.
"Species" is a fundamental biological classification. Related species are grouped into a
larger category called a genus and are members of a group having a high degree of mutual
resemblance. Species are usually able to interbreed only among themselves. Consequently
the members of a species form a reproductively isolated group. The abbreviation sp. is
placed following a genus, meaning that a particular species is undescribed or may be un-
known to a writer. Spp. is the plural form used to indicate several species in a genus. The
diversity and number of species requires a system of classification called taxonomy, or sys-
tematic biology. The principal taxonomic groupings can be found in basic biology texts.
The outline of species included in this and the next chapter covers only a small fraction of
the phyla harvested by humans. Rather, examples of species or species groups are discussed
based on their use by humans, emphasizing ecological relationships, habits, and habitats im-
portant to fishery scientists in their attempt to improve fishing and information necessary to
form good management plans. The habitats and degree of mobility relates to sizes of fish (ju-
veniles and adults) sought by fishermen. Habitats of certain species can vary seasonally or
during different life stages; for example, young penaeid shrimp spend their early life in estu-
aries moving offshore as they mature, bury in, and move about on the substrate.
The weight of catches of species groups of invertebrates worldwide, shown in Table 2.1,
constitute only about 15% of the total weight of seafood landed. Invertebrates have a greater
value per pound than do finfish.
Biological information on the species described may vary considerably throughout the ge-
ographic range of the species. The fishing methods and gears listed are usually the most
widely used ones, but for some species a variety of distinctly different gears and methods
may be employed.
Chapters 2 and 3 are resource chapters that review marine plants and animals of known
value to humans, as food or for reduction as food for domestic animals, and for use in a va-
riety of other products. It is well known that only a few of the thousands of marine species
have value to humans. Of these only a relatively few are abundant, sufficiently available, or
valuable enough to support major fisheries. This chapter begins with an example of a mac-

28 Part One Living Resources

Table 2.1 World Commercial Catch of Crustaceans and Mollusks, By Species Groups,

Species group 1987 (1) 1988 (1) 1989 (1) 1990 (1) 1991

Thousand Metric Tons

Live Weight
Crabs 974 l,055 1,167 1,143 1,348
Krill 376 371 396 375 233
Lobsters 229 226 211 216 222
Shrimp 2,365 2,497 2,508 2,575 2,647
Other crustaceans 327 312 320 323 340
Abalones, winkles, conchs 101 100 87 71 68
Clams, cockles, arkshells 1,490 1,465 1,459 1,463 1,540
Mussels 1,135 1,287 1,311 1,321 1,332
Oysters 1,112 1,093 1,041 1,005 1,007
Scallops 739 868 838 877 816
Squids, cuttlefish, octopus 2,318 2,290 2,722 2,328 2,560
Other mollusks 799 847 751 797 791
Miscellaneous 364 281 312 283 342
Totals 12,329 12,352 13,123 12,777 13,023
Source: Food and Agriculture Organization of the United Nations. (FAO)-Yearbook of Fishery Statistics, 1991,
Vol. 72. Rome.

roalgae, and the remainder of the chapter is devoted to the invertebrates (shellfish). But first,
it is important to understand how precise identifications of the species are made by taxono-
mists. Once this is provided, the biological, ecological, and geographic range can be sought
and the effect of fishing on the stocks can proceed. To this end, taxonomists' methods are de-
scribed. Distinguishing characteristics between species of invertebrates are generally less ev-
ident and more variable than in vertebrates.
Brief outlines of general characteristics of the major invertebrate groups, for example, mol-
lusks or crustaceans, are followed by a biological profile of some important species in the
group. These species profiles consist of geographic range and habitat, biology (size, growth,
reproduction, etc.), food eaten, predators and fishing gears, and methods of capture. The
species selected from various groups occupy different habitats and behavior patterns, infor-
mation that is necessary for their capture. The gear mentioned in these chapters is defined
and methods of their use described in greater length in Chapters 8 and 9.


Seaweeds and large marine algae (many of which are called kelp) are grouped by botanists
using a variety of characteristics, but mainly color. Some are blue-green, some brown, and
some red, the latter two being the most commercially valuable. Many species of algae occur
along the sea coasts, mostly in intertidal zones. In Canada, about 300 species grow along the
Atlantic Coast. Of the numerous species of algae that occur in Hawaiian waters, over 70 are
edible. Most algae contain important minerals and vitamins valuable to the human body.
Macroalgaes are also harvested for chemical extraction.
Chapter 2 Major Resource Organisms-Plants and Invertebrates 29

Nearshore Shallow Water-Red Algae

Nori Porphyra spp., was cultured in Tokyo Bay beginning in the
late seventeenth century (Fig. 2.1). It is now farmed in Kona, Ha-
waii, Washington State, Alaska, North Carolina, British Columbia,
Oregon, Prince Edward Island, Canada, China, Korea, the Philip-
pines, Norway, England, and Chile. It occurs in the intertidal zone
of temperate waters in the Northern Hemisphere.
Biology: Fronds may be long, narrow, and knifelike, or broad
and fan-shaped with ruffled edges. Color varies from purple to
deep rose to black. When water temperature reaches about 72°F
(22°C), plants release spores (called conchocelis) that bore into oy-
ster shells, leaving them at a specific stage of maturity to find other
attachment, where they grow to large plants. (Porphyra reproduce
sexually.) Fronds may reach about 3 ft (1 m) in length and 1 ft (0.3
m) in width in about 7 to 8 months. This algae photosynthesizes
and extracts nutrients from the shallow water. It is preyed upon in
nature by sea urchins, in culture by snails, isopods, and amphi-
Several of the numerous species known from this genus are
eaten; red algae is traditional food in Asian countries and is exten-
sively cultured, especially in Japan. Some is eaten fresh, but most
is sold sun dried. Due to high levels of pollution in Japan's coastal
waters and the increased demand for nori, foreign sources are be-
ing sought.
Nori Fisheries: Fronds are hand harvested. Nori is popular in
Asia where it is used to impart flavor to sauces, broths, and soups. Figure 2.1 Nori.

Irish Moss, Chondrus crispus, is another shallow water

nearshore species of red algae (Fig. 2.2), that ranges from
Nova Scotia to Rhode Island where it lives on rocks and
ledges to depths of about 30 ft (10 m). It is most abun-
dant in semiexposed open coastal sites and estuarine
Biology: Irish Moss grows on exposed beaches, at-
tached to plants or hard surfaces. Its holdfasts and
fronds are perennial; fronds usually grow about 2 or 3
years, but holdfasts may live longer. Growth is slow; in
summer the maximum growth is less than 1/2 mm per
day. It is preyed upon by sea urchins, other echinoids,
gastropods, and crustaceans.
Irish Moss Fishery: This algae is collected by rakes
from wind rows of plants washed up on beaches. Chem-
Figure 2.2 Irish moss. icals extracted are used in many industries, including
food processing.

Nearshore Deep Water-Brown Algae

Giant kelp, Macrocystis pyrifera, is found in northern California's cold waters, to 150 ft (50 m)
(Fig. 2.3).
30 Part One Living Resources

Biology: This large algae may attain a height of 100

to 150 ft (32 to 50 m) in 6 months and reach the surface
at 130 ft (40 m) by 9 months. It may attain an age of 6
or more years. Giant kelp pass through several stages
in their life history: Special reproductive leaves produce
spores that are released into the water where they swim
freely before settling on the bottom to grow into tiny
plants that produce eggs and sperm that unite to grow
into the adult plants. This algae is preyed upon by sea
urchins, other echinoids, gastropods, and crustaceans.
Kelp Fisheries: The kelp is harvested from vessels
with kelp-cutting equipment. Its many uses include
food stabilizing and emulsifying, and cosmetics. Indus-
trial chemicals are also extracted.


These animals, low on the evolutionary scale, are

simple in structure and range in size from micro-
scopic to about 50 ft (15 m). Many are asymmetri-
cal; some are spherical or are radial or bilaterally
symmetrical. Simpler forms lack appendages;
more advanced invertebrates have jointed append-
ages. Many have a chitinous exoskeleton that is
shed periodically for growth, for example, shrimp
and crabs. Others, mollusks such as oysters and
clams, are encased in heavy, protective shells.
Some species have incomplete digestive tracts, ex-
cretory organs, circulatory or respiratory systems
or organs, and most do not have true segmenta-
tion (repetition of parts).
Figure 2.3 Giant kelp. Invertebrates eat plants or other animals, with
few exceptions those smaller and lower on the ev-
olutionary scale than themselves. Invertebrates are eaten by a wide variety of fish and other
invertebrates; some are cannibalistic. As a rule, they serve as intermediate or para tonic
(transport) hosts for parasites. Those that are sessile, or very slow-moving with large shells,
have many other species of plants and animals living on them or in a close association with
them, as conch fish inside queen conchs. Invertebrates are important in the food web, con-
stituting a high proportion (about 90%) of all marine species.

Identification, Classification, and Naming of Fish

and Shellfish
Most fish and shellfish are reasonably easy to identify as to species. Finfishes usually have
obvious anatomical characteristics that are usually sufficient to identify them to larger taxo-
nomic groups such as families and genera, and sometimes even to species. Body shape,
color, and fins (their sizes and locations on the fish body) are a few of the obvious character-
istics used by both layman and scientists for fish identification. However, some species of
Chapter 2 Major Resource Organisms-Plants and Invertebrates 31

finfish and some of the crabs and shrimp differ in minute characteristics, so careful examina-
tion by trained specialists is usually needed to correctly identify them to the species level.
For ease of identification, scientists use artificial dichotomous keys to "key out" or identify
unfamiliar fish and shellfish.
The aim of taxonomy (general principles of orderly scientific classification of living things)
is to provide stability and uniformity in the names of groups such as species and genera, to
which biota belong. This scientific research area reduces the amount of confusion that would
occur in publications if only common names were used. Further, taxonomy provides an un-
derstanding of inter- and intraspecific relationships of a wide variety of species.
Studies of the classification of organisms would be easier if, when a scientist names a spe-
cies, that name would be universally accepted forevermore. Unfortunately, some species
cover broad geographic ranges, and unfortunately, in their early identification organisms new
to science were collected and given different names by researchers working in different loca-
tions within the geographic range of the species, but without the means to communicate the
results of their findings. Today this confusion in names is eliminated by giving priority to
the first name given to an animal, providing it is accompanied by a complete and careful
dated and published description of the species. Still, some old scientific names persist.
Some shellfish have no approved common names, and those that do are badly in need of
standardization. A first step has been taken on this enormous job in the publication of Com-
mon and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mol-
lusks (American Fisheries Society. Special Publication No. 16. 277 pp. 1988).
Mollusca. Mollusca have soft bodies enclosed in a fleshy mantle. Most members of this
phylum have an external hard calcareous shell. The unsegmented body has a head, a dor-
sally located visceral mass, and a large ventral foot that enables very limited movement. In-
ternal shells permit squids and octopuses (cephalopods) to have a streamlined shape that
enables them to swim rapidly. They are found from the shore to depths of 35,000 ft (5,833
Most Mollusca eat seaweed and microorganisms, except for squids and octopuses that eat
other mollusks, crustaceans, and fishes. Some gastropods (oyster drills, tulip snails) are car-
nivorous. Mollusks are eaten by a wide variety of predators, including crustaceans, notably
crabs, lobsters, other mollusks (drills, octopuses), and finfishes.
Most serve as intermediate hosts for parasitic worms, but some often-studied Mollusca,
like oysters, are parasitized by a wide range of other organisms from viruses to helminths.
Spaces within the shells provide convenient shelter for many small crustaceans. Mollusca
that bury have fewer symbionts present on the exterior of their shells than those that do not
(abalones, oysters, etc.); these provide a suitable substrate for many commensal encrusting
organisms, algae, tube worms, boring sponges, and other Mollusca.
Gastropods. Conchs (Genus Strombus), whelks (Genus Busycon), moon snails (Genus Po-
linices), and abalone (Genus Haliotis) are all members of the Class Gastropoda. Except for the
abalone, the species mentioned here have a single, coiled shell with a horny operculum that
seals the shell opening. These marine snails feed by rasping their food with a ribbonlike se-
ries of teeth called radulae. Their diet generally consists of plants and possibly some benthic
organisms. Some species move moderate distances by means of a "foot," that, when ex-
tended, pushes them along with a sort of hopping motion, while others slide or glide over
the bottom on a path of excreted slime. The geographic range is wide, with different species
found in limited areas. Whelks in the Atlantic Ocean extend from Cape Cod to Florida; the
commercially important stocks of queen conch are found from Brazil to Bermuda, but not in
32 Part One Living Resources

the Gulf of Mexico. The moon snail ranges from Massachusetts to the Gulf of Mexico. For-
merly considered pests by oyster farmers and fisherman, whelks and moon snails have re-
cently achieved economic importance in the New England area. Abalone and queen conch
are important to commercial, subsistence, and recreational fishermen, and fishing is presently
regulated on these species.

ON ROCKY SUBSTRATE. Abalone, Haliotis sp.

range from Sitka, Alaska to Baja California in intertidal
and sublittoral zones, on rocks, under ledges, and in
fissures having an abundance of drift algae and vegeta-
tion (Fig. 2.4).
Biology: This marine snail has a large and concave
shell with holes over the gill cavity. It moves over the
sea bottom using its large foot. Abalone eggs are de-
mersal, and the planktonic stages last up to about 2
weeks. Growth is slow, about 1 in (2.5 cm) per year
and may reach over 11 in (28 cm). Foods include dia-
toms, unicellular algae, and, occasionally, coralline al-
gae. Juveniles and adults eat macroalgaes such as Ulva
and Macrocystis, although different species prefer differ-
ent algaes. They graze on the bottom where they eat
small animals such as hydrozoans, copepods, foraminif-
era, and byrozoans. Their predators are rock fish, sea Figure 2.4 Abalone.
otters, starfish, octopuses, whelks, rock crabs, and rock
Abalone Fisheries: Divers use flat bars to pry abalone loose. They are prized for both meat and their
shells, which are sold intact or cut up and polished for jewelry. The abalone fishery is tightly regulated
in California because of heavy fishing pressure and vulnerability to over fishing.

ON SANDY SUBSTRATE. Queen conch, Strombus gigas,

has a range extending throughout the western Atlantic
Ocean from Bermuda through the Bahamas, southeast-
ern Florida, southern Gulf of Mexico, the entire Carib-
bean to Brazil, from intertidal zones to 3,900 ft (650
fathoms) deep (Fig. 2.5).
Biology: Large (about 1 ft or 30 cm in length) and
colorful, this marine snail moves short distances on a
heavy muscular foot, extending it to raise the shell off
the bottom, then pushing itself forward. The queen
conch usually become sexually mature at the "flared
lip" shell stage. Large egg masses containing hundreds
of thousands of eggs per spawning are laid down on the
sea floor several times during the summer where they
lie until the veligers (larval stages) hatch and drift
Figure 2.5 Queen conch. among the plankton about 3 weeks before settling to the
bottom. Queen conch graze on the substrate and will
bury apparently to avoid predators and perhaps heavy
weather. They reach market size of about 1 ft (30 cm) in about 3 to 3 1/2 years. After settling to the
bottom, juvenile and adults eat detritus and microalgae. A crystalline style releases digestive enzymes
(microprotein gel). They are preyed upon by small octopuses, hermit crabs, spiny lobsters, spotted ea-
gle rays, loggerhead turtles, tulip shells, and finfishes.
Chapter 2 Major Resource Organisms-Plants and Invertebrates 33

Fisheries: Queen conch are caught by divers who hook them using a long pole with a hook on the
end. The meat (the foot) is a high-value product in Florida and the Caribbean Islands, as well as a basic
subsistence food. The beautiful shells are valuable, alone or fashioned into ornaments as souvenirs for
tourists; damaged shells are burned to make lime for mortar. A type of pink "pearl" occurs infrequently
in wild conchs. Primitive early inhabitants made tools from the shells. Unsubstantiated claims abound
that conch possess antitumor properties, and that the crystalline style is an aphrodisiac.
CLAMS. The most commercially important species of bivalves, clams are also sought by sport fish-
ermen (Fig. 2.6). Like all bivalves, the two shells (or valves) are opened and closed by powerful mus-
cles. They feed by filtering algae and diatoms over the gills and passing the food by hairlike cilia to the
mouth. Movement is limited, but is achieved by extending a foot to propel the animal over the bottom.
The geographical range is wide; certain species are found on virtually every coastline (protected or ex-
posed) in the northern hemisphere, all along the Atlantic and Pacific coasts. The commercially valuable
species in the United States are surf clams, Spisula solidissima, hard clams, Mercenaria mercenaria, and
soft-shell clams, Mya arenaria.

pallial Hne
Exterior View of Right Side Interior View of Right Valve
Figure 2.6 Anatomy of clam.


abrupta (generosa), are found in the lower reaches of the
intertidal zone of the eastern Pacific Ocean down to
about 30 fathoms (180 ft), from Alaska to Mexico (Fig.
2.7). This large clam, weighs about 3 lb (1.4 kg), al-
though there is a record of a 20 lb (9.1 kg) specimen.
The geoduck grows 1.2 in (30 mm) per year for the first
few years and may live 15 or 16 years. After about 10
years, growth is reduced. They live in permanent bur- Figure 2.7 Geoduck.
rows about 4.3 ft (1.3 m) deep in sand and mud sub-
strate. Spawning occurs in late spring and early summer and the spawn feed upon detritus. Young
stages are eaten by fish, but there is no record of predation on adults.
Geoduck Fisheries: They are dug on tidal flats and subtidal areas (diving).
Hard clam, Mercenaria mercenaria, are found in intertidal zones to 50 ft (15 m) depth of the west-
ern Atlantic Ocean from the Gulf of St. Lawrence to the Gulf of Mexico. Hard clams have two thick,
hard, equal-sized shells with two adductor muscles and short siphons. They may live 20-25 years and
reach 5.5 in (14 cm). Eggs are released free into the water to be fertilized, and after about 12 days, the
young settle to the bottom. Hard clams feed mostly on phytoplankton. They are subject when young
to high predation primarily by blue crabs and whelks. The big-clawed snapping shrimp is also a pred-
Fisheries: They are taken by hoes, dredges (hydraulic and escalator), tongs, rakes, and by hand.
Commercial market grades are based on length: clams 2 to 2 1/2 in (50-65 mm) are called little necks,
clams 21/2 to 31/4 in (66 to 79 mm) are called cherry stones, and clams larger than 31/4 in (80 mm)
are called chowders.
34 Part One Living Resources

Ocean Quahog, Arctica islandia, range in western Atlantic Ocean north from Cape Hatteras to
Newfoundland in depths of 5 to 40 ft (1.5 to 12 m) on the continental shelf. They live in shallow bur-
rows on bottoms composed of sand, mud, clay, gravel, and shell mixtures. Ocean quahogs reach a
length of 4 to 5 in (10 to 13 cm) and a width of 3 to 4 in (7.6 to 10 cm). The exterior of the shell (pe-
riostracum) is wrinkled and is dark brown or black, which accounts for the names "black quahog" or
"mahogany quahog." Spawning begins in late June to early July, peaks in August, and is usually com-
pleted by October. Larvae are planktonic for about 1 month. A slow-growing and long-lived clam,
the ocean quahog is perhaps the longest lived pelecypod found on the continental shelf, reaching an
estimated age of about 100 years. It filter feeds on phytoplankton. Predators are drills (gastropods).
Fisheries: Ocean quahogs are harvested by rakes and tongs in shallow water. Commercial fishermen
with powered boats tow a type of dredge made of a box-like steel cage having teeth for digging into the
substrate. Such a dredge is commonly called a "dry" dredge, in contrast to a "hydraulic" dredge, which
uses water jets to wash clams from the substrate.
SUBSTRATE OR WATER COLUMN. The sea mussel, Mytilus edulis, is widely distributed in northern
temperate regions of the western Atlantic and eastern Pacific Oceans. Mussels are bivalve mollusks
characterized by byssus (holding "threads") along the shell edges that are secreted from a gland at the
base of the foot. Shells are thin, bluish-black to black. They grow about 1 in (25 mm) per year for the
first 3 years. From 5 to 12 million eggs are spawned annually and ciliated larvae form about 4 hours
after fertilization. All organs are present after 10 weeks. Sessile as adults, they live on various sub-
strates, or attach to rocks, pilings etc. by their byssus. Mussels eat plankton: small diatoms (29 species),
protozoans (9 species), and detritus, drawing this floating food into the mouth by currents caused by
their gills. Predators are crabs, especially blue crabs, starfish, whelks, birds, and mammals (rats, musk-
rats, and walruses).

Byssol and Pedal Muscle

H i nge
L igomenl


Ventral Figure 2.8 Blue (sea) mussel.

Fisheries: They are caught by dredges, tongs, and rakes, and by hand. Numerous mussel culture
fisheries exist.
Oysters. The oyster is a sedentary mollusk like the clam, having two hard shells, or
valves, attached by a hinge and held together snugly by a strong muscle. This calcium car-
Chapter 2 Major Resource Organisms-Plants and Invertebrates 35

bonate armor shields the fleshy part of the adult oyster against most predators and adverse
environmental conditions. In general, the various oyster species fall into two groups: flat
(Ostrea) and cup-shaped (Crassostrea).
The two shell valves are dissimilar in all oysters. In genus Ostrea, the right (upper) valve
is flat and the left (lower) valve is concave or saucer-shaped, and is generally equally
rounded from hinged end to shell margin. In genus Crassostrea, the left (lower) valve is
deeply cupped, and is elongate from hinge end to shell margin. Ostrea reproduces by releas-
ing larvae free in the water; Crassostrea by releasing eggs. Crassostrea can tolerate higher sa-
linities and turbidity than Ostrea. Oysters extend over a worldwide geographic range,
mainly in temperate waters.
Oysters feed by pumping water between their valves, filtering out microorganisms in the
water by specially adapted hairlike structures in the gills that also move food toward the
mouth. They are eaten by many predators including gastropods (oyster drills, conchs), star-
fish, flatworms, crabs, and fish. Oysters, in common with many other mollusks, are most
vulnerable to predation when they are small and their shells are thin.

The American Oyster, Crassostrea virginica (also called

eastern oyster) occurs along the Atlantic Coast of North
America from the Canadian Maritime Provinces south-
ward around Florida, around the Gulf of Mexico (Fig.
2.9). It has been introduced along the Pacific Coast in
California and Washington, and in Hawaii. The valve
closest to the substrate to which the oyster is attached is
cup-shaped; the upper shell is generally flat; shell shapes
vary greatly. Spawning takes place when the tempera-
ture increases within a range of 68° to 90°F (20 to 32°C).
Individual females release 14 to 114 million eggs free in
the water to be fertilized. After hatching, young drift as
veliger larvae for about 2 to 3 weeks until suitable sub- Figure 2.9 American oyster.
strate is found, settling to the bottom and attaching
themselves to remain for their adult life. In the Canadian Maritime Provinces it takes 4 to 7 years to reach
3 in (7.6 cm); it may reach 15 in (38 cm) in 20 years. In Long Island Sound, it takes 1 year to reach 3/4 in,
3 years to 3 in (7.6 cm); in the Gulf of Mexico, it takes 2 years to reach 3 1/2 in (8.8 em). Their predators
include gastropods (oyster drills, whelks), starfish, flatworms, crabs, and fish. Competitors for food and
dissolved oxygen include boring sponges, clams and mud worms, oyster crabs, mussels, tunicates,
sponges, hydroids, bryzoans, ascidians, and algae.
Fisheries: The capture fisheries use tongs, rakes, picking by hand, and dredges (including hydraulic
dredges). Oysters are well-suited for farming because of their demand; they grow rapidly, and because
they attach to the bottom, they need not be impounded. They can be easily spawned, and by off-the-
bottom rearing techniques to reduce predation, a greater production of oysters can be achieved.
Olympia Oyster, Ostrea lurida, dwells in the Pacific northwest. This native Pacific oyster is a small
species; at 4 years it may be only about 1 1/2 in (4 cm) long. About 1902, the oyster (Crassostrea gigas),
now known as the Pacific oyster, was transplanted from Japan to the Pacific Northwest where it thrives
but does not reproduce freely; therefore, to maintain harvestable stocks in some areas, the state of Wash-
ington imports oyster seed from Japan each year. Oysters filter feed on plankton. They are commonly
preyed upon by drills, snails, starfish, boring sponges, crabs, skates, ducks, and red tides. In addition
to natural predators, Japanese drills may arrive with the imported seed. Several kinds of microscopic
single-celled plants produce "red tides" by releasing a toxin that can cause mortalities to oysters, clams,
and fish.
Fisheries: They are collected on tide flats by rakes and by hand at low tide.
36 Part One Living Resources

Scallops. The Atlantic Bay scallop, Argopecten (Aequi-

pecten) i"adians, lives in nearshore waters of the western
Atlantic Ocean, off the coast from New England south to
North Carolina, and in shallow water from Nova Scotia to
North Florida and Texas (Fig. 2.10). Their range includes
the northern side of the Greater Antilles and throughout the
Gulf of Mexico to Bermuda, just north of Cape Hatteras,
and the eastern Pacific Ocean in deep waters off Alaska,
south to California. Like clams and oysters, scallops are bi-
valves, having two rather similar shells, one flatter or less
concave than the other and hinged by a very large adductor
muscle. After a drifting larval life, the recently hatched
young settle to the bottom. They are able to swim short dis-
tances by contracting their adductor muscle and clapping
their shells together. They filter feed algae and diatoms,
protozoans, and detritus. Occasionally crustaceans, polycha-
etes, echinoderms, and sea weeds are found in their stom-
achs. Starfish are a major scallop predator. They eat the
entire animal by inserting their stomach between the shells,
and prying them open. Snails, crabs, fish, and water birds
also feed on scallops.
Fisheries: Scallops are fished by otter trawls, towed
Figure 2.10 Scallop. dredges, and divers.

Cephalopods. These mollusks have well-developed heads with a crown of suckers

and/or horn-bearing mobile arms that surround the mouth, a rasping structure (radula), and
large, often complex, eyes. They are highly mobile, having prehensile tentacles, and a mus-
cular funnel (siphon) with gills; unlike other mollusks, the shell is absent or internal and re-
duced. They are soft-bodied animals with well-developed hearing and smell. Cephalopods
prey on fish and crustaceans, such as crabs, prawns, and shrimp. Some squids eat their own
kind if one is injured. Predators are dolphins (porpoises), sea birds, seals, turtles, and
whales. Squid attempt to protect themselves by rapid emission of clouds of "ink," color
changes, and with their strong beaks.
Cephalopods, mainly octopuses and squids, support an extremely large world fishery, par-
ticularly in the Orient, Mediterranean, and eastern Atlantic areas. They are an important food
in the western Atlantic and Caribbean islands, and in the United States. The commercial catch
from the western central Atlantic is presently low, but the catch is anticipated to increase.
Because cephalopods have highly developed brain and sensory organs, they have become
important experimental animals in biomedical, behavioral, and neurophysiological research
having direct application to humans.

WATER COLUMN. Squids, Loligo sp., Illex sp. are found in all oceans from shallow, inshore waters
to ocean deeps (Fig. 2.11). These cephalopods, close relatives to octopuses, have elongate, torpedolike
bodies with eight long arms and two longer tentacles, and can change color. They are jet propelled,
moving by means of a siphon that takes water in and expels it. The giant squid is 50 ft (15 m) or more
in length) and is the largest known invertebrate. They eat shrimp, crabs, fishes, and other cephalopods.
Predators are fish including the swordfish.
Fisheries: Squids are caught by jigs, trawls, purse seines, pound nets and gill nets.
Chapter 2 Major Resource Organisms-Plants and Invertebrates 37

" .

Figure 2.11 Squid.


of the Octopuses, Octopus spp., is wide (Fig. 2.12); they
can be found in all oceans in depths of 3 to 164 ft (1 to
50 m). These eight-armed cephalopods change in color
from mottled brown, green, white, and white-spotted to
deep shades of red. Some migrate vertically. The fa-
vored bottom type varies with the species. If they can-
not find a suitable hole for shelter, they will construct a
den of stones and shells. They move by jet propulsion
by alternately spreading and closing their mantle for
movement. They also "walk" or slither over the bottom
using their arms. Depending upon the species, fecun-
dity varies. One species produces between 30,000 and a
Figure 2.12 Octopus.
million eggs. The eggs usually attach to the bottom, or
to some structure. The young may be planktonic or re-
semble adults, depending on the species. They live only a few years. Octopuses eat shrimp, crabs, fish,
other cephalopods, bivalve mollusks, squid, and starfish roe. Predators include moray eels, groupers,
snappers, and other reef-associated fish.
Fisheries: They are caught in trawls and unbaited pots.


A strong exoskeleton composed of chitin covers crustaceans' exteriors. In addition to pro-

viding protection for the animals, the exoskeleton provides attachment for muscles and forms
levers for body parts. Growth necessitates shedding; this means that a new, larger exoskele-
ton forms inside the old before it is cast away. Crustacea eat fresh and decaying fish or meat;
vegetation, including roots, shoots, and leaves of common seaweeds; clams; oysters; worms,
and other crustaceans; and are in turn eaten by other crustaceans (some species are cannibal-
istic), and predatory fish, such as groupers, jewfish, snappers, bottom fish, and sharks.
The exoskeletons of Crustacea are suitable substrate for settlement by many symbionts;
they serve as host for many species of microorganisms, including viruses and fungi and a
wide variety of parasites and symbionts. Molting rids the host of unwanted symbionts, but
because older crustaceans molt less often, some symbionts can become established.
38 Part One Living Resources

The true crabs possess a small, symmetrical abdomen that is bent and tucked under the
depressed cephalothrorax (carapace). They have five pairs of legs (as do the lobsters), the
first pair modified into large pincers (chelate) usually larger than the others, and the most
posterior legs are either modified to aid in swimming or are held under the body. Unlike
some other crustaceans, such as lobsters and shrimp, the abdomen lacks legs (uropods).
Crabs vary greatly in size, from a few inches to over 5 ft (1.5 m) across the outstretched legs.
If a leg is injured or grasped by an enemy (frequently the enemy is a fisherman), the crab can
drop it at a "breakage" point where healing and regrowth occurs, then grow another (regen-
Crabs-Bottom dwelling. The blue crab, Calli-
nectes sapidus, range extends along the western Atlantic
Ocean and Gulf Coasts of the United States, from Mas-
sachusetts to Texas, and are normally found in shallow,
nearshore saltwater, but also occur in lesser numbers in
brackish and freshwater (Fig. 2.13). This popular crusta-
cean actually has a brownish to dark green shell; it
reaches about 5 to 7 in (12.7 to 17.8 cm) across the back.
The blue color, for which it is named, is seen mostly on
the legs and pincers. Its carapace is about 2 1/2 times
as broad as it is long. It has five pairs of walking legs,
the last pair modified into swimming paddles; the first
Figure 2.13 Blue crab.
pair are chelate (clawed). The female produces about 2
million eggs in a "sponge" that attaches to her abdo-
men. Hatching occurs between 11 and 14 days, but it takes about 2 1/2 years to reach 6 in (15.2 cm) in
width. Blue crabs eat vegetation, fresh and decaying fish, mollusks, crustaceans, other blue crabs, and
insect larva. Their predators include other blue crabs (cannibalism) and raccoons.
Fisheries: They are caught by trot lines, pots, fyke nets, dip nets, and dredges.
The Dungeness crab, Cancer magister, is found in the eastern Pacific Ocean from the Aleutian Is-
lands to Mexico. The upper side is reddish-brown and the underside is whitish. They are slow grow-
ing and may require 3 to 4 years to reach market size. Females attain over 6 in (15.2 cm) and males may
be more than 10 in (25.4 cm) in body width. Mating occurs in summer, usually May to June. After the
eggs are extruded, the female carries them attached to her abdomen until late fall. About 1 year after
mating the young crabs, resembling adults, appear on the bottom. They eat small fish, oysters and
cockles, clams, barnacles, and amphipods. The larval stages are eaten by salmon and sea birds; adults
are eaten by ling cod, great marbled sculpin, wolf eels, halibut, rockfishes and other voracious fish.
Fisheries: Baited traps are used in deep waters, generally 30 to 240 ft (5 to 40 fathoms), mesh baskets
are used in shallow waters. The basket with bait (usually dead fish or clams in the center ring) is low-
ered to the bottom where it opens and lies flat. At intervals the basket is quickly raised, trapping the
crab in the meshes. On tide flats, rakes are sometimes used to get Dungness crabs out of tide pools.
Crabs must be delivered to processing plants alive and in good condition, so the vessels have holding
tanks aboard. In addition to the high cost of fishing for these crabs, picking the meat is tedious and ex-
King crab, Para litho des camtschatica, adults live in the cold depths of the northern Pacific Ocean,
in the Bering Sea, the Sea of Japan, and in the Sea of Okhotsk in Siberia (Fig. 2.14). Young may be
found in shallow waters, but move to deeper water as they age. Of the three species of king crabs, P.
camtschatica, the most abundant, has the widest geographic distribution. P. platypus and P. brevipes have
limited distribution. These huge creatures, dwellers on the continental shelf, congregate mainly on
sandy bottoms but are found elsewhere, except perhaps in the most rocky areas. (They should not be
Chapter 2 Major Resource Organisms-Plants and Invertebrates 39

confused with horseshoe crab, Limulus polyphemus, that

live off the Atlantic coast of the United States, which are
also called "king crabs.") King crabs are enormous,
reaching about 24 lb (11 kg) and have a leg span of
about 6 ft (2 m). The average size, however, is about 10
lb (4.5 kg). The shell is heavy and spined. The second,
third, and fourth pair of legs are larger than the first
pair, unlike other crabs whose first pair is largest. They
eat starfish, sea cucumbers, sea urchins, small crusta-
ceans, barnacles, bivalves (family Mytilidae), hermit
crabs, clams, and snails. Little is known about their en-
emies, but it is unlikely that adults have many because
of their large size.
Fisheries: They are caught almost entirely by large Figure 2.14 King crab.
pots (traps) 7 x 7 x 2 1/2 ft (2.1 x 2.1 x 0.7 m). Tangle
nets are used mainly by the Japanese. Handling large traps and giant crabs alive aboard the vessels un-
til delivery to the processing plant, coupled with heavy weather encountered on the fishing grounds, re-
quires large vessels. Stocks have been heavily overfished (discussed in Chapter 14).
The stone crab, Menippe mercenaria, inhabits the west-
ern Atlantic Ocean, southward from North Carolina
through the Gulf of Mexico to Mexico, in shallow, near-
shore waters 10 to 60 ft (3 to 18 m) deep (Fig. 2.15). Its
body is a dark reddish brown, spotted with gray, two-
thirds as long as it is wide. A large crustacean, it grows
to about 4.6 in (116 mm) across the carapace and has
powerful black-tipped claws and yellow and red
banded walking legs. A mature female can produce as
many as six million eggs in a spring-summer spawning
season. She carries the egg masses on her abdomen like
an apron for about 2 weeks, then migrates offshore to
saltier water where the eggs are released to hatch. It
takes about 3 years to reach adult size with claws large
enough for the commercial market, and about 1 year, in
nature, is necessary to regenerate claws to legal size. Its
Figure 2.15 Stone crab. diet is varied and includes polychaetes, sea grasses, oys-
ters, oyster drills, barnacles, conchs, flat worms, blue
crabs, hermit crabs, jellyfish, carrion, and other stone crabs (cannibalism). This species is preyed upon
by horse conch, sea turtles, octopuses, and fishes (cobia, grouper).
Fisheries: They are caught in baited pots (traps) and dipnets. Only a single claw is to be taken and
the rest of the body returned to sea to regenerate a new one. Stone crab claws are a desirable seafood.

Lobsters. These crustaceans range in length from a few centimeters to more than a meter.
The body is elongate, tubular, or flattened and has a prominent "tail" or abdomen made up
of six movable segments that terminates in a fan (telson). The anterior end (carapace) is rigid
and is about the same length as the abdomen. Some species are covered with spines, and
some have rather smooth exoskeletons. Lobsters have stalked, movable eyes, and most have
long, strong antennae; there are five pairs of walking legs; in some groups the first pair are
modified as pincers (chelate). Legs on the abdomen are short and biramous (pleopods). Lob-
sters occur off both coasts of the United States.
40 Part One Living Resources

LOBSTERS-BOTIOM DWELLING. The northern lobster (Ameri-

can lobster), Homarus americanus, ranges along the Atlantic
east coast of North America from Labrador to North Carolina
(Fig. 2.16). It is easily identified by two large, well-developed
claws that have teethlike forms and sharp spines. About 5 years
are required for them to reach marketable size, 3 3/16 in (8 em)
head length. Eggs are not released free in the ocean, but are ce-
mented to the hairs of the swimmerets while embryo develop-
ment takes place. Up to 60,000 eggs may be extruded during
one spawning. Young lobsters, upon hatching, pass through
several molts, and, after a short drifting period, take up a bot-
tom-dwelling existence. The northern lobsters, eat clams, fish,
scallops, other lobsters, and some vegetable matter. Frequently
they are scavengers. When young, many species of bottom fish
prey upon them; later they become prey to larger fish, such as
cod and sharks.
Fisheries: The northern (American) lobster brings Maine to
mind, and rightly so because that is where most of them are
caught (about 60%). Massachusetts is second (about 26%), fol-
lowed by Rhode Island (about 6%). The commercial fishery em-
ploys traps in the nearshore shallow waters and predates the
Revolutionary War. The fishery expanded during the latter part
of the 19th century, and by 1900, Maine had become the leading
lobster-producing state, a role it has maintained ever since. Dur-
ing the early period, lobsters were either canned or sold live.
About 1950, an offshore fishery for lobsters began using otter
Figure 2.16 Northern lobster. trawls in canyons along the continental slope. It peaked in 1970
with landings of about 3,200 mt, but declined to about 600 mt by
1976 when many fishermen returned to the traditional nearshore trap fishery.
Spiny Lobster, Panulirus argus, range the central western At-
lantic Ocean, extending from Bermuda and North Carolina
southward through the Caribbean and Gulf of Mexico, in the
Antilles and coasts of Central and South America to Rio de Ja-
neiro (Fig. 2.17). Related species are found in tropical and sub-
tropical waters worldwide. The spiny lobster is similar in
appearance to the northern lobster but lacks large claws and a
rigid fantail. It has long, whiplike antennae, and its body is cov-
ered with forward-projecting spines. Eggs attached to the abdo-
men of the female are incubated for 3 to 5 weeks. After the
larvae (phyllosomes) are cast free in the ocean, they drift among
the plankton for 6 to 12 months. Within each larval stage they
undergo instars (a stage between two molts) during which they
experience high mortality. The spiny lobster has a long larval
life. After settling, it may take between 18 months to 3 years to
grow from egg to market size; maximum size may reach 10 Ib
(4.5 kg). Water temperature plays an important role in its devel-
opment. Spiny lobsters eat small mollusks and crustaceans,
worms, seaweed, fish, coelenterates, echinoderms, and sponges.
The spiny lobster is cannibalistic and is a scavenger. Its preda-
tors and competitors are pelagic fishes including tuna (who eat
larvae), groupers, snappers, other bottom fish, sharks and rays, Figure 2.17 Spiny lobster.
octopuses, and turtles.
Chapter 2 Major Resource Organisms-Plants and Invertebrates 41

Fisheries: Commercial fishermen use wooden slat traps; recreational fishermen dive and catch lob-
sters by hand. A popular species, it is an important fishery in terms of value in the Caribbean islands,
Bahamas, and the Florida Keys. Major fisheries for related species are found worldwide.

Shrimps and Prawns. The terms "shrimp" and "prawn" are often used interchangeably;
however, in some areas prawns are considered to be larger than shrimp. Prawn is often used
in Europe rather than shrimp. In any case, in the United States, prawns live in freshwater
and shrimp live in salt or brackish water. The freshwater prawn included in this text is
somewhat larger than its marine relative. The shrimp body is nearly always laterally com-
pressed, long, and narrow with stalked eyes and antennae. The carapace bears a rostrum
that is usually compressed and toothed. The legs or pereopods are usually slender; but in
some shrimp, a single leg or pair of legs may be heavy, and some legs end in pincers. The
abdominal legs are used for swimming.
The life histories of commercial cold water shrimps are quite similar. Mature shrimp
breed in the late autumn or early winter. The developing eggs that appear on the abdomen
of the female shortly after breeding are carried over the winter months. The young shrimp
hatch in early spring and swim for about 3 months before settling to the bottom to begin
adult life. They mature first as males in their first or second year, depending on the species.
After one or two seasons as sexually active males, they change sex and function as females
for the rest of their lives.
Penaeid shrimps are usually abundant in warm water areas characterized by an inland
brackish marsh that is connected by passes or outlets to an offshore area having relatively
high salinity. The Florida Everglades is a perfect example of this environment. Eggs are laid
in the offshore water, and, after hatching, the larvae move inshore to "nursery" areas, usually
large estuarine bays. Later, the young juveniles move offshore to mature and remain in the
deep saline offshore waters.
iterus, species is found worldwide; in the United States it occurs in the warm waters of the western
Atlantic Ocean and Gulf of Mexico characterized by in-
land brackish marshes and connected by passes to an
area offshore of relatively high salinity (Fig. 2.18). Es-
timated to live about 1 year (perhaps as long as 20
months), females may reach about 11 in (28 cm), males
7 1/2 in (19 cm) total length. The male attaches a
packet of sperm cells to the female's sex organ soon af-
ter she molts. Between 500,000 and 1 million demersal
eggs are released free in the water upon fertilization.
The white shrimp may spawn more than once during its
life. In some areas, spawning takes place throughout Figure 2.18 Penaeid shrimp.
the year. Drifting larval stages last about 3 weeks.
Young juveniles live in nursery areas, usually large estuarine bays, moving offshore as they mature, to
deeper saline waters where the cycle repeats. Shrimp eat plankton, detritus, worms, and small mol-
lusks. Many fish species including croaker, spot, flounder, seatrout, and silver perch prey upon shrimp.
Fisheries: Otter trawls, beach seines, dip nets, butterfly nets, and channel nets are used.
dalus borealis, are found off the northwestern coast of North America. The two cold water genera,
Pandalus and Pandalopsis, contain 14 species of pandalid shrimps, but only about seven are caught or are
of sufficient abundance to support commercial fisheries. The two most important species are the north-
ern pink shrimp (also called pink shrimp), Pandalus borealis, fished on the continental shelves of the At-
42 Part One Living Resources

lantic and Pacific coasts of the United States in depths usually shallower than 650 ft (200 m), and the
ocean shrimp (also called pink shrimp), P. jordani, fished in the waters from California north through
Washington. Some enter the fishery at about 2 years of age. The following life history is based prima-
rily on the northern pink shrimp, P. borealis, in the Gulf of Maine. The life history of the cold water
shrimp (family Pandalidae) is different in several aspects from the warm water shrimp (family Penaei-
dae). Most pandalid shrimps begin life as males, that between 1 and about 5 years of age depending
on species and location, change to females. In both families, the male deposits a spermatophore (sperm
capsule) on the female prior to release of the eggs. However, after fertilization the female pandalid
shrimp attaches her eggs to her pleopods (swimming legs), but the female Penaeus shrimps releases her
eggs freely in the water. The fecundity of pandalid shrimps is much lower than Penaeus female
shrimps. A single female pandalid shrimp may extrude only about 2,000 eggs at a single spawning that
she carries on her abdominal appendages for about 7 to 8 months. The larvae that hatch in spring have
no resemblance to the adult, and are tiny, about 0.2 in (5 mm) long. During their planktonic, drifting
stage, they feed on planktonic organisms at middepths. Juveniles and adults feed mainly on euphausi-
ids and copepods. A wide range of bottom-dwelling finfish including cod, flounders, soles, halibut,
rock fishes, salmon, skates, rays, and harbor seals prey on shrimps.
Fisheries: The majority of cold water shrimp are caught with beam and otter trawls. In Alaska,
beam trawls are mainly used in southeastern Alaska; pots (traps) are used in both areas where rocky
bottoms prohibit use of trawls. Pots account for only a small portion of the total cold water shrimp
catch. On the east coast of the United States some shrimp traps are used, but it is mainly an otter trawl
fishery. The abundance of the northern pink shrimp, P. borealis, fluctuates widely, probably more so
than any other commercial shrimp species in the Gulf of Maine. This is generally attributed to the fact
that Maine is the southern end of their geographic range. Their abundance appears to be dependent on
water temperatures, being higher in cooler waters.


Note: Most references below pertain to commercial invertebrate species discussed in this chapter. General refer-
ences covering a wide variety of marine species can be found at the end of Chapter 3.
Ault, J. S. 1985. Species profiles: life histories and environmental requirements of coastal fishes and in-
vertebrates (Pacific Southwest). Black, green, and red abalones. U.S. Fish and Wildlife Service bio-
logical report 82(11.32); U.S. Army Corps of Engineers TR EL-82-4. U.S. Fish and Wildlife Service,
Slidell, LA.
Barr, L. 1970. Alaska's fishery resources: The shrimps. U.S. Dept. Interior, Fish. Leafl. 631. 10 pp.
Breisch, L. L. and V. S. Kennedy. 1980. A selected bibliography of worldwide oyster literature. Univer-
sity of Maryland Sea Grant publication UM-SG-TS-80-11. University of Maryland Sea Grant Pro-
gram, College Park, MD.
Brownell, W. N. and J. M. Stevely. 1981. The biology, fisheries and management of the queen conch,
Strombus gigas. Mar. Fish. Rev. 43(7):1-12.
Butler, T. H. 1980. Shrimps of the Pacific Coast of Canada. Can. Bull. Fish Aquat. Sci. No. 202. 280 pp.
Cheney, D. P. and T. F. Mumford, Jr. 1986. Shellfish & seaweed harvests of Puget Sound. Washington Sea
Grant Program, Seattle.
Cobb, J. S. and B. F. Phillips (eds.). 1980. The biology and management of lobsters. 2 vols. Academic
Press, New York.
Darcy, G. H. 1981. Annotated bibliography of the conch genus, Strombus (Gastropoda, Strombidae) in the
western Atlantic Ocean. NOAA technical report NMFS SSRF-748. National Marine Fisheries Serv-
ice, Seattle.
Dore, I. 1991. Shellfish: A guide to oysters, mussels, scallops, clams and similar products for the com-
mercial user. Van Nostrand Reinhold, New York.
Chapter 2 Major Resource Organisms-Plants and Invertebrates 43

Fay, C. W., R. J. Neves, and G. B. Pardue. 1983. Species profiles: Life histories and environmental re-
quirements of coastal fishes and invertebrates (Mid-Atlantic)-surf clam. U.S. Fish and Wildlife
Service FWS/OBS-82/11.13. U.S. Army Corps of Engineers TR EL-82-4. National Coastal Ecosys-
tems Team, U.S. Fish and Wildl. Serv., Slidell, LA.
Fischer, W. (ed.). 1978. FAO species identification sheets for fishery purposes. Western Central Atlantic
(Fishing Area 31). Vol. 6. Invertebrates and turtles. FAO Rome (Pages not numbered sequentially).
Galtsoff, P. S. 1964. The American oyster Crassostrea virginica Gmelin. Fish. Bull. 64:1-480.
Goodwin, L. and B. Pease. 1987. The distribution of geoduck (Panope abrupta) size, density, and quality
in relation to habitat characteristics such as geographic area, water depth, sediment type, and asso-
ciated flora and fauna in Puget Sound, Washington. State of Washington, Department of Fisheries
technical report no. 102. Washington Department of Fisheries, Shellfish Division, Olympia, WA.
Harvey, M. J. and J. McLachlan (eds.). 1973. Chondrus crispus. Nova Scotian Institute of Science, Halifax.
Holthuis, L. B. 1980. FAO species catalogue Vol. I-Shrimps and prawns of the world. An annotated
catalogue of species of interest to fisheries. FAO Fish. Synop. 125, 271 pp.
Holthuis, L. B. 1991. Marine lobsters of the world. FAO Species Synopsis No. 125, Vol. 13. 292 pp.
Iversen, E. S. and D. E. Jory. 1985. Queen conch at the crossroads. Sea Frontiers. Int. Oceanogr. Found.,
Vol. 31(6):150-159.
Iversen, E. S., D. E. Jory, and S. P. Bannerot. 1986. Predation on queen conch, Strombus gigas in the Ba-
hamas. Bull. Mar. Sci. 39(1):61-75.
Iversen, E. S., E. S. Rutherford, S. P. Bannerot, and D. E. Jory. 1987. Biological data on Berry Islands (Ba-
hamas) queen conchs, Strombus gigas, with mariculture and fisheries management implications. Fish.
Bull. 85(2):299-310.
Kato, S. and S. C. Schroeter. 1985. Biology of the red sea urchin, Strongylocentrotus jranciscanus, and its
fishery in California. Mar. Fish. Rev. 47(3):1-20.
Mottet, M. G. 1976. The fishery biology of sea urchins in the family Strongylocentrotidae. State of Wash-
ington, Department of Fisheries technical report no. 20. Department of Fisheries, Olympia, WA.
McHugh, J. L. et al. 1982. Annotated bibliography of the hard clam (Mercenaria mercenaria). NOAA tech-
nical report NMFS SSRF-756. National Marine Fisheries Service, Seattle.
McHugh, J. L. and M. W. Sumner. 1988. Annotated bibliography II of the hard clam, Mercenaria merce-
naria. NOAA technical report NMFS 68. National Marine Fisheries Service, Seattle.
Parker, P. S. and E. D. McRae, Jr. 1970. The ocean quahog, Arctica islandica, resource of the northwestern
Atlantic. NMFS. Fish. Indus. Res. 6(4):185-195.
Perez Farfante, 1. 1969. Western Atlantic Shrimps of the genus Penaeus . U.S. Fish. Wildl. Serv., Fish. Bull.
Roper, C. F. E., M. J. Sweeny, and C. E. Nauen. 1984. FAO species catalogue. Vol. 3. Cephalopods of
the world. An annotated and illustrated catalogue of species of interest to fisheries. 277 pp.
Stachowitsch, M. 1992. The invertebrates: An illustrated glossary. John Wiley & Sons, Inc., Somerset, NJ.
Shumway, S. E., (ed.). 1991. Scallops: Biology, ecology and aquaculture. Developments in aquaculture
and fisheries science 21. Elsevier, Amsterdam.
Taylor, H. 1984. The lobster: Its life cycle. Rev. ed. Pisces Books, New York.
Turgeon, D. D., et al. 1988. Common and scientific names of aquatic invertebrates from the United States
and Canada: Mollusks. Am. Fish. Soc. Sp. Pub. 16. 277 pp.
Wild, P. W. and R. N. Tasto (eds.). 1983. Life history, environment, and mariculture studies of the
Dungeness crab, Cancer magister, with emphasis on the central California fishery resource. Fish bul-
letin 172. California Department of Fish and Game, Long Beach.
Williams, A. B. 1965. Marine decapod crustaceans of the Carolinas. Fish. Bull. 65:1-298.
Yancey, R. M. and W. R. Welch. 1968. The Atlantic coast surf clam-with a partial bibliography. U.S. Fish
and Wildlife Service circular 288. U.s. Fish and Wildlife Service, Washington.
Chapter 3

Major Resource Organisms


The importance of U.S. commercial finfishes in this volume is based on total weight landed
of at least 100 million pounds (50,000 tons) per year. Annual landing records reported by the
National Marine Fisheries Service, using a 5-year average (1977-1981), include anchovy, At-
lantic herring, Gulf menhaden, salmon (pink and red), and tuna (skipjack and yellowfin).
Short histories of the fisheries are presented below for some of the species that have pre-
sented special fishery problems associated with their past management. These (species
groups) are Pacific sardine, pollock, American tuna, Pacific halibut, cod, fur seals, and


Vertebrates (Vertebrata) are a subphylum of a larger group (Phylum Chordata) that includes
all animals having a vertebral column or backbone. This large group includes fish, amphib-
ians, reptiles, birds, and mammals.
Vertebrate classes of importance to capture fisheries are finfishes, reptiles, and mammals
(Table 3.1). Numerous vertebrate species are of value to small scale fisheries in many coun-
tries throughout the world. Various vertebrate species are important predators on other ver-
tebrate species, on invertebrates, and on their own kind (cannibalism). These kinds of
feeding behaviors regulate the stock sizes of many species in the sea of value to humans.
Feeding behavior is discussed in Chapter 6.
As in Chapter 2, invertebrates, examples of principle distinguishing characteristics of fin-
fishes are used to identify finfishes. Distinguishing characteristics between closely related
species of finfish are often more obvious than for invertebrates.


Example: Salmo trutta Linnaeus 1785, brown trout

Salmo is the Genus to which the brown trout belongs. The first letter is always capitalized
and the name is underscored or printed in italics. Trutta is the species to which the brown
trout belongs. Note that the first letter is not capitalized. The word species is collective and

Chapter 3 Major Resource Organisms-Vertebrates 45

Table 3.1 World Commercial Catch of Fish by Species Groups, 1987-1991.

Species Group 1987 (1) 1988 (1) 1989 (1) 1990 (1) 1991

Thousand Metric Tons

Live Weight

Cods, hakes, haddock 13,786 13,636 12.905 11,827 10.476

Flatfish 1,292 1,342 1,204 1,223 1,113
Herrings, sardines, anchovies 22,375 24,387 24,800 22,183 21,406
Jacks, mullets, sauries 8,296 9,128 9,350 9,728 10,077
Mackerel, snoeks, cutlassfish 3,644 3,862 3,771 3,505 3,480
Redfish, basses, congers 5,713 5,705 5,917 5,656 5,742
River eels 103 116 110 120 121
Salmons, trouts, smelts 1,103 1,174 1,455 1,455 1,638
Shads 808 658 739 646 663
Sharks, rays, chimaeras 668 689 678 690 698
Sturgeons, paddlefish 24 21 19 18 15
Tunas, bonitos, billfish 3,644 4,065 4,082 4,373 4,478
Other fish 15.506 15,956 16,295 17,074 17,523
Totals 76,962 80,739 81,325 78,498 77,421
Source: Food and Agriculture Organization of the United Nations (FAO) Yearbook of Fishery Statistics, 1991,
Vol. 72, Rome.

never appears without the final s, unless coinage is being referred to. Then the word specie
is used. Like generic names, the specific name is underscored or printed in italics.
Linnaeus is the Latinized name of a Swedish botanist, Linne, who first named the brown
trout. He established the system of binomial nomenclature (genus and species) for naming
organisms. If the brown trout were moved to a different genus or species than that used by
Linne because of an erroneous or incomplete description, the name Linnaeus would be en-
closed in parentheses. 1785 is the year when the scientific description and name of the fish
was proposed in publication.
Brown trout is the acceptable name in common usage over the years, and is approved by
the American Fisheries Society Committee on Names of Fishes (Common and Scientific
Names of Fishes from the United States and Canada. Fourth Edition, 1980).
The International Commission on Zoological Nomenclature prepared a code of rules for
naming and describing organisms as long ago as 1780. The Commission continues today and
attempts to settle disputes that arise on scientific names. For example, in the early 1980s the
Commission ruled that the accepted scientific name of the tarpon was not Megalops atlantica
but rather Megalops atlanticus. This is a seemingly small change, but it insures that the name
is correct and will eliminate confusion in fish systematics.
Identification of certain species is difficult where differences appear in anatomical structure
or color, or are related to size and/or sex. Juveniles and adults of the same species may ex-
hibit anatomical differences. If taxonomists do not have a range of sizes of both sexes of an
unnamed species to examine when preparing a description, they may coin an unacceptable
name because of an incomplete description.
46 Part One Living Resources


Even though scientific names have been used for some 200 years, common names go far back
in history. Common names given to one species of fish may vary widely between distant ge-
ographic areas within the range of a fish or shellfish. Common names may reflect use by
man; for example, halibut means holy flat fish and is said to come from the fact that this spe-
cies was mainly eaten on holy days. The name may come from another familiar animal that
it resembles, such as the parrot fish that has fused teeth resembling the beak of a parrot. Or
the name may come from a location, like the mangrove snapper that lives in mangrove
swamps; or by the sound it produces, as with grunts and drums. Undesirable fish are some-
times given disparaging names, mud suckers, for example.
In Alaska, individual species of salmon are known by several different common names.
Chum salmon is the recommended common name for Oncorhynchus keta, but other common
names for this species are keta salmon and dog salmon, the latter name because it connotes
that the species is considered a low quality fish. The common name mullet (Mugil sp.) was
recommended to be changed to "liza" or "lisa" to appeal to Latin American buyers purchas-
ing mullet in Miami or for sale in South America because the name appeals to this ethnic
Since 1957, the American Fisheries Society Committee on Names of Fishes has published
lists of recommended scientific and common names of finfishes from United States and Can-
ada to prevent confusion by the standardization of finfish names. Common names are not
easy to standardize, so the Committee selects the one most widely accepted. Common
names are used by recreational and commercial fishermen, so standardization should avoid
problems and mistakes in recording catches and in marketing fish. In 1968, a "Multilingual
Dictionary of Fish and Fish Products" was published (revised in 1978), giving common
names in 15 languages.
The importance of exact identification of fishes is emphasized by the occurrence of highly
poisonous fishes in the same families that resemble fishes of commercial importance (rose
fish or rock fish), and in puffer fishes where misidentification has caused deaths.

Sharks-Continental shelf to deep water. Various species occupy a wide range of habitats
from deep ocean to very shallow coastal areas and in both cold and warm seas, with the greatest
number of species in tropical and subtropical waters (Fig. 3.1). Estimates place the number of shark spe-
cies at over 250 worldwide. Most biological information on sharks is divided between species harmful
to and those of value to humans. In general, sharks are long-lived, some known to live 25 years, having

L-------TOTAL L.ENGTH--------j


Figure 3.1 Shark anatomy.

Chapter 3 Major Resource Organisms-Vertebrates 47

a slow growth rate, late in life sexual maturity, and low fecundity. Many species tend to be top level
predators. Their apparent ecological role in the sea has been likened to wolves on land in which they
catch and consume any fish that is diseased, or injured, that thereby improves the gene pool of the prey
fish species because only the stronger, healthier fish will survive and reproduce. This notion is sug-
gested by their ability to perceive blood in the water, by their highly developed sense of smell, and also
by their attraction to sounds made by spear-wounded fish or fish struggling to get off a fishermen's
hook or out of his net.
Fishing: Certain shark species have been fished by man for many years using gill nets, hook and
line, and long-line gear. As food for human use, sharks have been ranked slightly below swordfish in
quality. The vitamin and mineral content of shark meat is high, it is relatively fat-free but has lower
protein values than bony fishes. A disadvantage is that urea must be removed to make shark meat
palatable. Sharks were fished formerly for the vitamin A in their livers, but when it became produced
synthetically, the fishery collapsed. Shark hides (called shagreen) were once used by cabinet makers to
polish wood; this shark product, too, was replaced by synthetic abrasives that could be produced more
cheaply and dependably. Other products obtained from sharks include using the hides for leather, the
teeth for jewelry, and the meat for bait for fish and crab traps. Sharks have been sought by recreational
fisherman for at least the last 50 years. This resource use has been promoted by some fishing clubs so
that heavy fishing pressure on certain popular sport fish could be reduced somewhat. Also recreational
fishermen find satisfaction in catching sharks because they are seen as a nuisance and a danger to man.


Nearshore-Estuarine Dependent Species

Atlantic Menhaden, Brevoortia tyrannus, oc- SPI NOUS DORSAL FIN
cur from Nova Scotia to central Florida. Another (I" DORSAL!

species, Gulf or large scale menhaden, B. patronus,

is found from southern Florida to Veracruz, Mexico
(Fig. 3.3). Both species live near the surface on the
coastal part of the continental shelf in summer
months. During colder months they move to
deeper waters off the shelf. Like other members of
the herring family, menhaden are somewhat flat-
tened laterally with a body about three times as
long as it is deep. The head is scaleless and large
in proportion to the body. It has a toothless large
mouth and a protruding lower jaw. The tail is
Figure 3.2 Anatomy of bony fish.
deeply forked. Body color varies from dark blue
to green, blue gray, or blue brown above, with sil-
very sides and belly, and fins with a yellow or
brassy luster. A conspicuous spot appears on each
side of the body behind the gill openings where a
number of smaller spots also can be found. Adults
weigh from 1/3 to 1 Ib (0.15 to 0.45 kg) and are
between 12 to 15 in (30.5 to 38 cm) in length.
Some mature at from 1 to 2 years; all are mature
by 3 years. Between 63,000 and 631,000 eggs may Figure 3.3 Menhaden.
be released by a single female, depending on her
size. Over their geographic range some groups of
menhaden spawn every month of the year. Some may live as long as 10 to 12 years, but 6- and 7-
year-old fish occur most frequently in the catches. An outstanding characteristic of the menhaden is
48 Part One Living Resources

the schooling behavior evident throughout their lives. They feed on plankton, primarily microscopic
plants and small crustaceans. Menhaden are preyed upon by nearly all larger inshore fishes, sharks,
marine mammals, and birds.
Fisheries: Purse seining is the principle fishing gear. The method depends wholly on sighting sur-
face schools from menhaden vessels and/or aircraft. A small fraction of the landings come from pound
nets. The catch is processed into fish meal, oils, and solubles that are used in dozens of ways. Recently,
attempts have been made to develop menhaden surimi in the United States. History of the fishery and
fishing methods are discussed in Chapters 8 and 9.

Reef Fishes-Nearshore, Rocky Bottom

Reefs are made of the skeletons of billions of tiny soft-bodied corals. The only true coral
reefs in the continental United States are in Florida. The number of resident fish species on
reefs probably exceeds 150. They are usually pretty fish, brightly colored with stripes or bars,
etc., and have many different feeding habits. They are night feeders and day feeders, some
are algal feeders and some predators. Reefs are host to numerous invertebrates, including
many species of crustaceans, shrimps, crabs, lobsters, conch, and sea urchins to name a few,
that serve as food for predatory fishes.
Red grouper, Epinephelus morio/Black grouper,
Myeteroperca bonaei, are found on rocky or muddy bot-
toms (Fig. 3.4). Young fish tend to inhabit inshore wa-
ters. Groupers occur to depths of 625 ft (104 fathoms)
where temperatures range from 59° to 86°F (15° to
30°C). The red grouper has small scales, large eyes, and
a robust body. Coloration is highly variable but is gen-
erally uniformly brownish-red with a lighter ventral col-
oration and a transient pattern of whitish spots;
Figure 3.4 Grouper. coloration can change to blend with surroundings.
These species are distinguishable from other groupers
by a long second spine on the dorsal fin, and by the unnotched interspinous membrane. Some groupers
are hermaphrodites, changing from female to male between ages between 5 and 10 years. Females ma-
ture between 4 and 6 years. Between 200,000 and 700,000 eggs are produced per female. Spawning oc-
curs late April to early May when the temperature of the water reaches about 7J'bF (25°C). Larvae
metamorphose to juveniles after about 1 month. They grow rapidly in cages, reaching about 21 oz (0.60
kg) in about 6 months, and may live as long as 30 years. The growth rate for each sex is similar, but
males reach a slightly larger ultimate length than do females. Opportunistic carnivores, groupers feed
on cephalopods, crustaceans, other invertebrates and fishes, competing with other groupers for food be-
cause of habitat overlap, but also with jacks, snappers, barracudas, and sharks. They are possible prey
to other groupers and sharks.
Fisheries: A hardy, popular, and expensive food fish, grouper has many good attributes for aquacul-
ture including rapid growth. They can be induced to spawn in captivity. There are numerous species
of groupers and flavor can vary widely by species. Just over 10 species are sold in the United States,
many from South and Central America. In some locations, certain grouper species are implicated in
ciguatera poisoning to man. Capture fishermen use hook and line (electric reels), traps, and spear guns.
Arrowhead traps are popular in the Caribbean Islands.
Chapter 3 Major Resource Organisms-Vertebrates 49


Anadramous-Salt Water to Fresh Water to Spawn

Pacific salmon, Oncorhynchus Spp., common and sci-
entific names are chinook, Oncorhynchus tshawytscha;
coho, O. kisutch; and steelhead, O. mykiss (formerly
Salmo gairdnen) (Fig. 3.5). Spawners and young juve-
niles range from California to Alaska in freshwater riv-
ers and streams with a marine lifestage in the north
Pacific Ocean generally east of the International Date Figure 3.5 Salmon.
Line. These anadramous fish are large and powerful,
reaching over 4 ft (1.2 m) in length, and 90 lb (41 kg) in weight. Females start nests in streams in gravel
and generally lay about 3,000 to 5,000 eggs fertilized externally by the male, who helps build the nest
then covers it with gravel. Young spend from 1 week to a year in freshwater as fry and fingerlings then
go to sea, later returning to the freshwater stream from which they came. With a few exceptions, Pacific
salmon die after spawning. In addition to time spent in freshwater (up to about I year), they may re-
main at sea for 3 to 4 years. As young in freshwater, chinook eat plankton. As adults at sea, they eat
herring, small shrimp, needlefish, and pilchards. Salmon are cannibalistic when young; adults are eaten
by fur seals and large finfishes.
Fisheries: Salmon in general have a high potential for successful aquaculture because of substantial
background on biological aspects, spawning, etc. They are easy to culture and hold in captivity. It is a
high-priced product for which strong markets exist. Net pen culture using Atlantic salmon is now a
proven industry in Norway, Canada, and Chile, yet salmon ranching in the Pacific Northwest, under-
way for many years, has been only moderately successful. Capture fisheries use gill net, purse seines,
and trolling. They are extremely valuable fish and support large capture fisheries in temperate waters
of the Northern Hemisphere.
American shad (roe shad-white shad), Alosa sapidis-
sima (Fig. 3.6) range along the east Coast of United
States from St. Lawrence, Canada to St. John's River,
Florida. Like salmon, shad are anadromous. The great-
est abundance occurs from North Carolina to Connecti-
cut. They have been successfully transplanted to the
Pacific Coast and are found from the Mexican border
north to Cook Inlet, Alaska. Shad are typical herringlike
fish. They have reached 15 lb (6.8 kg), but recent years
Figure 3.6 Shad. indicate a lower caught weight. Most of their life is
spent in the ocean, but, like salmon, they return to fresh-
water rivers and streams to spawn when 3 or 4 years old. In Florida, the southern end of their range,
they move up rivers to spawn as early as November, and at the northern extreme in Canada as late as
July. Stream mortality is high at the southern end of their range and low at the northern extreme.
Young eat insects and crustaceans in freshwater, and after shedding their teeth, they become plankton
feeders at sea and feed using their gill rakers. Eels and several species of freshwater fish such as carp,
catfish, and large-mouth bass, feed on juvenile shad. In California, striped bass stomachs have been
found to contain shad. In North Carolina, sea shad are eaten by sharks.
Fishery: They are caught by gill nets, pound nets, fyke nets, and haul seines.

Catadromous-Freshwater to Saltwater to Spawn

Eels. The geographic range of the American eel, Anquilla rostra (Fig. 3.7) in the Atlantic Ocean ex-
tends from Greenland to the Gulf of Mexico. Males tend to remain in estuaries, but females journey far
upstream into lakes and headwaters of rivers. The major portion of their lives are spent in fresh or
50 Part One Living Resources

brackish water. Juveniles are dark brown or olive green

on the dorsal surface and white to golden underneath,
becoming an all-over silver color at maturity. Eels
spawn in the Sargasso Sea hundreds of miles from
shore. Each female lays between 1 to 20 million eggs at
a depth of about 450 ft (75 fathoms) usually from Janu-
ary through March. The eggs float to the surface and
Figure 3.7 Eel. hatch into tiny prelarvae approximately 0.039 in (1 mm)
in length, and soon metamorphose into a larva called
leptocephali, which are transparent and ribbon shaped. They are weak swimmers and drift with the
ocean currents until they reach the coastline where they transform into elvers, and, carried by currents,
swim up freshwater streams and rivers along the coast where they mature into adults. Females live 10
to 13 years in freshwater and commonly reach a length of 3 ft (0.9 m). Lengths of 5 ft (1.5 m) are rare.
Males are much smaller; any that are longer than 20 in (51 cm) are rare. Weights of about 0.5 to 2 Ib
(0.2 to 0.9 kg) are common for eels 15 to 130 in (38 to 330 cm) long. After hatching, the young eels go
through a brief embryonic stage and derive nourishment from the yolk sac. Larval stages feed on
plankton near the sea surface. When in freshwater they prey on small fish. Plankton feeders consume
eels in their early larval stages; large fish eat juveniles and adults.
Fisheries: They are captured in cylindrical or square pots, by anchored nets placed in the streams, or
by hand-held dipnets or pushnets. Gear type is determined by their life stage (elvers or adults) and by
the time of year when gear is set. Strong markets exist in Europe (European eel, Anguilla anguilla) and
Japan (Japanese eel, A. japonica).

Bottom Fishes-Close Proximity to,

or On the Bottom
Atlantic Cod, Gadus morhua, (Fig. 3.8) are found in
American waters, from the southern tip of Baffin Island
(Canada) to North Carolina, the coastal waters of
Greenland, and the coast of Europe. Cod live in cold
waters close to the bottom over a wide range of
depths, 6 to 1,500 ft (1 to 250 fathoms). They are simi-
lar to hake and haddock, but the cod has three separate
dorsal fins and two anal fins. It is further character-
ized by a large head, stocky body, blunt nose, and wide Figure 3.8 Cod.
mouth. Cod can also be distinguished from haddock
by a pale lateral line, and from pollack by a squarish caudal fin, projecting upper jaw, and mottled
color pattern. Their winter spawning peaks in December. Females reach maturity as early as 2 to 4
years (coastal stocks) and as late as 7 years (northern oceans), depending on geographic locations of
the stocks, and produce up to 9 million eggs. They may live 30 years. There is a single record of a
cod 6 ft (1.8 m) long that weighed 200 Ib (90.7 kg), and several recorded weights of over 100 Ib (45.4
kg). Cod are bottom feeders; adults eat shellfish, clams, mussels, crabs, squid, and small fish. Preda-
tors are seals and fishes.
Fisheries: The cod fishery is of great economic value worldwide. It is Norway's main round fish.
Fry can be reared artificially with reasonable survival; however, rearing young stages is expensive. The
capture fishery uses many different kinds of gear, the most important of which are otter trawls, traps,
and hook and line.
Pacific Ocean perch, Sebastes alutus (one of about 50 species in the genus Sebastes), are found on the
eastern and northern rim of the Pacific Ocean from La Jolla, California to Kamchatka and the Bering
Sea. This species is common along the outer continental shelf and the upper continental slope, usually
in or around gullies, canyons, and submarine depressions; it is also present in deeper areas of the north
Atlantic. A small- to medium-sized fish, it may reach about 39 in (100 cm) in length. Some members
of the family can inject a poisonous venom through a hollow spine that may be fatal to humans. Pacific
Chapter 3 Major Resource Organisms-Vertebrates 51

Ocean perch are sought after food fish; although ugly, it is usually brilliantly colored, often shades of
red or rose, and has been called "rose fish" This somewhat slow-growing species may not reach sexual
maturity until about 6 to 9 years (males may mature at an earlier age). Highly fecund females may pro-
duce over 100,000 eggs at each spawning, thought to occur in late spring and early summer. Estimates
for large females suggest a maximum number of ova at twice that. Fertilization is internal, and individ-
ual males are not monogamous. They are carnivores that feed on a wide variety of food items, but as
juveniles they eat mostly copepods. Upon reaching maturity, they feed mainly on euphausiids. The Pa-
cific Ocean perch is an important food item for the Pacific halibut. They have been reported in the
stomachs of sperm whales.
Fisheries: They are caught by long line and otter trawls.

Flatfish. Flatfish have unusual life-styles. They begin life swimming upright like most other fishes
and later turn and lie on one side or the other, depending on the species. One eye migrates across the
forehead to the upper side, and skin pigment is lost on the side in contact with the bottom. The pig-
ment on the top side of older fish blends with the color of the bottom it lives on. Flatfish bury in
muddy bottoms with just their eyes exposed. When suitable prey species venture close, the flounder
will dart out of the mud with surprising speed and capture the victim. Transformation from swimming
upright to the bottom dwelling habits takes place in some flounder before they are half an inch long.
Once transformed they will spend the remainder of their lives swimming on their side. Identifying
characteristics are eye location and body outline. These fish are sometimes called flounders. The left-
eyed species includes the turbot (family Bothidae). Bothidae and Soleidae (true soles) are generally cir-
cular in outline when viewed from above. Right-eyed species are dab, plaice, and halibut (family Pleu-
ronectidae) and are thinner in outline. The halibuts are the largest and important commercial species of
flatfish. They occur in both the Atlantic and Pacific Oceans.
Pacific halibut, Hippoglossus stenolepis, are boreal
in distribution (Fig. 3.9). The female Pacific halibut is
fecund, laying between 2 and 3.5 million eggs. Females
become mature between 8 and 16 years of age (average
12 years); males reach maturity earlier at about 7 to 8
years. Large, flat fish, they can reach 8 ft (240 cm) and
weigh over 500 Ib (227 kg). Spawning takes place off
the continental shelf during the winter, usually Novem-
ber to March, depending somewhat on the area. Esti-
mates of about 6 months in the plankton stage have Figure 3.9 Halibut.
been made. They may live over 30 years. They are pis-
civorous as adults. Halibut are apparently not heavily preyed upon; there is some predation by canni-
balism, Pacific cod, and sand sole. Sleeper sharks have been observed to feed on halibut hooked on
ground lines. The extent of predation of halibut not hooked on fishing gear is unknown.
Fisheries: They are caught by two gear types: hook and line (ground line), a more desirable gear
from a biological standpoint because it tends to take larger fish and only halibut; and bottom trawls, un-
desirable gear that takes small to average sized halibut, plus nontarget species, that die.
Haddock, Melanagrammus aeglefinus, live in the At-
lantic Ocean off Newfoundland, Nova Scotia, the Gulf
of Maine and on Georges Bank (Fig. 3.10). They are also
found in northern Europe and in the British Isles.
Haddock are distinguished from their close relatives,
cod and pollock, by a black lateral line and black blotch
on the shoulder. A streamlined fish with three dorsal
fins and two anal fins, it is dark purplish-gray on the
dorsal side and silvery gray with pink below. It lives in
Figure 3.10 Haddock. deep water at 150 to 450 ft (25 to 75 fathoms). Rel-
atively nonmigratory, haddock are usually associated
52 Part One Living Resources

with smooth bottoms or those with pebbles, gravel, or broken shell. Spawning takes place from Febru-
ary to May in the New England area. Single females may spawn 150,000 to 2 million eggs. Haddock
may live about 15 years and attain a weight of 37 lb (16.8 kg). Early life is spent near the surface; at
about 5 months they return to the ocean floor. After absorbing their remaining yolk, the young feed on
zooplankton. Adults feed on a wide variety of invertebrates, some of which they dig from the bottom
with their strong lips.
Fisheries: They are caught by otter trawls, gill nets, and long lines.

Seamounts-Atop Undersea Mountains

Arrnorhead, Pentaceros richardsoni (Fig. 3.11), is widely
distributed in the north Pacific from California north to
Vancouver and Queen Charlotte Islands. The species is
also found on seamounts in the north central Pacific
northwest of the Hawaiian Archipelago. Armorhead
also occur off South Africa, New Zealand, and Japan.
As the name implies, the major characteristic is the unu-
sual armature of the head, which is almost completely
enclosed in exposed, rough, striated bones. Those sam-
pled on seamounts were about 12 in (30 cm) in average
length. Coloration and condition of the fish (fatness)
Figure 3.11 Armor head. differs considerably from seamount to seamount. Little
is known of their biology other than they are pelagic de-
mersal fishes in the vicinity of seamounts. They appear to be pelagic spawners. Stomach analyses
show they feed heavily on crustaceans associated with the deep-scattering layer. Vertical night migra-
tions are necessary for armorheads to feed on this layer that ascends at night and is usually deep during
the day.
Fisheries: Soviet and Japanese began fishing them with trawls, gillnets, or bottom longlines in late
1967. United States fishermen did not fish for armorhead until U.S. exploratory fishing vessels deter-
mined that the fishery could be profitable. Fishery scientists had been unaware of the kinds and abun-
dance of fishes associated with seamounts. They are usually caught together with a somewhat smaller
fish 8 in. (20 cm) long called the aifonsin, Beryx splendens, a bright red-colored fish that also inhabits
rocky bottoms on seamounts.
Fishing: Bottom longlines were used to catch the less common aifonsin, before a moratorium on
fishing for armorhead on the Hancock seamounts, due to declining catches, was imposed in 1984. This
has since been extended.

Pelagic Fishes-Open Sea

Yellowfin tuna, Thunnus albacores (Fig. 3.12), are found in tropical to temperate waters in the At-
lantic, Pacific, and Indian oceans. The tuna is a large, powerful, streamlined, fast-swimming fish that
may reach 8 ft (2.4 m) in length and 450 lb (204 kg). Its color fades rapidly when caught, but when
alive it is brilliantly iridescent with a golden or bright yellowish stripe along the sides and, as the
name implies, the fins are yellow. The back is bluish-black and the undersides white. Yellowfin tuna
grow rapidly, attaining 5 ft (1.5 m) in about 5 years. The species probably feeds more actively in day-
light than at night, randomly on pelagic small fishes and squid. It is cannibalistic on smaller yellowfin
tuna. Large yellowfin eat pelagic fish, about 60% by volume, and squid, about 30%, with some mol-
lusks and crustaceans. Larvae and juveniles are eaten by many kinds of fish. Billfish and sharks eat
adult tuna.
Fisheries: The yellowfin, Thunnus albacares, and skipjack, Euthynnus pelamis, tunas support an impor-
tant fishery in the eastern Pacific Ocean; they aggregate into schools near the sea surface, a habit that is
the basis of the U.s. tuna fishery, because both pole and line, and purse seining operations depend for
economic operation on fishermen locating and harvesting fish from large, relatively compact schools
Chapter 3 Major Resource Organisms-Vertebrates 53

(Atlantic bonito)

.. udal

caudal fin

Figure 3.12 Tuna.

swimming near the surface. The first United States west coast tuna boats probably evolved from sar-
dine lampara seine boats that used jigs and baited handlines. The large [260 ft (79 m) long] multimillion
dollar tuna boats in use today came much later. In the early fishery, tuna were caught close to Mexico
and either landed at shore installations, or sold to freezer vessels. Also, live bait (anchovette) necessary
for fishing was caught near the Mexican coast. When Mexico raised the export duty on tuna taken in
her waters from $5 to over $37 per ton in 1924, the United States tuna industry responded by develop-
ing a high seas vessel that did not have to deliver catches to a shore station or obtain supplies or live
bait. A tuna vessel with a live bait well that could range several thousand miles was launched in 1924,
freeing United States fishermen from dependency on Mexico. To preserve tuna catches on these long
trips, brine refrigeration was installed to precool fish before stowing them below decks. Heavy nets
made from natural fibers (before the invention of nylon) were difficult to set around schools of fast sur-
face-swimming tuna and retrieve (before hydraulic power blocks). About 1948, a tuna purse seine made
of lightweight nylon was successfully used in the fishery. These nets became popular with tuna fisher-
men because they avoided the problems of searching for, capturing, and holding live bait. Today, tuna
is the most valuable fishery in the United States. In 1992, landings of tuna by U.S. fishermen at ports
in the 50 states, Puerto Rico, American Samoa, and other U.S. territories and foreign ports were about
580 million lb (263 million kg), valued at about $270 million.
Billfishes. The word "billfish" is a term encompassing several kinds of large pelagic fishes, some
exceeding 1,000 lb (454 g). They belong to two families, Istiophoridae (marlins, sailfish, and spearfish),
and Xiphiidae (swordfish). The Istiophoridae are primarily recreational fish because of their large size
and fighting characteristics when hooked (Fig. 3.13). In a short burst of speed a sailfish has been
clocked at 68 mph (110 km/h). The sword fish is primarily a commercial fish although it is popular
with recreational fishermen for the same reasons as the sailfish. In addition, its abundance in an area
draws tourists. Billfish are capable of long migrations and are found in all the warm seas. The sword-
fish is described below as an example species. Marlins, sailfish, and spearfish will be referred to later
in the recreational fishing chapters.
Swordfish, Xiphias gladius (Fig. 3.14), occur in all tropical, subtropical, and temperate seas generally
between 45°N and 45°S latitude, from the Chile to Los Angeles, California, in and around the Hawaiian
Island in the Pacific, and from the West Indies to the Grand Banks. They travel singly or in small
schools, and may go to depths of about 1,000 ft ( 167 fathoms). They normally inhabit waters of 5S'F
(13°C) or warmer. Thirty or more common names, each originating from the various countries where
swordfish are caught, have been recorded. Even within certain fisheries several different common
names may be used. The body of the swordfish is streamlined with the greatest thickness at the shoul-
der area tapering back to a tunalike caudal fin. Like the tuna, it has keels on either side of the caudal
54 Part One Living Resources

T.trapturus a/bidus
(white marlin) -


-- -- _.. _n,. ...

-.. -
Figure 3.13 Billfish.

peduncle just in advance of the caudal fin. The outstanding characteristic from which the common
name comes is the protrusion of the upper jaw into a stout, sharp swordlike appendage that may reach
up to about one-third of the total length of the fish.
Swordfish are large fish that normally weigh between
200 and 400 Ib (90.7 to 181 kg) and may reach 15 ft (4.6
m) in length. A swordfish caught off Chile weighed
over 1,100 Ib (500 kg). Members of this species may live
as long as 9 years. The dorsal body surface is a dark
metallic purplish color, becoming almost pure white on
the sides and lower body. Swordfish spawn in surface
waters; spawning time varies by geographic location.
They are believed to spawn year round in some waters.
Swordfish eat primarily pelagic fish and squid. They
prey on a wide variety of fish and some invertebrates.
300 em They are eaten when young by tunas, marlin, dolphin
fish, and sharks. Adult swordfish have few enemies be-
cause of their speed and size, mostly sperm whales,
killer whales, and large sharks.
120 ell! Fisheries: Longlines and harpoons are the principal
kinds of gear in most swordfish fisheries. Harpoons

were for many years the only gear used until large
swordfish catches were made by the Japanese in their
tuna longline fishery, and by the Norwegians in their
38 ell!
shark fishery. Fishing takes place at night. Swordfish
Figure 3.14 Swordfish. are powerful swimmers and when hooked or har-
pooned, they put up a gallant fight. Their large size and
fighting qualities make them popular with high seas
sport fishermen. Rod and reel is used by recreational fishermen. Restrictions on the sale of swordfish
because of high mercury levels in their flesh were imposed in Canada and the United States in the early
1970s. The U.S. fishery landing dropped markedly and the Canadian fishery collapsed. Allowable mer-
cury levels have been raised and the fisheries continued. Recent U.S. catches have exceeded 11 million
Ib (4950 mt).
Chapter 3 Major Resource Organisms-Vertebrates 55

Pelagic Fishes-Coastal
Sea herring (Atlantic sea herring), Clupea harengus,
are found in the western Atlantic Ocean from northern
Labrador to Block Island, Rhode Island, and as far south
as Cape Hatteras (Fig. 3.15). In the eastern Atlantic they
range from the Strait of Gibraltar to Norway. They
grow to about 20 in (50 cm) maximum size. Spawning
takes place nearshore and on banks and shoals offshore,
sometimes as far as 25 miles. The females lay 20,000 to
40,000 adhesive eggs that attach to the substrate. Sea Figure 3.15 Sea herring.
herring travel in large schools in open water. They eat
small shrimp, copepods, occasionally small larvae, and
eels, and may be cannibalistic on young herring.
Fishing: Herring are caught by gill nets and purse seines.
Pacific sardine, Sardinops sagax, is characteristically
found in cool waters in upwelling areas of the eastern
Pacific Ocean from southeastern Alaska southward to
the tip of Baja, California, and also in the Gulf of Cali-
fornia (Fig. 3.16). Young sardines 1 and 2 years old oc-
cur along central California; larger, older sardines occur
mainly along the Pacific Northwest. This sardine may
reach a length of about 1 ft (30 em). Highly fecund, the
Figure 3.16 Sardine. numbers of ova counted when nearly ready to be re-
leased number about 27,000. In the southern end of its
range, spawning occurs throughout the year with peaks during the winter and summer months over
deep water, varying somewhat with latitude. Sardines remain in the larval, planktonic stage for 1 or 2
months, and may live about 13 years. The fishery normally exploits sardines at ages 1 to 7 years. They
tend to form large schools, some estimated to weigh several hundred tons. Pacific sardines are filter
feeders that prey upon planktonic animals, mostly copepods. Their predators include a number of fil-
ter-feeding nekton, fish, and crustacea. Adults are eaten by tunas, sharks, yellowtail, barracuda, bonito,
marlin, hake, mackerel, and mammals like sea lions, porpoises, and whales. Birds, sea gulls, pelicans,
and cormorants also prey upon sardines.
Fisheries: Nearly all sardines are taken by round haul nets, purse seines, or ring nets. Beginning
with the 1945-1946 fishing season there was a sharp decline in the landings of Pacifie sardines. Regu-
lations were unsuccessful in preventing further declines. (See Chapter 12).
Northern (California) anchovy, Engraulis mordax,
are found along the eastern Pacific coast from southern
Alaska to lower California (Fig. 3.17). The northern an-
chovy is a small fish, reaching a maximum length of
about 9 in (22 em), usually not over 7 in (17.8 em). The
northern anchovy is believed to be short-lived, perhaps
only 3 years. Many reach sexual maturity at about 1
year, and spawning occurs several times each year. Figure 3.17 Anchovy.
Population abundance fluctuates widely from year to
year. The species forms massive schools that may mi-
grate over long distances (tagged anchovies moved 360 miles in 129 days, or about 3 miles per day).
Anchovies feed on plankton organisms including larval stages of crustaceans and mollusks. They are
random filter feeders and are occasionally particulate feeders. Larger fish prey upon them.
Fisheries: These oily-fleshed anchovies, generally sought for reduction to fish meal, are also canned
for human consumption and pet food. They are caught using round haul nets. From 1949 to 1952 Cal-
ifornia restricted the use of anchovies as baitfish.
56 Part One Living Resources

Pacific hake, Meriuccius pro ductus, range the eastern

Pacific Ocean from the Gulf of Alaska to the Gulf of Cal-
ifornia (Fig. 3.18). They are found on the upper conti-
nental slope in large schools. Hake may reach about 3 ft
(1 m) in length. Spawning takes place mainly offshore
along the coasts of southern California during December
to April. Seasonal migration is northward in spring and
Figure 3.18 Pacific hake.
summer and southward in the fall. Females grow faster
than males and both may live 10 years. Hake eat fish
and invertebrates; primary food may be euphausiids. Predators on larval hake are zooplankton; numer-
ous finfish species, birds, seals, whales, and porpoises feed on juveniles and adults.
Fisheries: They are caught with otter trawls.
Walleye pollock, Theragra chalcogramma. (Fig. 3.19),
is a Pacific Ocean species whose range extends from
Central California north through the Bering Sea to St.
Lawrence Island, and on the Asian coast to Kamchatka,
the Okhotsk Sea, and the Sea of Japan. The walleye pol-
lock, also known as the Alaska pollock, is a demersal
species in the family Gadidae. Fertilization is external.
Figure 3.19 Walleye pollock.
Fecundity is high; estimates made from many different
areas range from 29,000 to 670,000 eggs per female.
Spawning occurs in summer. Pollock grow rapidly during their early years, as much as 3 in (7.6 cm)
per year. Beyond year four, growth slows considerably. The oldest pollock captured in American wa-
ters was a 17-year-old female. Foods are shrimps, sand launce, and young salmon. Adults are eaten by
Fisheries: The Gulf of Alaska pollock fishery was composed of a multispecies foreign bottom-trawl
fishery (a fleet of freezer and surimi trawlers operated by foreign nationals) during the early years
(1964-1971), at which time this species supported important fisheries in Japan, the USSR, and a smaller
fishery in the Republic of Korea. The Japanese use pollock to make surimi, a minced product used for
imitation crab, scallops, and shrimps. Before the Magnuson Fisheries and Conservation Act of 1976, the
fishery was exclusively worked by foreign fleets that were mainly after ocean perch. During a second
period foreign fleets targeted pollock. During a third period (1981-1985), the fisheries shifted from for-
eign only to joint venture fisheries wherein u.s. catcher boats sold fish at sea to foreign ships. Begin-
ning in 1986 through 1988, the fishery became a sole venture of U.S. fishermen and expanded
dramatically. This movement of U.S. fishermen into this fishery was induced by declines in the king
crab fishery (a result of overfishing), the formation of joint ventures, and the discovery of large concen-
trations of spawning pollock. Shoreside and at-sea processing of pollock was begun in the United States
to produce surimi. Since the collapse of the overfished Peruvian anchovy fishery, walleye pollock has
been the world's largest single-species fishery. In the North Pacific there may be as many as 12 major
stocks, three of which are found around North America. It is not an old fishery, but was limited to
coastal waters off Japan and Korea until the late 1950s when it became a distant water fishery. Little is
known about the biology of the pollock in the United States, perhaps because the fishery is compara-
tively new, and was formerly exploited in U.S. waters only by foreign nationals. The United States is
taking a more active role in fishing this species within its 200-mile limit, and fishery biologists have in-
itiated studies on the pollock. Landings in the Gulf of Alaska increased from 1,000 metric tons in 1964
to a peak harvest of 307,000 in 1984.


Reptiles are cold-blooded vertebrates usually covered with scales or bony plates; they breathe
with lungs, and are most abundant in warmer regions of the world. Reptiles raised by aquacul-
Chapter 3 Major Resource Organisms-Vertebrates 57

turists are crocodiles and alligators, for the meat and skin (leather), and turtles; freshwater tur-
tles for pets and food, and sea turtles for commercial products, mainly for shells and meat.
Reptiles may be completely terrestrial, semiaquatic, or aquatic. Those included here spend
nearly all of their lives in the sea. Females emerge briefly to lay their eggs in nests on beaches.
The Atlantic green sea turtle, Chelonia mydas (Fig.
3.20), is a migratory species. Individuals tagged at As-
cension Island, South Atlantic Ocean traveled to the
coast of Brazil, a distance of about 1,400 nautical mi
(2,593 km). Their range is confined to tropical and
subtropical latitudes. Mating is external and takes
place in the sea. Eggs are laid in sand nests on
beaches and covered over; upon hatching the young
crawl to the sea where they live until they return to
the same beach to spawn where they hatched. This
large animal may reach 308 lb (140 kg) and may live at
least 15 years. Sea turtles are mainly herbivores and Figure 3.20 Sea turtle.
eat a variety of sea grasses including Thalassia or tur-
tle grass. Some small crustaceans and mollusks are also eaten. Newly hatched young turtles are often
eaten by sea gulls before they can reach the sea. Some mammals dig up turtle nests on beaches and eat
the eggs.
Fisheries: Turtle meat, oil, and shell can be used with little waste. In recent years the turtle market
has suffered a serious setback because green turtles have been declared an endangered species. How-
ever, fishing is still carried out around the world using tangle nets, hand-held harpoons, and seines at
sea. One unusual method of capture is using one or more remoras, Echeneis naucrates, attached to lines.
When the remoras find sea turtles and attach themselves to the carapace they are hauled aboard the
boat by the lines. Capturing egg-laying females or collecting their eggs on the beach is practiced in
some remote areas, although it is illegal in many countries. Although trade in farmed or ranched spe-
cies that are declared endangered is allowed, certain regulatory controls must be satisfied. Furthermore,
there is emotional pressure against trading in species where the wild populations have been declared
threatened or endangered by the Convention on International Trade in Endangered Species of Wild
Fauna and Flora (CITES).


Pelagic Mammals-Whales
Thanks to television documentaries, conservation ori-
ented magazines, and the activities of environmental
groups the pathetic story of the destruction of thou-
sands of whales has been documented and has alerted
the public to the need for conservation measure (Fig.
3.21). The belated struggle for proper whale conserva-
Figure 3.21 Whales. tion measures is not over because of traditional fisher-
ies in countries where whale meat has been a staple
food in the population. Today, only the sei and sperm whales are caught in any numbers. Whales are
divided into two groups: toothed whales (Odontoceti) and baleen whales (Mysticeti). As the name im-
plies, an outstanding characteristic of the toothed whales is teeth. They feed on sea animals. Baleen
whales feed on groups of animals by straining water through a series of transverse triangular horny
plates with soft bristles, an apparatus attached to each side of the upper jaw that acts as a sieve to filter
planktonic organisms, mostly small crustaceans such as krill and small fishes.
58 Part One Living Resources

Sperm whale, Physeter macrocephalus (a toothed whale), is cosmopolitan in distribution, and are
usually found between latitudes 400N and 400S. The sperm whale is large, having a maximum length
of 60 ft (18.3 m). The upper jaw has from 18 to 30 teeth. A single calf produced once every 3 years will
measure up to 14 ft (4.3 m). The mother nurses its young for 16 months. Whales eat mainly large
squid, 30 to 40 in (76.2 to 102 cm) long, cuttlefish, sharks, fish, and octopuses. Their large size keeps
them relatively safe from most predators. The killer whale, Orcin us orca, is strongly suspected of prey-
ing on other whales, but evidence for this is not well-substantiated under natural conditions. Killer
whales prey on large adult whales being towed to a factory, or when moored in the water prior to being
hauled out. They also have been observed attacking calves. Sharks are logical suspects in whale pre-
dation. The total effect of this type of natural mortality on whale stocks is not well known.
Fisheries: Whales were caught formerly by hand-held harpoons from skiffs, and later, by harpoon
guns from catcher boats. They were heavily hunted from the 18th century until after World War II.
Large males were selected by the fishing fleets that seems to have reduced the average size of sperm
whales by genetic control.
Sei whales, Balaenoptera borealis (a baleen whale), have worldwide distribution in all oceans.
Highly migratory, they travel long distances north-south from higher latitude summer feeding grounds
to lower latitude winter areas. The sei whale is as large as the sperm whale, about 60 ft (18.3 m) in
length. It may have conspicuous white spots, but its coloration is rather variable. A sickle-shaped dor-
sal fin is characteristic of the species. When they blow, the water spout may rise as high as 8 ft (2.4 m)
into the air. The sei is capable of swimming at speeds of 20 mph (32.2 km/h). The mating season lasts
about 5 months during the winter. Usually a single calf is born after a gestation period of 1 year. Calv-
ing occurs every 2 or 3 years. Populations north and south of the Equator are believed to be separate.
In contrast to the sperm whale, which eats large foods, the sei whale strains small planktonic organisms
from the water, mainly small crustaceans like copepods, and in the southern ocean, krill. Predators are
the same as for sperm whales.
Fisheries: As with other commercially important whale stocks, the sei whale has been overexploited.
Years ago, the baleen plates were split for use as supports in ladies' corsets. They are caught as de-
scribed above.
Northern fur seal, Callorhinus ursinus, are found in
the North Pacific Ocean, and the Bering, Okhotsk, and
Japan Seas (Fig. 3.22). Tagging studies have shown that
they may swim 2,000 mi (3,218 km) south to the Mexi-
can border. Male and female northern fur seals can be
distinguished by several characteristics. Males are sub-
stantially larger than the females and may reach a
length of 8 ft (2.4 m) and weigh up to 600 lb (272 kg).
They are dark brown as adults and emit a sustained,
deepthroated bellow. The females reach a length of
about 5 ft (1.5 m) and weigh about 130 lb (60 kg).
Ashore they are dusty brown that turns silvery-gray in
the sea. Females emit a sheeplike bleat. Sexual matu-
rity is reached at an average age of 3 years for females
and 4 to 7 years for males. Usually the older, large bulls
Figure 3.22 Seal. about 7 to 8 years old are the only ones that can take an
active part in the mating and breeding. Mating and
breeding takes place ashore in rookeries. Harems average about 40 females to each male able to assert
his dominance. Harem size may range from as few as 10 to as many as 100 females. Each female or
"cow" gives birth to a single pup after a gestation period of about 1 year. The mother nurses her pup
for about 3 months, going into the sea to feed, then returning to her pup. After this time, she returns
to the sea, leaving the pup to fend for itself. Maximum life can be 20 years, perhaps longer. Because
seals are air breathers, they occur most frequently at or near the surface. Northern fur seals feed on
squid and small schooling fish such as herring and capelin. Fishermen have accused seals of eating
Chapter 3 Major Resource Organisms-Vertebrates 59

large numbers of salmon, thus reducing their catches. However, studies of stomach contents of the fur
seal have shown this to be false even in areas where salmon are heavily concentrated. Predators on seal
pups are sea lions, Eumestopias jubata. However, the impact of predation by sea lions or other predators
on fur seal stocks in unknown.
Fisheries: For many years seals were captured in rookeries for their valuable fur by fishermen using
rifles or clubs. The northern fur seal has the finest fur, but sea otters and the Guadalupe fur seal are
also killed for their fur. Other seals are killed for oil and for their hides for leather. Some seals, mainly
the California sea lion, are trained for circuses and ocean aquarium acts. Fishermen also hate seals be-
cause they eat salmon corralled in their nets, destroy their nets, and scare schools away from their boats.
Some seal species are final hosts for parasites found in desirable fish such as the salmon; hence, reduc-
ing the size of seal stocks can reduce the number of parasitized fish brought to market.


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Allen, G. R. 1985. FAO species catalogue. Vol. 6. Snappers of the world. An annotated and illustrated
catalogue of lu~anid species known to date. 208 pp.
Audubon 1975. Special issue on whales and dolphins. 77(1). 108 pp.
Baker, R. c., F. Wilke, and C. H. Baltzo. 1970. The northern fur seal. U.S. Fish. Wildl. Serv. Cire. 336,
20 pp.
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137 pp.
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Cushing, D. H. 1975. Marine ecology and fisheries. Cambridge University Press, Cambridge, England.
Dadswell, M. J., et al. 1984. Synopsis of biological data on shortnose sturgeon, Acipenser brevirostrum
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Everson. I. 1977. The living resources of the southern ocean. FAO (F.A.O.U.N.)/U.N. Dev. Programme.
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(Fishing Area 31) Vols. 1-5 Fishes. FAO Rome (pagination not continuous).
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60 Part One Living Resources

Gulland, J. A. (ed.). 1970. The fish resources of the ocean. FAO Fisheries Technical Paper No. 97.
425 pp.
Gunderson, D. R 1993. Surveys of fisheries resources. John W. Wiley & Sons, Inc., Somerset, NJ.
Gusey, W. F. 1978. The fish and wildlife resources of the Gulf of Alaska. Shell Oil Co. 580 pp.
Hart, J. L. 1973. Pacific fishes of Canada. Fish. Res. Bd. Canada, Bull 180. 740 pp.
Jory, D. E. and E. S. Iversen. 1989. Species profiles: Life histories and environmental requirements of
coastal fishes and invertebrates (South Florida). Black, red and Nassau groupers. Biological Report
82 (11.110). National Wetlands Research Center, U.S. Fish and Wildlife Service, Slidell, LA. 21 pp.
Joseph, J., W. Klawe, and P. Murphy. 1988. Tuna and billfish-fish without a country. Inter-American
Tropical Tuna Commission, La Jolla, CA. 69 pp.
Kenyon, K. W. and V. B. Scheffer. 1955. The seals, sea-lions, and sea otter of the Pacific coast. U. S. Fish
Wildl. Servo Circ. 32. 34 pp.
Lane, J. P. 1978. Eels and their utilization. Mar. Fish. Rev. 40(4):1-20.
Leim, A. H. and W. B. Scott. 1966. Fishes of the Atlantic Coast of Canada. Fish. Res. Bd. Canada, Bull
155. 485 pp.
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State Museum of Natural History. Raleigh, N.C. 376 pp.
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Palko, B. J., G. L. Beardsley, and W. J. Richards. 1981. Synopsis of the biology of the swordfishXiphias
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decades of change. International Center for Living Aquatic Resources Management (ICLARM). 351
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Chapter 3 Major Resource Organisms-Vertebrates 61

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Chapter 4

Life Histories
of Resource Species


I have chosen to discuss separately the history of fisheries research from two aspects of fish-
eries that rely on basic research: fish population dynamics and fisheries management. Fish-
eries research encompasses effort directed to obtaining knowledge on all life history aspects
of commercial and recreation fish and shellfish life histories such as mating, spawning, early
development, food, feeding habits, age estimation, growth, extent of migration, survival, ef-
fects of parasites, and diseases.
Ecological studies, including the role of the environment and prey and predatory species,
are also necessary for rational management. An important advance in effective fisheries re-
search was made when we learned how important studies on the total ecology of exploited
species are. Fisheries research provides information that enables population dynamists and
fisheries managers to make educated decisions on the best utilization of fish stocks.



The voyage of the Endeavor in 1768, an expedition led by Captain James Cook, brought fishes
home to England from many areas of the ocean to be studied, classified, and cataloged. The
science of oceanography was born later, between 1850 and 1900. The HMS Challenger Expe-
dition (1872-1876), traveled over 70,000 miles of ocean, led by pioneering Englishmen whose
curiosity drove them to expand their knowledge of the seas. On this expedition, basic phys-
ical observations (for which equipment and techniques were then available) on such param-
eters as temperature and salinity were made that gave great impetus to pursue studies of the
world's oceans, including its physical features, biological aspects (both fauna and flora), and
the benthos, plankton, and nekton (swimming organisms). Fish, fish products, and fishing
gear from around the world were shown for the first time at the 1883 International Exhibition
in London.
Despite increasing concern about overfishing of natural stocks, many continued to be-
lieved that this was a myth. One person with considerable influence at the time was a sur-
geon, Dr. Thomas H. Huxley, who lived from 1825 to 1895. Huxley, who was interested in
science and was a strong supporter of Darwin and his theory of evolution, proclaimed:

Chapter 4 Life Histories of Resource Species 63

1. I believe that the cod, herring, pilchard, mackerel and probably all great sea fisheries are inex-
2. Nothing man does seriously affects the number of fish.
3. The reproductive potential is so very high that any mortality caused by fishing is insignificant.
4. Therefore, any attempt to regulate the fisheries would seem to be useless.

These observations, no doubt were fitting in Huxley's time, but as we look back to those
days with brilliant hindsight, several faults leap out from the arguments he espoused and
that others believed, perhaps because of the attitudes of their day.

1. They did not foresee the increased technology that would become available to future fishermen,
such as more efficient gear, more efficient vessels, and, most importantly, more efficient vessel
2. The concept then was that the "great and wide sea" had limitless resources. We now realize
that our vast oceans are not uniformly productive; for example, about 90% of our fishery pro-
duction comes from continental shelves alone.
3. The production of extremely large numbers of eggs by a Single fish or shellfish may have mis-
led early fishery experts to believe the reproductive potential of pelagic spawners was so great
that fishing pressure on ocean resources appeared to them to be insignificant. We have since
learned of the extremely high mortality between the egg stage and adult that occurs in the
ocean, where survival is less than one-tenth of 1 percent for many species.

1884: The Marine Biological Association was formed at Plymouth, England. Scientists
begin considering fish as populations rather than as individual fish, a major step in
the formation of the science of fishery biology.
1892: C. Hoffbauer (Germany) studied scales and bones to determine age.
1893: T. W. Fulton (Scotland) studied sea fish catches and effort.
1898: C. J. Petersen (Denmark) used tags on fish to study growth, migration, and mortal-
The science of fishery biology was not available before the 1900s. Any study concerning
fisheries depended much upon one's personal knowledge of marine biology and oceanogra-
phy, because books on these subjects were also unavailable. For fish stocks to be properly
managed in those days, it was necessarily to have input of several kinds:

1. collection of specimens from the sea, their description and classification;

2. life history and ecological studies such as growth rates, age, methods of reproduction, and the
interrelationships between species, individuals, and their environment;
3. acceptance of the concept that stocks needed management, that is a conservation movement
based on biological aspects of conserving the stocks of fish and shellfish in terms of yield in
weight or numbers of fish. Management, based on the greatest economic yield from marine
fisheries, was to come much later.

In the latter part of the 19th century, and very early 20th century, the pendulum began to
swing toward conservation. This trend was partly due to decreasing catch rates, despite fish-
ermen working harder and fishing in new areas.
By 1900, there was mounting evidence that certain fisheries were declining. After patterns
of decrease in total catch and catch per unit of effort, fisheries research came into being. In-
64 Part One Living Resources

terest was so strong that in 1902 the International Council for the Exploration of the Sea was
formed in Sweden.
Across the Atlantic, in 1908 Canada established biological stations on the Atlantic coast at
St. Andrews, New Brunswick, and on the Pacific coast at Nanaimo, British Columbia, begin-
ning what developed into high quality research in Canadian marine biology.
In 1914, J. Hjort in Norway, published "Fluctuations in the Great Fisheries of Northern Eu-


In 1871, the U.S. Fish Commission carried out scientific research on the vessels Blake, Fish
Hawk, and Albatross. Then, in 1903 it became the U.S. Bureau of Fisheries, under the Depart-
ment of Commerce, evolving still further in 1940, when it became the U.S. Fish and Wildlife
Service under the Department of the Interior. Since 1970 it has been known as the National
Marine Fisheries Service, returning to the Department of Commerce, and was given respon-
sibility for marine and anadromous fish species.
As one scans this brief history an evolution becomes apparent. By the early 1900s, evi-
dence of decreases in total catches year after year suggested overfishing of the stocks in cer-
tain fisheries causing concern in important fishing countries. The question nagged whether
fishermen were harming the stocks of fish by overfishing because of improvements in fishing
methods, gear, and vessels, as well as by increased numbers of vessels. The old belief that
the supply of sea fish was unlimited now began to appear to be wrong, or at least very ques-
tionable, and about the tum of the century it became apparent that the question had to be ad-
dressed. Fisheries science was born.


Aspects of fish and shellfish reproduction important to fishery biologists include: the effects
of fishing on spawning stocks, size, and age of fish at first spawning, how mating takes place,
how young are produced, kinds of eggs and their identification, the extent of care afforded
young, time of spawning during the year, frequency of spawning within a year by a single
female, geographic location of spawning, and methods used in reproduction studies.
To perpetuate their species, of course, fish and shellfish must be allowed to grow to matu-
rity, mate (except for hermaphroditic species), and spawn. Information gathered from age
and growth studies is valuable, together with size at first maturity, to insure that a fishery
does not remove a high percentage of spawners from a stock before they have spawned. As
was discussed earlier, large, slow-growing, late-in-life maturing animals are the most suscep-
tible to this danger. Some whale species, for example, may be large enough for harvest but
still may not have reached maturity and reproduced. Whale stocks are very sensitive to de-
struction from the technology fishing methods used today because of the few offspring each
female bears (some species produce only about 10 during their life) and of their slow growth.
One must understand all aspects of reproduction of exploited stocks of aquatic animals to de-
velop a management technique in order to obtain maximum sustained yields of any particu-
lar fishery (Fig. 4.1).
When we discuss management and regulation later in the book, it will become apparent
why the effects of fishing on stocks of spawning fish must be controlled in certain fisheries.
Chapter 4 Life Histories of Resource Species 65



Figure 4.1. Life history of crab. The two

larval stages are planktonic and their survival
is dependent on location to which the currents
carry them. From Goode (1884). U.S. Bureau
of Commercial Fisheries.

In earlier discussions on gear and fishing methods it was pointed out why large schools are
necessary to make fishing economical. These schools may consist of spawners or fish migrat-
ing to the spawning grounds. A good example of the result of overfishing is the Pacific
salmon, Oncorhynchus spp., fish that (with the exception of the Japanese who fish them on the
high seas), are caught by Americans and Canadians a short distance from their spawning
grounds. Another is the Pacific herring (Clupea harengus) that is fished during its spawning.
Before discussing the various aspects of reproduction, a few definitions are in order. In
general, reproduction is when part of the parent's body is segregated by sexual or asexual
means into a new individual. Spawning describes reproduction in aquatic animals that pro-
duce or deposit eggs, usually in large numbers. The eggs of aquatic animals may be collec-
tively referred to as spawn. Mating is when males and females join together for the purpose
of reproduction that may involve courtship, then copulation.
Finfish employ a truly amazing variety of reproductive behavior patterns or strategies,
each of which is apparently directed at survival of the species and is balanced against energy
cost. As a matter of interest, fishes have been classified according to their reproductive strat-
egies. Most unusual strategies are used by freshwater fish and include such behavior as se-
lecting certain types of substrate for spawning, nest spawners, guarding young, mouth
brooders, pouch brooders, live bearers, and brood hiders. Catadromous fishes leave freshwa-
ter and go to sea to spawn, and, conversely, anadromous fishes leave the sea and go into
freshwater to spawn (Fig. 4.2).

Size of Fish at First Spawning

As mentioned earlier, it is important to allow a portion of fish stocks to spawn to replace
losses from fishing and natural mortality. If fishing is heavily concentrated on immature
fishes, serious harm can befall the stocks.
66 Part One Living Resources

Figure 4.2. Life cycle of chum salmon. From

Merrell (1970). U.S. Fish and Wildlife Ser. Fish.
Leaflet No. 632.

To determine the approximate size at which fish first spawn, samples from a stock must be
measured over a wide range of sizes and the stages of maturation noted. A curve represent-
ing accumulated percent of either females or males (usually females give best results) plotted
against body length will show an S-shaped (sigmoid) curve. This curve reveals that all fish
do not mature at some particular body length, but rather only a small percent do at small
sizes. Maturity increases as the body length increases, up to 100% of the females. In prac-
tice, the body length at which 50% of the females are mature is usually used to indicate the
average size and age at first maturity (see Fig. 4.3).

Mating Procedure
Marine fishes segregate into pairs or congregate in large schools during mating. Salmon
typically will pair together when nest building and the male will defend the nest and its area
for the brief remainder of his life. Fishes that spawn in large schools such as mackerel, cod,
herring, etc., release sex products in response to those released by the opposite sex within the
school. When the spawning process is completed, the adults go on their way, leaving the fer-
tilized eggs and young on their own.
In most populations the ratio of males to females is about equal, but uneven sex ratios do
exist. In some cases, the sexes may be separated most of the year and come together only
during the spawning season. Male soupfin sharks, Galeorhinus zyopterus, occur in the north-
ern part of their range and females in the southern part, except during the spawning season.
This occurs in invertebrates as well; blue crab males may stay well up in the upper reaches
of an estuary, but the fertilized females go down to high salinity to spawn.
Chapter 4 Life Histories of Resource Species 67

10 e e._--

~ ~--------------------~
Figure 4.3. Determination of size of yellowfin
sole females at first maturity, when 50% are
mature at about 10 in (253 mm) total length.
Age at first maturity can be calculated from a
length/ age relationship. Modified from
Wakabayashi (1974). Japanese Fishery Agency,
ISO 300 3SO
Far Seas. Fish. Res. Lab. Unpublished
Manuscript. Total length in mm

Unequal sex ratios may also be due to sampling or spawning migration, or to the effect of
fishing; for example, some fishing gears such as gill nets seemingly select one sex over the
other when it is actually the larger size of one sex that gets caught. Northern shrimp (pan-
dalids) metamorphose into females at larger size and the fishery harvests them almost exclu-
As we have seen, size differences occur simply because one sex grows faster than the
other. Also, body shapes between sexes can be different as in the dolphin fish where the
male has a straight forehead and the female's slopes. Each sex may be colored differently, as
in certain reef fishes. Fishes that fertilize internally require that the male have a copulatory
organ. Some sharks and rays and in livebearers, Gambusia spp., are examples. The deep sea
angler fish, where the tiny male is attached to the head of the female, exemplifies the extreme
size difference between sexes in certain marine fish. For some species that show no obvious
outward signs of sexual dimorphism, the only way to determine the sex of these species is to
cut the fish open and examine its gonads.
Fishery scientists frequently need estimates of the numbers of each sex in the population
of fish they are studying (Fig. 4.4). Occasionally shifts in sex ratio can indicate overfishing on
one sex or the other due to selective gear and/ or differential growth rates in both length and
weight between sexes. To obtain data on sex ratios in fish stocks where the fish are landed
whole and the sexes show sexual dimorphism is relatively easy. However, when species are
gutted at sea this information is difficult to obtain because of the loss of external genitalia
and gonads. The Pacific halibut, Hippoglossus stenolepsis, fishery is an example of the latter
fishing practice.
It was found that otoliths of males and females of halibut differ somewhat in weight, but
also, they appeared to be shaped differently. Using a computer program, otolith images were
digitized, using Fourier shape descriptors, and were analyzed together with their weights.
However, even then, successful classification of sexes reached only about 75 percent.
Otolith shapes have been found useful in fish stock identification (see Chapter 6).
Selective fishing for a single sex may be a useful management tool. If it is known that one
male mates with more than one female, the excess males can be harvested without reducing
the reproductive potential of the species. A study of Alaskan red salmon found that a single
male could mate with up to 16 females. Sex ratios from 1:1, up to 30 females to 1 male, were
68 Part One Living Resources

Figure 4.4. Research scientists determining

the sex of Pacific cod, Gadus macrocephalus,
collected during a resource assessment of the
Bering Sea. Photo D. Smith, Alaska Fisheries
Center, NMFS, Resource and Fisheries
Management Division.

tested. This management technique of protecting females while allowing fishing on males is
used on whales, fur seals, and berried (attached eggs) female lobsters.

How Young Are Produced

One category, where eggs released by the female are fertilized and develop outside of her
body, is called oviparous reproduction (Fig. 4.5). Spawning in dense schools, as with herring
and sardines, is believed to insure fertilization of the majority of the eggs. Other species, like
salmon, freshwater bass, and sunfish, build nests for the eggs, and the fish mate in pairs, pro-
tecting the nest and its surroundings. In some cases, the early stages of young are protected
as well. Tilapia, Sarotherodon spp., allow their young to swim into the mouth of one or the
other of the parents (depending on species), when danger threatens. This behavior is called
"mouth brooding."
The second spawning category refers to when the female is fertilized internally and the
eggs develop inside her body. This is termed ovoviparous spawning. The young are born
alive, but during their stay with the mother, nourishment comes from a yolk sac and not di-
rectly from the body of the female. Ocean perch reproduce in this manner (Fig. 4.6).
The last spawning category, called viviparous, is similar to the ovoviparous method, but
differs because the young receive nourishment directly from a vascular connection not con-
sidered to be a true placenta. Guppies, halfbeaks, sharks, and surf perches are all examples
of fish that reproduce in this manner.

Kinds of Eggs and Their Identification

A biologist must be able to identify eggs of various fish species in order to determine
spawning season, location, and spawning success. Most fish eggs are spherical or subspher-
ical, but some are elliptical (anchovies) or oval (gobies). The eggs of species such as skates
and hag fish have tendrils. Most marine fish eggs are pelagic and therefore are subject to dis-
persal by currents, but some are able to attach to seaweeds, dock pilings, and the like, as
with the herrings. Demersal eggs are those that remain near the bottom of the sea.
Eggs are generally identified on the basis of their size and shape, and, in some cases, on
the amount of yolk material present.
Chapter 4 Life Histories of Resource Species 69

cO ~ 'lbUI19 halibut
Late pasllanlll 1.38 inch
1.00 inch

~ Mid-postlarw
.88 inch


Figure 4.5. Life history of Pacific halibut, a

pelagic spawner. From the International Pacific
Halibut Commission.

Egg Size
Egg size of fishes is important for survival in the wild (and in some types of aquaculture)
because size influences the development, growth, and activity of larvae until they begin ex-
ogenous feeding. Among 40 marine and freshwater species, those with the largest eggs pro-
duced larger larvae at the onset of feeding. Large larvae can avoid predators because they
can swim faster; their food needs may be satisfied more readily because they can capture
more and larger prey. They can survive food shortages longer than small larvae. In aquac-
ulture facilities, fishes with large eggs are generally much easier to care for during larval
stages and usually have a higher survival rate than those species with small eggs.

Care Afforded the Young

The kind and extent of care afforded the young fish and shellfish is important in manage-
ment practices. As mentioned above, marine fish that cast their eggs and sperm into the sea
offer no parental care. As might be expected, eggs and young from such spawning suffer ex-
tremely high (mass) mortalities. Anadromous fishes like salmon, which build nests and pro-
vide some care and protection for their eggs and yoke sac young, provide them a better
survival chance than the young of the pelagic spawners get. The small freshwater stickle-
back fish makes a nest in the sand and cements it together with mucus from its buccal cavity.
The male chases the female into the nest, fertilizes her eggs, then guards the nest against in-
truders. Male seahorses and pipefishes protect the small number of eggs laid per year, about
200, in a pouch on the abdomen in which the females places the eggs. Some of these are un-
usual examples, but show the wide care given.
70 Part One Living Resources

Figure 4.6. Some elasmobranchs bear their

young alive. Longline caught pregnant shark
from the Central Equatorial Pacific. Photo by


Fecundity is generally defined as "capable of producing offspring in abundance." Fishery

scientists qualify and make this definition more exact: "Fecundity for egg bearing females is
the number of eggs being readied for the next spawning in the body of the female." This
number may be relatively easy to determine if the species has a contracted spawning period,
or season. A female that releases all of the eggs for one season at one time is called a "total
spawner." On the other hand, the fecundity (number of eggs) released may be hard to deter-
mine for a "partial spawner"; for example, fishes that spawn smaller numbers of eggs on
several occasions throughout the year, are partial spawners.
Relative fecundity is often used as an index of the capability of production and is the
number of eggs per unit of weight of the female. The total number of eggs laid during the
entire life of the female is called "total fecundity."
There are apparent reasons for the wide range in fecundity found among various species
and within species. The hazards and hardships faced by the eggs and young stages corre-
lates with the fecundity of the species. The greater the mortality of the species caused by a
harsh environment, the higher the fecundity; therefore, those species that release eggs free
Chapter 4 Ufe Histories of Resource Species 71



~ ~o
~ '
, '
.. 40

~ 30
...o ,
20 , '
~ 10
Figure 4.7. Relationship between fecundity
and fish length. From MacGregor (1957). U.S.
170 180 190 200 210 220 230 240 2S0 260 270
Fish and Wildl. Servo Fish. Bull. 121:427-449. STANCARD LENGTH IN MILLIMETERS

into the sea and provide no protection or parental care to their eggs and young will lose
countless numbers of young to the vicissitudes of nature. Conversely, those species that pro-
vide parental care, such as the salmon and sticklebacks (nest builders), the tilapia (mouth
brooders), and the mammals such as whales and porpoises (suckle and protect their young)
have very few offspring during a breeding season and throughout their lives.
Four general patterns have emerged from fecundity studies.

1. Fecundity is approximately inversely related to the sizes of eggs and care given to eggs and lar-
val stages of fish.
2. Fluctuations in abundance are greater for species with high fecundity and conversely.
3. Within the range of ova laid by a species the fecundity increases (positive correlation) with the
size of the female (Fig. 4.7). Studies on the Pacific sardine, Sardinops sagax, showed that the
number of eggs produced per batch was more closely related to fish weight than to length or
age. In fact, the number of eggs produced is almost directly proportional to fish weight. The
typical relationship between fecundity and fish length shows exponential relationships, for ex-
ample, tunas.
4. Fecundity is related to mortality imposed by the environment; for example, even within a spe-
cies, variation may exist from one race or stock to the next. This is evident for females from
salmon streams in Alaska that have different ranges of fecundity depending on the environ-
mental conditions affecting survival of their offspring in the home stream.

Fecundity information is used by fisheries scientists to better understand the relation be-
tween the survival of a species and its environment. Natural fluctuations in abundance (see
Chapter 7) can be explained in part by the fecundity of a species. Species of low fecundity,
such as sharks and whales, can be biologically overfished more readily than those of higher
From the standpoint of providing maximum recruitment to a fish stock, it is important to
determine if fish, or mammals, should be undisturbed during spawning periods. Actually
most pelagic spawners are harvested most heavily when they are schooled for spawning.
Table 4.1 lists the fecundities of some marine fish and shows dearly the considerable var-
iation between species.
72 Part One Living Resources

Table 4.1 Examples of Variation of Fecundity Among Species

Species Fecundity Range

No. of Ova, or Young/Female/Season"
Northern (Maine) lobster, Carry, eggs: 3,000-100,000, on swimmerettes
Homarus americanus
Salmon, Oncorhynchus sp. Nest builders, 2,000-8,000, in streams
Tilapia, Tilapia, Sarotherodon Mouth brooder, 75-250 each spawning, may spawn 6
Shark, Galeorhinus australis Bear young alive, 17-41
Fur seals, Callorhinus ursinus Bear young alive, l/year
Whales Bear young alive, 10 young during 30-year life span
Oyster, Crassostrea virginica 14-114 million
Giant clam, Tridacna 500 million/day
Cod, Gadus morhus 2-9 million
Plaice, Pleuronectes platessa 16,000-700,000
Atlantic herring, Clupea harengus 20,000-40,000
Dolphin fin, Coryphaena hippurus 80,000-1 million/spawning, spawn 2 or 3 times per year
Bluefin tuna, Thunnus thynnus 10 million
• Fecundity varies with size of female and geographic location.

Frequency of Spawning per Season

By a Single Female
In some fish species, spawning by an individual female takes place in a short period of
time (within a few months). This is especially true where mass spawning occurs. In other
species, a single female may spawn several different times during a season, releasing small
batches of eggs at intervals. The number of spawning characteristic of a species can be de-
termined by measuring egg diameters of females and plotting the frequency of various egg
diameters against time in months (Table 4.2). In total spawners (a single big spawning per
season) a group of eggs will break off from the ovarian pool of developing eggs and grow
until they are readily to be released. There usually will be a definite valley or space between
the pool of small eggs that will be spawned during the remainder of the females life and the
ripening eggs that will be spawned during that spawning period.
The eggs of many partial spawners may be measured to locate the number of modal
groups representing groups of eggs to be spawned during that season. In contrast to the to-
tal spawner, groups of eggs are seen to increase in size and are released together when the
optimum size is reached. Dr. F. Clark, in her 1934 study, suspected that the Pacific sardine
was a partial spawner, and proceeded to study the problem in the following way.
She found a multiplicity of modes of egg sizes (diameters) suggesting that individual sar-
dines spawn more than once in each season when perhaps three batches of eggs are released.
To demonstrate partial spawning by the sardine she had to show that the secondary modes
did not represent eggs to be spawned in subsequent seasons, or eggs that would be resorbed.
Evidence for this was that from September to October in the species studied, only immature
eggs were present. She found a multiplicity of modes of egg diameters in her size-frequency
Chapter 4 Life Histories of Resource Species 73

Table 4.2 A Five-point Maturity Scale for Partial Spawners.

Stage State Description

Immature Ovary and testis about 1/3 length of body cavity. Ovaries
pinkish, translucent; testis whitish. Ova not visible to naked
II Maturing virgin Ovary and testis about 1/2 length of body cavity. Ovary
and recovering pinkish, translucent; testis whitish, more or less
spent symmmetricai. Ova not visible to naked eye.
III Ripening Ovary and testis is about 2/3 length of body cavity. Ovary
pinkish-yellow color with granular appearance, testis whitish
to creamy. No transparent or translucent ova visible.
IV Ripe Ovary and testis from 2/3 to full length of body cavity. Ovary
orange-pink in color with conspicuous superficial blood
vessels. Large transparent, ripe ova visible. Testis whitish-
creamy, soft.
V Spent Ovary and testis shrunken to about 1/2 length of body cavity.
Walls loose. Ovary may contain remnants of disintegrating
opaque and ripe ova, darkened or translucent. Testis
bloodshot and flabby.

distributions that showed a strong correlation with the growth of the successive groups of
eggs. As the breeding season progressed there was a decrease in the numerical ratio between
successive batches of eggs and the largest size groups. The evidence suggested to Dr. Clark
that the California sardine is a partial spawner. Later studies on the number of batches of
eggs per female per year were not in agreement with Clark's results which might mean that
the number of eggs spawned per batch is not a good indicator of number of eggs spawned
per season.
Studies like this are difficult because egg growth cannot be studied from live fish. Large
samples of eggs from many deceased fish must be collected at different times of the year and
carefully measured for proper conclusions to be drawn.

Time of Spawning

The length of time fish and shellfish spawn and at what time of year, are important in de-
termining when to permit fishing and when to restrict fishing to protect spawners. The con-
dition of both females and male gonads indicate when spawning takes place, as is shown in
maturity criteria for both sexes. Adult fish must be examined yearround for changes in go-
nadal condition to determine the beginning, peak, and ending of the spawning season.
Time of spawning is extremely variable between species; however, some general trends are
apparent. Fishes in cold waters spawn mostly during autumn and winter, which is believed
to enable their young to reach a sufficient size to take advantage of periods of abundant
foods. As a rule, warm water species spawn year around.
To illustrate the degree of variation in spawning time, let us consider the herring, a clupe-
oid. Cold water herring spawn yearround; however, different races spawn at different times
of the year, spring spawners, summer spawners, etc. The possible increased survival value
74 Part One Living Resources

resulting from different spawning times versus a single protracted spawning season is ob-
scure but may be related to competition for food by the larval herring.. If the entire species
spawned at one time and food were in short supply, herring mortality would be disastrous
to the species. Another possible reason is that massive spawning in a single location could
attract many more predators than spawning by different races at different seasons. Herring
may also hedge their survival bets by increasing the spawning period to cover a wider vari-
ety of environmental conditions.
At the other extreme is the grunion in southern California which spawns only on the full
moon during the highest tides. A very restricted set of environmental conditions governs the
time when these fish spawn, and the success of spawning.
Important from the standpoint of species survival, time of spawning requires that it coin-
cide with favorable growth conditions. When below optimum environmental conditions
such as cold water temperature, and/or scarcity of food, slows the growth of the young of a
species, more time is needed to pass through the larval stages, at a period in their lives when
they are highly vulnerable to predators. Survival generally will be much higher when the
young find conditions optimum for rapid growth.

Place of Spawning

The spawning place of a stock of marine fish can be done with the help of fishermen, who
recognize fishes heavy with roe in their catches and/or may actually see running ripe fish (a
condition where eggs or milt may be released when the fish land on the deck of the fishing
vessel or can be expressed with slight pressure on the abdomen).
Locating the place and time of spawning to give more information on time of spawning
for marine fish requires expensive oceanographic surveys. Collecting samples from a multi-
tude of stations sampled periodically at different depths and to search all the plankton mate-
rial collected for eggs and larvae is a massive and tedious undertaking. An undesirable
aspect of this work is that it adds greatly to the expensive nature of this type of research. It
has to be done throughout the year to establish time of spawning, and requires the use of a
station grid that covers an area much larger than the suspected spawning grounds. Without
these requirements, the data collected would be of little value to establish the spawning area.
An extensive study was done for the California sardine in which plankton samples were col-
lected and examined for eggs and larvae of sardines and other fish species year around over
a large geographic area off the California coast (Fig. 4.8).
From a recruitment standpoint, information on time and spawning location for a species
makes it possible to restrict fishing during certain times of the year and in certain locations
to prevent overfishing on the spawning stock. Furthermore, it tells investigators the environ-
mental conditions during which the species spawns. If enough data are collected over a
number of years, optimum spawning conditions will be better known and will help explain
the variation in recruitment that occurs between years.


From the sections above it can be seen that studies must be undertaken to count eggs to de-
termine certain aspects of reproduction; that is, fecundity by species and size of fish and for
pelagic spawners that release hundreds of thousands or even millions of eggs require the use
of sampling techniques. The formula to determine fecundity is as follows. The number of
Chapter 4 Life Histories of Resource Species 75

... ... "",



" .0

..,. ••

'- "..t

>II •• I
,;,.0 • l
"t •
• "" . .. .
• .;IP. •• ..
• • II!.

. .". . . ..
-"". f//t.
"'" ~.
/' ...
...::'" io\
• . "..
4~ • . ..
/ . toO· ..

• "..
• • • 4·

...f .,'- • ~".'~P"'!.:--.~
,'f .~,,~ tP.:.... •
. ." ... .
4-. •••
• ...
/,; /J>.

•• / •• .•oJ!!.
II! •• / . .

• • - / "' • .!'I'"0
~f / •• ,.tIl" • ••
. . .
.".f •
SOUTHERN CALIFORNIA .....".. oC!~. • •
.....' . WI!. -'! •• ••
/ ~ •• 01.1 •• / •• -
#. .".. ........
,. .• ,rP~ ""0 . " . .• / •
1! ••
.,.,. .
.p'....... ,. •
II! •
•• ok
• IP..

." •
4 • ....f •• tI.!~~ ••
/"'. .".. "-llll·,·

/ ~~
•• ..tP. ••
....0. til· •

" • • •~,.<!.
•• If'i.
• ••....." ••
...;"\. .,...r.. ••
--'.. .. ..
#. • •
./ '\,o!
/ ~. •• ,.<! • • •


• • ~•

• ••
~''f •

.. " ... .... ...

Figure 4.8. Sardine egg and larval sampling station grid off California and Baja, California. From
California Cooperative Oceanic Fisheries Investigations Reports. Vol XI. 1967.
76 Part One Living Resources

eggs in a sample of known volume or weight is determined and the volume or weight of the
whole gonad is determined. The number of eggs in the sample is multiplied by the propor-
tion the sample is of the whole. If, for example, the portion of the ovary sampled has 1,000
eggs by actual count and the sample represents 1/10 of the total weight of the ovaries, the
fecundity of the fish of that species and that size would be estimated at 10,000. Because the
ovaries of most fish are bilobate, the number would usually double if the weight of both
lobes is similar.
Egg diameters must be measured, as indicated earlier, to determine the number of spawn-
ing within a single season for a certain species of fish. For large eggs this is not difficult, but
for very fecund fish with tiny eggs magnification of the ovary is necessary. In some cases,
tissue sections of gonads are made to measure egg diameter under a microscope. The impor-
tance of spawning studies should be rated on a level with fish growth measurements.


The number of fish of a single year class entering the exploitable phase of a stock in a given
period through growth is called recruitment. Substantial numbers of fish may also enter the
exploitable phase from outside the fishery (immigration), which can seriously bias the suc-
cess of the parent-progeny relationship. The success of the breeding stock to provide young
to the fishery determines the subsequent yield from a fishery. Reproductive success is very
important for short-lived species with few age classes in the fishery, because high egg and
larval mortality of more than one spawning can greatly reduce the abundance of fish availa-
ble to the fishery.
Recruitment processes are complex, and attempts to describe these relationships and ac-
count for the wide variation seen in stock-recruitment relationships are not very convincing.
This wide variation has led to the statement that "no relationship exists between stock and
recruitment." Possibly some of this variation between numbers of parents and numbers of re-
cruits may lie in the difficulty of accurately measuring both stock size and numbers of re-
cruits over wide areas of the sea. This failure to satisfactorily describe the relationship has
been called one of the big unsolved problems in fisheries science today.
Generally two types of relationships between the size, of the parent stock (numbers) and
the numbers of recruits are suggested. An asymptotic relationship (Fig. 4.9) between the
size of the spawning stock and the number of recruits has been explained by the shortage of
suitable sized food for large numbers of recruits.

c 3 r-----------r----------,r----------.

Figure 4.9. Returns (corrected) on number of
18 sockeye salmon spawners for the 1929-1948
periods. The limiting factor on the number of
recruits (asymptotic relationship) may be
availability of suitable sized food. From
Rounsefell (1958). Fish. Bull. 53:83-169.
Chapter 4 Life Histories of Resource Species 77

The dome-shaped relationship (Fig. 4.10) of number of recruits is believed to be the result
of heavy cannibalism by adults on young, on the high density of young, and on the time lag
between predator response to prey density. As the size of the spawning stock increases, so
does the number of recruits up to a dome. Then, with increases in the size of the spawning
stock, the numbers of recruits decreases (Fig. 4.11).

Figure 4.10. Effect of environment and

spawning stock size on year-class production:
curve A represents average environmental '"
conditions; curve B, best conditions; and curve
C, poorest conditions. Domed relationship --...
where cannibalism of adults on young, high
density of young, and the time lag in predator
response to prey density. From Radovich

(1962). Calif. Fish Game. 48:123-140. srUNINC STOCK SIZE ---...



Figure 4.11. Pacific sardine survival ratios .41 .31
(year-class divided by spawning stock size) plot- • 31 .32

ted as a function of spawning stock size shows .40

that above 1 billion spawners survival • 33

decreased. From Radovich (1962). Calif. Fish

e" .41 •! 41

00 1 2 3 4 5 • 1 10
Game. 48:123-140. SPAWNING STOCK SIZE (s) IILLioNS or rlsa

Food availability to larval fish is probably the single most important factor in survival,
with predation next, followed by adverse currents and temperature.
Numerical relationship between parent and progeny produces a relationship with ex-
tremely high variability. To improve the estimates, the next logical step is to collect data on
numbers of early larval stages of a fish species and numbers that subsequently enter the fish-
ery as recruits.
It is well known that early larval stages of marine organisms that hatch from highly fe-
cund females who provide no parental care die in extremely large numbers in their early lar-
val stages. Recent studies show that the precision of correlations of even early larval stages
of marine fish is too low to test whether abundance or mortality rates of early larvae are re-
ally correlated with recruitment. After larval fish have metamorphosed, the strength of rela-
tionship and precision of sample correlations increase sufficiently to permit more reliable
forecasting of recruitment for some fisheries. Australian studies have provided reliable re-
cruitment of spiny lobsters using postsettlement stages. (See Chapter 12).
78 Part One Living Resources

As a rule, assuming that the sampling data is reliable, the strength of recruitment esti-
mates improves the closer in time the fish are to the size and age of recruitment. Species
such as cod or herring where juveniles can be sampled for a year or two prior to the time
they are recruited into a fishery, can be expected to provide more accurate data. (See
Chapter 12.)
Many difficulties are associated with attempting to study parent-progeny relationships, es-
pecially on marine species. Chief among these are obtaining reliable estimates of the size of
the parent stock and the number of recruits over many years. These data are difficult and ex-
pensive to collect, especially for migratory open sea fishes. A variety of different environ-
mental factors (adequate food, predation pressure, adverse currents) can affect survival in
different years causing wide variation in the relationship, thereby obscuring a trend with dif-
ferent sizes of parent stocks. This is especially true in species such as salmons, Oncorhyncus
and Salmo, and warm water shrimps, Penaeus, that spend a portion of their lives in freshwater
where additional environmental factors may greatly influence their survival.
Using spawning ground surveys to estimate size of spawning population recruitment and
forecasting future harvest is illustrated in Figures 4.12 and 4.13.

~ 6

~ 5
::I 4


c 3

Figure 4.12. Index of production of pink

salmon for the years 1951-1955 in the Skeena
River and total October rainfall. Survival of
eggs from the spawners appears to be
________ ________ ________ __
0L-~ ~ ~ ~

influenced by discharge during early

20 30 40
incubation. From Wickett (1958). J. Fish.
NEW HAZELTON, PRINCE RUPERT a KITIMAT Res. Bd. Can. 15(5):1103-1126.
Chapter 4 Life Histories of Resource Species 79


Estimated total Estimated

number of eggs and survival to
early larvae market size

Estimated size Parent/progeny Estimated

of spawning relationship abundance of
population (Recruitment) progeny


Hypothetical data below can be used to illustrate the value of spawning ground surveys:
Average fecundity/female = 1,000 eggs
Sex ratio 50:50
Survival-egg to market size = 0.2 percent
Rate of harvest = 50 percent
Correlated Population. The total number of eggs and early larvae as estimated from spawning ground surveys
is 200,000:
Dividing 200,000 eggs and early larvae by the average fecundity per female gives an estimate of 200
female spawners. Knowing that the sex ratio is 1 to 1, add 200 males to the number of females, which
is the estimated number of spawners (population size at the time of spawning) to produce the 200,000
eggs and early larvae, e.g. 400 fish.
Parent/Progeny Relationship (Recruitment). Using the total number of eggs and early larvae as estimated from
the spawning ground surveys:
(200,000) and knowing that survival to market size is 0.2 percent, gives a figure of 400 progeny
(200,000 x 0.002 = 400 progeny). Determination of survival rates is discussed in Chapter 7.
Forecasting Future Harvest. From an estimated spawning population of 400 fish, 400 offspring are produced, as
shown above:
Of these offspring, 50 percent will be harvested by fishermen, 200 fish.

Figure 4.13. Determining abundance of spawners, parent/progeny relationship (recruitment), and

future harvest from spawning grounds surveys.


History of Fisheries Research

Benson, N. G. 1970. A century of fisheries in North America. Am. Fish. Soc. Spec. Pub. No.7. Wash-
ington, D.C.
Deacon, M. 1971. Scientists and the sea, 1650-1900: A study of marine science. Academic Press, Lon-
don and New York.
80 Part One Living Resources

Fishing Gazette. 1979. The American fisheries (1620-1979). Fishing Gazette 96(11):51-90.
Graham, M. 1954. Half a century of fishery biology in Europe. Proc. Gulf Carib. Fish. Inst. 6:70-90.
Gulland, J. A. (ed.). 1971. The fish resources of the ocean (section 5, pp. 246-255). Fishing News
(Books) Ltd., West Byfleet, England.
Herrington, W. e. 1954. 50 years of progress in solving fishery problems. Proc. Gulf Carib. Fish. Inst.
Johnstone, L. 1977. The aquatic explorers: A history of the Fisheries Research Board of Canada. Uni-
versity of Toronto Press, Toronto and Buffalo.
Roos, J. E 1991. Restoring Fraser River Salmon: A history of the International Pacific Salmon Fisheries
Commission 1937-1985. The Pacific Salmon Commission, Vancouver, Be.
Thomasson, E. M. (ed.). 1981. Study of the sea: The development of marine research under the aus-
pices of the International Council for the Exploration of the Sea. Fishing News (Books) Ltd., Farn-
ham, England.

Reproduction and Fecundity

Dadswell, M. J., R. J. Klauda, e. M. Moffitt, R. L. Saunders, R. A. Rulifson, and J. E. Cooper (eds.). 1987.
Common strategies of anadromous and catadromous fishes. Proceedings of an International Sym-
posium, Boston, MA, March 9-13,1986. American Fisheries Society Symposium 1.
Diana, J. S. 1994. Biology and ecology of fishes. Biological Sciences Press, Carmel, IN.
Forsberg, J. E. 1993. Estimating sex of Pacific halibut, Hippoglossus stenolepsis, using Fourier shape anal-
ysis of otoliths. International Pacific Halibut Commission, Seattle, Washington. Technical Report
No. 29. 23 pp.
Lagler, K. E, J. E. Bardach, and R. R. Miller. 1977. Ichthyology. John Wiley & Sons, Inc., New York.
Potts, G. W. and R. J. Wootton. 1984. Fish reproduction. Strategies and tactics. Academic Press, Or-
lando, FL.
Royce, W. E 1984. Introduction to the practice of fisheries science. Academic Press. New York.

Cushing, D. H. 1973. Recruitment and parent stock in fishes. University of Washington Sea Grant
Publ. WSG 73-1. 197 pp.
Jones, R. 1982. Population fluctuations and recruitment in marine populations. Philos. Trans. R. Soc.
Lond. Ser. B. 297, No. 1087.
Lasker, R. 1978. The relation between oceanographic conditions and larval anchovy food in the Cali-
fornia current: Identification of factors responsible for recruitment failure. Rapp. Proc.-Verb. Reun.
Int. Comm. Expl. Mer. 173:212-230.
Lasker, R. 1981. The role of a stable ocean in larval fish survival and subsequent recruitment. In Ma-
rine Fish Larvae-Morphology, Ecology and Relation to Fisheries. R. Lasker (ed.). Washington Sea
Grant Program. 79-87.
Pitcher, T. J. and P. J. B. Hart. 1982. Fisheries ecology. AVI Publishing Co., Westport, CT.
Sinclair, M. 1988. Marine populations: An essay on population regulation and speciation. University
of Washington Press, Seattle.
Skud, B. E. 1982. Dominance in fishes: The relation between environment and abundance. Science.
Smith, A. D. W. and e. J. Walters. 1981. Adaptive management of stock-recruitment systems. Can.
Fish. Aquat. Sci. 38(6):690-703.
Chapter 5

Age and Growth

of Resource Species

Estimating ages of fish and shellfish is necessary to determine aspects of their biology that
are fundamental to fisheries management. These include growth of individual fish and the
biomass of the species, age at maturity, and rates of mortality. Age studies form the basis of
our increased knowledge of fluctuations in the strength of year classes, and the abundance
that might be caused by predation, adverse weather conditions, and density of populations.
Variation of growth rates associated with latitude (temperature) require aging of fish before
meaningful relationships can be developed. Methods have been discovered and developed
to estimate ages of aquatic organisms, and the literature that has accumulated on the subject
during the last 90 years is voluminous.


The age estimation method in most widespread use at the present time is reading fish
scales (Fig. 5.1). It is based on interpretation of microscopic structures visible on fish scales.
Each scale is covered with roughly concentric lines or ridges, known as circuli, which are
irregularly distributed and show breaks at various intervals. These breaks are the year
marks or annuli by means of which age is determined, and most juvenile and adult fish
having these year marks may be easily observed with a hand lens or low-power micro-
Some general characteristics of annuli are that they are concentric with the margin of the
scale; they usually can be traced around the sculptured part of the scale; sometimes they
can be followed around the unsculptured part; they are separated from each other and do
not ordinarily meet at any point; and they are present on all normal scales of an individual
Scales are formed when newly hatched fish complete their larval stages, and cover the en-
tire body, with the exception of the head and fins. In most species, they lie in an overlapping
pattern much like shingles on a roof. Their growth begins with the formation of the scale
center, or focus, and growth is outward from this point, although it is greatest toward the
forward margin of the scale. Fine ridges called circuli are laid down in a circular pattern
around the focus as growth proceeds. Many circuli are added to the scale each year.

82 Part One Living Resources

""'-"---- 9
i31--- &



Figure 5.1. Annual marks on a ctenoid scale

from a 10-year-old Pacific Ocean perch,
Sebastodes alutus. From Alverson and
Westerheim (1961). International Commission
for the Northwest Atlantic Fisheries. Special
Publication No.3. pp 12-27.

The two types of scales found on most bony fishes are: cycloid scales that are subcircular,
disclike, thin, and are to be found in most soft-rayed fishes; and ctenoid scales, that are sim-
ilar to cycloid scales but possess cteneii (small teeth) on the exposed surface. These are char-
acteristically found on most spiny-rayed, bony fishes.
The scale method of age estimation rests upon the following conditions:

1. After formation, the number of scales on the body of a fish remains constant throughout life.
2. The same scales are retained throughout life, unless lost accidentally, whereupon a new scale
forms. New or regenerated scales do not exhibit a record of growth from time the scale was
lost, but instead show a mottled, opaque area in the center. These regenerated scales cannot be
used for age determination.
3. The growth of the scale and the growth of the body of the fish are roughly proportional.
4. The rate of growth of a fish, except in tropical and subtropical latitudes, varies throughout the
year. Growth is most rapid during the warm summer months and practically ceases during the
winter. This seasonal variation in rate of growth is recorded in the spacing of the circuli on the
scale giving rise to the bands of growth that represent each year of life (Fig. 5.2).

Scale reading is a very complex procedure that requires considerable interpretation. A

long training period with an experienced scale reader is required for the beginning scale
reader to make the most accurate age estimate. It is by no means mechanical, and is rarely
simple. Many stumbling blocks arise because of the presence of accessory checks, spawning
marks, or false year marks, as well as occasional abnormalities in the configuration of the
scale make precise age determination difficult, and often impossible.


Five major categories should be used when estimating age from scales or "hard parts" (other
body structures that are used for age estimation). These are interpretation, validation, col-
Chapter 5 Age and Growth of Resource Species 83

Figure 5.2. Blownup section of a red salmon

scale showing slow freshwater growth and
more rapid growth in the sea. Photo courtsey
of U.S. Fish and Wildlife Service.

laboration, automation, and innovation. These categories are briefly described below. For a
full description see paper by J. Casselman in Prince and Pulos (1983).
Interpretation means the hard parts are readable (being able to see or read "annuli" on the
scale), and that, it must be determined that the marks observed are laid down on the scale in
a regular fashion, either a yearly or lesser period. Descriptions of methods used to read cal-
cified tissue should be precise. To check the readability of the number and location of annuli,
a reader should read a sample of scales, place them aside, and later reread them without ref-
erence to the results of the first reading. Agreement of the number of annuli on each scale
should be good and the location of each annuli should be the same in both readings. Dis-
tance from the focus of the scale out to each ring can be measured on a viewing screen. Dis-
crepancies between readings may result due to false marks (mentioned earlier), difficulty in
locating the first annuli, and crowding of year zones in later years on long-lived fish because
of slow growth.
Validation means establishing that marks read on scales or other hard parts are indeed
valid indicators of age, and cannot be overemphasized. If marks read are not valid indicators
of age, but are only assumed to be, growth data based on these marks will be suspect or
Recent studies have shown that daily marks are laid down on the otoliths of very young
(larval) fish that are very useful in determining the length of larval life and early growth of
fishes. In this discussion here, however, we use annual marks on juvenile and adult fish for
determining the validity of the method. A correlation between the number of marks and the
84 Part One Living Resources

size of the fish scale and/or fish length provides evidence of validity of age marks because
the larger, older fish would be expected to have more marks than small, young fish. The
number of marks on a scale can be compared with a graph showing the distribution of sizes
of fish (called length frequency distribution discussed later). The number of modal groups
that represent year classes in the fishery should agree with the number of marks on the scale.
Measuring the distance between the last annulus on the scale and the scale margin at various
times of the year will show if the annulus is formed only once yearly and during a certain
season (approximate birth date of the fish). As the scale grows, the marginal increment will
increase until a new annulus is laid down, at which point the increment will drop to zero.
Tagging wild fish and taking scales at the time of tagging can provide evidence of the valid-
ity of marks when the fish are recaptured and the scales are again taken and compared with
those taken earlier (the scale of a tagged fish at liberty for 2 years would be expected to have
two additional marks on its scale).
Another technique to validate that marks on hard parts are indeed age marks is to hold
fish in tanks, ponds, or aquaria and check the number of marks in relation to time held in
captivity. A disadvantage is that in this technique, environmental conditions in the enclo-
sures might be radically different from those in the wild and thereby cause false rings to ap-
Collaboration means that more than one reader (interpretation) or several experienced
readers is preferable to, read scales and compare their interpretations. This technique is not
really proof but rather collaboration and it, is to look for agreement between the number of
rings of more than one hard part, say between scales and otoliths.
Automation can be used such as image analysis. This procedure removes the possibility
of personal bias and reads all scales objectively.
Innovation means finding new techniques to improve the accuracy of aging fish hard
parts. Most of this research has been done with otolith microstructure and biochemical

Some fishes do not have scales or have epidermal (ganoid or placoid) scales that possess
the structural features of dermal (cycloid and ctenoid) scales employed in age and growth
determinations. For such fishes the age may be determined by examining the otoliths or "ear
stones" correspond to a part of the inner ear of man and other mammals. All bony fishes
have otoliths, although their appearance ranges from small sandlike grains to large ringed
pearls. The exact shape and size of otoliths are usually characteristic in a species. Fishes
have three pairs of otoliths, although only one pair is usually large enough to be used for age
estimation. They consist of calcium carbonate in an organic matrix and are associated with
orientation, balance, and hearing in fishes.
Archaeological researchers found that fish otoliths, primarily sagittae, were known as
early as 300 BC and believed to be used as good luck charms, ornaments, and for the aphro-
disiac properties they were believed to possess (Fig. 5.3). For many years paleontologists
identified otoliths from fossil deposits and assemblages. Not until 1898 did fishery workers
realize the potential for studying the age and growth of fishes using otoliths. Cod otoliths
were used even then to determine age at first spawning. Subsequently, herring otoliths were
used in racial studies. Since then otoliths have received wide attention by fisheries scientists.
The otolith has ridges arranged in a somewhat concentric fashion that may be interpreted
in the same manner as the circuli and the year marks or annuli of fish scales.
Chapter 5 Age and Growth of Resource Species 85

Figure 5.3. Drawing of a hypothetical red

grouper otolith with 10 annual marks. From
Moe (1969). Florida Department of Natural
Resources. Professional Papers Series Number
Lateral view Dorsal view
10. 95 pp.

Aging of larval finfish, unheard of in the past, has been successfully used on many species.
Two examples give some details on the technique. In a study of California halibut, Paral-
ichthlys californicus, laboratory-raised juveniles having sagitta of known age were mounted in
resin on a microscope slide. To obtain a relationship between the number of increments on
the sagitta with the known age in days, a microcomputer interfaced to an electronic digitizer
was used to measure and count increments on a projected image of the otolith from a high-
resolution videocamera on a compound microscope. Analyses showed that increments were
formed daily.
In another study, on the larvae of Dover sole, Microstomus pacificus, otoliths were embed-
ded in casting resin and polished to the core. Next they were etched with 5 percent buffered
EDTA (ethylenediaminetetraacetic acid). A scanning electron microscope (SEM ) showed
that the growth rate was about 0.2 mm/day.
Recent developments in age and growth studies include improvements in a computerized
system using a videocamera to scan zones in skeletal structures of fish, biochemical methods
(RNA-DNA ratios and amino acid uptake by scales) for determination of short-term fish
growth, and the use of otolith microstructure to study daily growth of larval fish. Otolith
microstructure has attracted the attention of many fishery biologists who have mainly fo-
cused on daily growth increments to determine hatching dates and the influences of environ-
ment variables on growth and survival of larval fish.
The ratio between RNA (ribonucleic acid) and DNA (deoxyribonucleic acid) provided a
short-term index of growth in a variety of organisms in the 1960s and 1970s. RNA is respon-
sible for protein synthesis and DNA is the primary carrier of genetic information; the ratio
between these nucleic acids correlates with known growth rates of larvae in laboratory exper-
iments. The method works well on larval fish because of the characteristic continuous
growth during that life stage. Experiments in the mid-1980s confirmed the value of this
method on larval fish that promises better understanding of larval growth and mortality, and
their relation to environmental parameters.
86 Part One Living Resources

Other Hard Parts

Other hard parts (calcified structures) of fish such as some of the bones also exhibit char-
acteristics that may be used for age estimation. Cross sections of vertebrae (catfish that are
scaleless, and albacore) and fin rays (sturgeons and some catfish) exhibit concentric markings
that have been found to correspond to the annular markings of scales and otoliths. Even
opercula have been used in some studies of aging in fishes. However, because of the dis-
tinctness of the structure of scales in comparison with other bony parts of fishes and the ease
of collection, scales usually give the most satisfactory and reliable result, if they can be em-
ployed in age estimation.
Calcified structures to determine age estimates have been widely used over a 50 year
period. What has been described as a technological revolution in aging fish, began in the
1980s. New techniques are being developed that include biochemical methods of measuring
instantaneous scale growth to provide a direct measure of growth rate, and radiometric age
determination that provides a more objective age assessment. These developments now pos-
sible with new powerful and sophisticated tools and techniques promise to provide badly
needed objective aging of fish.


An important question that bears on using age marks is what causes them to be laid down
on hard parts. Unfortunately, as in many biological phenomena, no single reason appears to
apply to a range of species. There are theories concerning changes in growth, metabolism,
and spawning periods. For example, high temperature increases the growth rate in summer
and may cause wide spacing in the circuli; conversely, cold temperatures cause slow growth
and the appearance of annual winter marks. Weaknesses in the temperature theory are
found in experiments where fish are held in warm tanks yearround yet produced age marks.
Some warm water species in the tropics also lay down annual marks. Temperature as the
sole factor causing age marks is thought by some scientists to be overemphasized. It may be
the greater availability of food in combination with high summer temperatures, or physiolog-
ical rhythms related to the onset of spawning in some species. Long summer photoperiods
when temperatures are high and food is usually more available have also been suggested as
the cause. Whatever environmental parameters or physiological factors of the species cause
age to be recorded, fishery biologists can be confident in the knowledge that validated age
marks that appear on many species of fish and shellfish are a good determinant of age. The
development of this method has been a significant contribution to management of fish stocks.
It should be noted that crustaceans lack hard parts that can be used for aging, so other
techniques have to be used, such as tagging or size frequency analyses.


The oldest method of fish age estimation is known as the Petersen method, or the length-fre-
quency distribution method. It is fairly simple, but is useful only in dealing with large pop-
ulations or random samples of fishes as units. The method tabulates the length
measurements of fish on a frequency basis and interprets the modes in the distribution as the
age groups of the population, or sample, used. It is unreliable for the obvious reasons that it
Chapter 5 Age and Growth of Resource Species 87

cannot take into account unusual conditions of growth, nor does it reveal reliable information
concerning the growth history of an individual fish. The method is mostly used only for
fishes where another current method of age determination may be applied, or to gain a
rough preliminary estimate of age composition, or age segregation of a sample of a popula-
Length-frequency distribution is based on the premise that fish spawned over a relatively
short (contracted) time will all grow at about the same rate throughout their lives, following
a particular growth pattern. These groups of fish can be identified in a size-frequency distri-
bution and when plotted with time the modal groups will progress suggesting growth.
However, because some fish of a certain year class are spawned earlier than others and some
later, and conditions for growth vary between fish, modal groups will have a symmetrical or
normal curve appearance in size-frequency distributions.
One of the difficulties of this method is identification of the modal groups in the total dis-
tribution, also called dissection of multimodal distributions (Fig. 5.4). The complex multimo-
dal curves representing in some cases many age groups of fish are formed by a combination
of a number of "normal" curves. Some researchers simply draw freehand normal curves on
multimodal distributions for a gross estimation of the number of age groups present and
their average size at each age. The appearance of the same modal groups in about the same
position, allowing for growth, on repetitive samples gives some confidence in the method.
Another method for locating modal groups requires considerable mathematical manipula-
tion, but provides numerical values for modal position and standard deviation of the modes.
Length distributions are converted to percentages, accumulated, then plotted on probability
paper. Points of inflection indicate valleys between the modes.
Modal groups can be identified by the use of standard normal curves. Different normal
curves constructed with different size distributions and areas are placed over histograms of
fish distributions as illuminated below. The standard normal curves that best fit the histo-
grams selected by eye are sketched. The differences between the theoretical curves and the
observed curves are tabulated and subjected to a chi-square statistical test. The lower the
chi-square value, the better the fit between the theoretical and observed data; a perfect nor-
mal curve would show zero chi-square value.
A computer program for dissecting polymodal distributions called ELEFAN (Electronic
Length Frequency Analysis) combines Petersen and modal progression analysis.
As a general rule, if the modal groups are clearcut on the basis of size frequency distribu-
tions, methods of dissecting the modal groups out will prove their identification, but if
modes cannot be identified in the distributions even considerable manipulation of data may
not give accurate age readings.

Limitation of the Size-Frequency Method

1. Length-frequency distributions do not always approximate normality, e.g. because of prolonged

spawning periods, or size selection by fishing gear.
2. Two or more modes from one age group may occur because of prolonged spawning or inter-
mixture of two groups of different sized, same age individuals.
3. If age of first capture is unknown, what is the age of fish in the first modal group? New tech-
niques for reading larval and juvenile otoliths can provide this data for some species.
4. Growth increments may be too small for distinct modes to appear, particularly at older ages, es-
pecially in long-lived fish.
5. Population technique requires big samples. The method cannot determine age of individual fish.
88 Part One Living Resources


Figure 5.4. Monthly weight frequency

O~~TT~~~~~~,,~~~~~~~~,~.~~ distributions of bigeye tuna from the Honolulu
market for 1950. From Iversen (1955). U.S.
Fish and Wildlife Service, Spec. Sci. Report,
WEIGHT IN POUNDS Fisheries No. 162, 40 pp.

6. Fish of a size (not necessarily of an age) tend to school together.

7. One or more age groups may be missing or poorly reported.
8. For best results, a unit stock or closed population is needed. Migration of fish into or out of the
stock can confound interpretation of size-frequency distributions.

In summary, aging of fish and shellfish can be accomplished by several different tech-
niques, all of which are estimates. Whatever method is employed, it is important that it be
critically evaluated by an able investigator who can identify age marks or modal groups of
fishes. The investigator must provide evidence that he is using valid indicators of age. The
Chapter 5 Age and Growth of Resource Species 89

extreme variation that aquatic animals demonstrate due to behavior aspects and environment
parameters makes age determination a skill that takes time and patience to master.


Growth rings appear as concentric striae on the

outer surface of the shell of some mollusks. The
giant scallop, Placopecten magellanicus, shows ring
formation only once a year (Fig. 5.5). Shells of the
ocean quahog, Arctica islandica, indicate the pres-
ence of growth "rings" or ''bands.'' Over 500 recov-
ered marked clams released off Long Island, New
York, provided evidence of annual periodicity of
growth line deposition. Many were over 100 years
old, with one specimen estimated to be about 225
years old. The ocean quahog, which reaches about
5.2 in (130 mm) in length, is perhaps the longest
living bivalve species known. Ages averaging
about 100 years have been estimated for both
freshwater mussels and marine mussels from the
Far East.
Marking of bivalve mollusks consists of inscrib-
ing identifying numbers on their valves, using a
dentist's drill. The number of growth rings are
counted at the time of marking, then they are re-
leased. When subsequently recovered, the time at Figure 5.5. Drawing showing age rings
liberty can be checked with the number of addi- on the shell of a sea scallop, Placopecten
tional rings put down. magellanicus. From MacKenzie (1979).
Aging estimates can even be done on some spe- NMFS Tech. Ser. No. 19.
cies of mollusks having no shells; for example,
daily growth increments can be determined using squid statoliths (that particle of sand or
calcium carbonate within the hollow cell that acts as the squid's organ of balance). To check
the validity of these increments, squid are fed shrimp soaked in strontium chloride solution.
This chemical, when deposited in the statolith of the squid, appears in sequential bands that
correlate with time elapsed since marking.
Recent studies have shown that otoliths even show a record of daily larval growth in some
fish species. Important management decisions are made on the basis of fish and shellfish
ages such as those of size at first spawning, numbers of age groups in exploited stocks, age
to market size, age-mortality relationships, etc. Validating age marks is very important as
noted earlier in this chapter.


Growth rates of fish and shellfish are basic parameters for any yield equation that provides
estimates of the weight of the exploitable stock. When a single fish is taken from a popula-
tion an important question is "how long will it take to replace this fish?" If the target species
90 Part One Living Resources

is a penaeid shrimp that reaches market size and sexual maturity and spawns at an age of
less than 1 year, the management strategy for that fishery is quite different than that for the
Arctic clam, for example, that may take about 100 years to replace.
When finfish or mollusks are aged as described, graphs and equations describing their
growth can be prepared. Examination of growth curves shows periods of rapid and slow
growth during the life of the fish that suggest the best time to catch fish in a capture fish-
ery, or when to harvest them in a culture fishery (fish farming operation). If we are to
manage stocks properly there is no question that reliable growth rate estimates must be

Types of Growth
The following discussion concerns finfish that exhibit continuous growth because this type
of growth occurs most widely in finfishes. Students will encounter in their readings other
kinds of growth that have to do more with understanding how certain animals grow.
Crustaceans show a stepwise growth rather than continuous growth, as seen in finfish
(Fig. 5.6). In stepwise growth, increases are made during a short period of time during the
molt. In effect, crustaceans grow little during the intermolt period.





". (x) Male
() Female
(-) Cruschers Figure 5.6. Example of stepwise growth
(--) Pincers
typical of crustaceans as shown by stone crabs.
From Ehrhardt and Restrepo (1989). In Marine
invertebrate fisheries. Their assessment and
0 4 management. J. F. Caddy. (ed.). 225-240. John
Age (years) Wiley & Sons, Inc., New York.

Endogenous growth simply means that the growth rate is controlled internally, as is the
ultimate size of the animal. Even with abundant food and living in an ideal environment, a
fish will grow only to an approximate length that is characteristic of its species. This fish will
probably be somewhat larger and fatter than its cohorts who have not had the benefit of
abundant food and ideal environment, but they will not be three or four times larger than
their cohorts from the benefits the environment provides. This increase is called exogenous
growth. In other words, growth is greater for this animal because of abundant food and
ideal living conditions and is controlled by exterior conditions. It exerts a relatively small ef-
Chapter 5 Age and Growth of Resource Species 91

feet on the size of the animal in comparison with endogenous growth. Other terms used for
growth patterns of more interest to a taxonomist than a fishery biologist, are:
Isometric (isogonic ) growth literally means equality of measurements. In growth termi-
nology, this means that all body parts essentially grow proportionally.
Allometric (heterogonic) growth describes changes in the relative proportion of the size of
one part of an organism in comparison with the growth of the whole organism. Proportion-
ally, changes may be temporary or short term; for example, ovary size of shrimp during the
spawning season; or long term, such as length of the rostrum of a penaeid shrimp in relation
to the body length, for example, the tendency for the rostrum to be proportionally shorter on
older, larger shrimp than on young, small shrimp.
Growth, then, for individual fish is similar to a fish within a population. It is the sum of
the increases over the losses. The general pattern of this growth is called "sigmoid": A plot
forms a sigmoid curve (from the Greek letter sigma) when weight of fish is plotted against
time. The growth rate of an animal is slow during its embryonic larval stages, then increases
rapidly in its juvenile life. When growth starts to slow down the change is called the point
of inflection. The animal continues to increase in weight but at a decreasing rate. At both
extremes, then, the very early life and the late life of the fish, increases in weight are very
slow and approximate asymptotes.
Growth of fishes can be described in different ways (Fig. 5.7; Ricker 1975):

1. Absolute growth. Using lengths of fish the increase over 1 year can be described for lengths
L2 - L} where L} = length at time 1 and L2 = length at time 2. Weights can be substituted for
lengths in this simple formula.
2. Relative growth. This increase of fish in either weight (W) or length (L) is here expressed as a
proportion increase for the year. Therefore:

relative growth = L2 - L} / L} .
92 Part One Living Resources

Using the same notations as above. Again weight can be substituted for length.
Instantaneous rates of increase are also called logarithmic, exponential, or compound-in-
terest rates. Instantaneous rates of increase of weight or length can be obtained using the ap-
propriate units in this equation

One problem scientists have struggled with over the years is that of devising mathematical
models or means of describing the growth of all or almost all fishes. An equation published
in 1934 by Von Bertalanffy describes the growth above the point of inflection and is based on
physiological processes responsible for the mass of an organism. It involves catabolism
(breakdown) and anabolism (synthesis). This equation has been rather widely accepted
since it was "discovered" some years after publication. However, many published accounts
urge caution when using this equation as a generally applicable growth law.
Von Bertalanffy's coefficient of catabolism is proportional to the role of body destruction
per unit of weight and time. The greater part of catabolism involves protein breakdown. He
used the rate of nitrogen excretion of starving animals as an estimate of catabolic rate and ob-
tained experimental values of the breakdown coefficient K that agreed well with those ob-
tained by fitting curves calculated from his growth rate applied to that animal.

where 100 = length at infinity, It = length at time t, e = base natural logs = 2.71869, k = logk
slope, to = time zero.
In practice, k is determined by using log to the base e of the slope of the Walford line.
(This technique will be discussed in the next section.) The fact that Von Bertalanffy developed
this equation from bioenergetic principles gained it wide acceptance. It is continuously ac-
cepted because proponents say it is flexible, will fit a wide range of data, and is easy to use.
Some also say that despite its disadvantages it is so imbedded in the literature that we
should continue to use it primarily for the sake of comparison of early growth rates of fish
species with more recent rates.
That the Von Bertalanffy curve only describes growth above the point of inflection does
not substantially reduce its utility to the fishery scientist because this change in growth rate
at the point of inflection usually occurs at an early age of a fish, and, as a rule, before recruit-
ment into a fishery. Therefore, the important part in the growth curve of greatest interest to
the fishery biologist is that past the point of inflection.
Another technique, called the Walford growth transformation, can be used to describe or
illustrate growth above the point of inflection that involves a transformation that produces a
straight line relationship (Fig. 5.8). This is in contrast to the Von Bertalanffy equation that de-
scribes a curvilinear relationship. If the length of an animal above the point of inflection is
measured, then again at a later time, say 1 year, a series of these points will fall on a straight

where k = slope of line and 1 = length at age n.

On this graph with the abscissa length at age x, and the ordinate length at age x + I, a 45°
line can be drawn that is, in effect, a "no growth line." In other words, at year 3 the fish is a
Chapter 5 Age and Growth of Resource Species 93

Age (years)


y ~ .7021 X + 250.236

Figure 5.8. Walford line of observed standard

length in male king mackerel. From
Beaumarriage (1973). Florida Dept. Nat. Res. 1100

Res. Lab. Contribution No. 226. 45 pp. Standard Length (mm) at Age N

certain length, and if measured 1 year later at year 4, the point will fall on the 45° no growth
line. The points on a curve at this time and as the fish or sample of fish gets longer or in-
creases in growth, will diminish and the line drawn through the points will be a straight line.
Therefore, by using the Walford growth transformation, the slope of this decrease in growth
rate estimates of the rate of growth above the point of inflection, and the point where the
growth line crosses the no growth line estimates the ultimate length for the species of fish
being studied.
For both the Von Bertalanffy and the Walford growth transformation estimate, the age of
fishes must have been determined by one or more of the methods described earlier, such as
reading hard parts: scales, otoliths, finrays, or vertebrae. The Walford technique also requires
that the age, or some measure of the passage of time, be available. The Walford curve is ad-
vantageous in that it can be used to describe growth in a group of aquatic animals that can-
not be aged using hard parts, that is, crustaceans. By using tags and recording length for
short periods, 10 days or 30 days, for example, it is possible to estimate growth. This esti-
mate, of course, requires that the animal be growing rapidly enough to be able to indicate
growth for such short periods. The Walford technique has been used on fast-growing short-
lived penaeid shrimp. In some cases, analyses of length frequency distributions can also be
used to estimate growth of crustaceans. Age estimation of fish is not necessary to study their
growth by length frequencies if prominent modes can be identified that progresss in an or-
derly fashion with time.
94 Part One Living Resources


As stated earlier, because there is an approxi-

mate relationship between scale length and the
size of fish scales, the lengths at an earlier age of
old fish whose scales have been read can be es-
timated (Fig. 5.9). This being true, past growth
history of a fish can often be worked out from
its scale through a technique called ''back calcu-
lation." (Table 5.1). With this technique, if the
length of a fish at capture is known, it is possi-
ble to calculate lengths at earlier ages from
measurement of the scale at each year mark.
The increase in length of a sardine in each year,
in relation to increase in scale length is illus-
trated in Fig. 5.9. Length at each of the three
year marks were determined from back calcula-
Back calculation is of importance in fisheries
studies because it permits an evaluation of
growth rate of fish in all years of life from a
smaller sample than otherwise would be possi-
ble. However, many years ago Einar Lee found
that calculated lengths of fish at a particular age
become progressively smaller when calculated
successfully from older, larger fish. (Fig. 10).
Lee's phenomenon, seems to be due in part, at
least, because simple proportionality formulas
assume that the line describing the relationship
o between body length of a fish and a scale diam-
eter passes through 0, 0 as in 0 length fish and 0
size scale. Studies of fish scale formations show
Figure 5.9. The growth of fish and the in-
dividual scales occur at about the same rate. that scales may not appear until the fish reaches
If it can be demonstrated that a ring is a particular size, e.g., when salmon become 1
added to each scale each year it is possible 1/2 to 2 in (38 to 51 mm) long, scales start to de-
to calculate the size of a sardine at a former velop. Another possible reason is that the scales
time interval of its life. From California Co- exhibit allometric growth.
operative Sardine Research Program (1952). To calculate the body-scale regression for cal-
Progress Report January 1, 1951 to June 30, culated lengths,

where Sc is the scale measurement to the edge of the scale, Si is the scale measurement to
each annulus, L is the length of the fish at capture, and a is the intercept of the body-scale re-
gression as previously determined from an adequate sample.
It is reasonable to assume that warm waters stimulate more rapid growth in fishes than do
temperate waters. There are species exceptions to this notion, e.g. soles and scrombrids. An
Table 5.1 Example of Back Calculating of Lengths of Female King Mackerel From Annual Marks on Otoliths

Mean Back-calculated Lengths at Successive Annuli
No. of Length
Speci- at
Age mens Capture 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

1 16 548 462
2 24 719 503 668
3 19 820 519 681 775
4 43 832 494 635 727 794
5 51 878 500 650 739 804 856
6 37 897 484 634 716 775 829 876
7 30 937 487 632 718 778 831 876 919
8 14 977 491 646 731 790 836 878 917 956
9 39 990 477 616 709 768 815 857 896 937 975
10 42 1,005 477 629 699 759 804 847 883 919 956 991
11 32 1,052 482 630 710 765 812 855 893 929 964 998 1,034
12 27 1,063 466 620 701 755 801 840 877 912 946 977 1,013 1,044
13 17 1,035 482 610 674 724 767 801 837 870 902 931 961 990 1,022
14 12 1,041 458 599 663 713 753 791 824 856 885 921 948 976 1,007 1,035
15 9 1,145 500 643 718 772 818 858 899 931 962 993 1,022 1,050 1,077 1,105 1,134
16 3 1,189 513 668 753 807 847 883 919 954 981 1,008 1,043 1,070 1,097 1,123 1,154 1,181
17 2 1,216 504 629 708 766 811 850 889 928 960 992 1,018 1,056 1,088 1,120 1,152 1,177 1,203
18 2 1,272 480 642 737 785 834 871 920 950 981 1,018 1,049 1,074 1,105 1,130 1,159 1,191 1,215 1,246
19 2 1,075 416 551 626 663 704 740 766 791 812 843 869 889 916 936 962 983 1,003 1,034 1,054
20 2 1,151 464 603 671 723 764 795 825 856 881 906 937 957 987 1,007 1,032 1,052 1,077 1,106 1,126 1,146
21 1 1,220 421 602 686 727 768 796 823 865 892 920 947 974 988 1,015 1,043 1,070 1,084 1,111 1,138 1,165 1,192
Total number 424 408 384 365 322 271 234 204 190 151 109 77 50 33 21 12 9 7 5 3 1
Weighted mean 486 635 716 772 817 852 886 917 949 975 1,002 1,017 1,032 1,064 1,110 1,118 1,120 1,126 1,100 1,152 1,192
Growth increment 486 148 81 56 45 34 34 31 31 26 26 15 15 32 46 8 2 6 -26 52 40
Note that the estimated fork length of a young mackerel at a specific age usually decreases as the age at capture increases. Read the mean back-calculated columns
vertically with increasing age gives evidence of Rosa Lee's phenomenon. Sample sizes are very small in fish above 16 years of age so mean back-calculated lengths
are unreliable. From Collins, Schmidt, Waltz, and Pickney (1988). Fish. Bull. U.S. 87:49-6l.

96 Part One Living Resources

Assumed Relationship

Actual Relationship

Figure 5.10. Hypothetical graph showing one

A 1e_ltlll
lacernct Correct possible reason for the occurrence of Lee's
F1shLength phenomenon. Graph by E. S. Iversen, Jr.

Australian study shows growth in mean weight with age comparing species from temperate
and tropical waters. Fish were chosen with similar trophic positions and ecological habits.
Bottom-dwelling temperate species generally grew faster than tropical counterparts. A
number of possible reasons for this greater growth of tropical fish might be because of higher
metabolic costs in the tropics. Off-the-bottom species (mesopelagic and pelagic) showed sim-
ilar growth in temperate and tropical climes.

Length-Weight Relationship
Weight is the best way of depicting growth in fish because the fishery scientist is most in-
terested in biomass of a population, but weighing fishes accurately in the field, likely aboard
vessels under adverse weather conditions, is a troublesome task (Fig. 5.11). However, usually
a reasonably close relationship between length and weight exists. Hence using the more eas-
ily obtained data on lengths allows a transformation to be made from average length to av-
erage weight. The approximation shows that a species of fish of a certain length will weigh
a certain amount. Also, its weight will increase to the third power of its length. The formula
used is:

The W stands for weight, L for length, k for a constant. Plotting the length on the abscissa
and weight on the ordinate, it will be seen that the curve climbs rapidly upward (convex
curve or curvelinear relationship).
Not all fishes grow according to this equation. The shape in the example indicates that
this is an equation for ideal shaped fish such as tuna. A needlefish or a large swordfish or
marlin will not grow in the same way. Thus, for many species of fish it must be determined
first what the exponent is by taking large numbers of fish, measuring their length and
weight, then determining the value of the exponent. Hence, the general equation is:

W = kLn or logw = logk + nlogl

Chapter 5 Age and Growth of Resource Species 97

Spotfin Croaker Weight-Length Relationship

n-291 g'
W. 0.000032095 L2.94436







:!: 1000


~ 800




100 150 200 250 300 350 400 4110 500 550

Figure 5.11. Weight-length curve of 291 male and female spotfin croakers, Roncador stearnsi. From
Joseph (1962). Calif. Dept. Fish Game. Fish Bull 119.

To avoid a curvelinear relationship, logarithms can be used that will cause these data to
form a straight line on a log-log plot. That is to say, both the weight and the length are con-
verted to logs and plotted on arithmetic paper to show a straight line. If log paper is used,
then the numbers can be plotted directly and will form a straight line.

Condition Factor
The last matter of importance about growth has to do with a condition factor (or condition
index), sometimes called an index of well-being. The condition factor simply is a standard to
enable fishery biologists to compare the condition of fish from different bodies of water or
under different environments. The equation
98 Part One Living Resources

has the K equal to the coefficient of condition. The number obtained from this equation is, of
course, affected by how robust or heavy the fish is. When there is a large weight or W value
in the numerator, the condition value will be higher than when the fish is very thin, where
the W probably will be a smaller number, and hence the K value also will be smaller. The
condition factor is merely a standard means or index of determining the condition or well-be-
ing of fish.
Length-weight relationships and condition factors have been used extensively on freshwa-
ter fishes for years. Freshwater perch and bass living in isolated lakes in United States have
been the objective of many studies to rate the productivity of the lakes using growth in
weight or condition of the fish. Once done, it was possible to examine the environmental
and/ or biological conditions in the lakes to determine which were high and which were low.

Growth Measurements
In describing the growth of fish and shellfish, care must be taken in selecting the most
appropriate measurements (fork length, total length, etc.) for the species under study. The
measurement selected must be described fully in any publication reporting the results of
the growth study, so that other investigators can compare their measurements on the same
species from other geographic areas, or those made during other seasons, whatever the
Not only must the measurement technique be defined and standardized, but the condition
of the fish or shellfish measured must be reported. It is known that lengths (shrinkage) be-
tween live fish and the same fish after they have been dead for a time are reduced. Shrinkage
also occurs when fresh fish are dried, smoked, iced, or frozen. When sampling from a fishery
where landings may include live, dead, or preserved fish, the investigator has a responsibility
to determine if changes in fish length may have resulted from the condition of the fish he is
measuring. These factors are critical in his study. If these conditions exist, he should record
fish condition for all measurements and obtain a conversion figure, if necessary.
An important rule to follow when beginning a growth study, is to measure lengths or
weights to a greater precision then you would expect to need. If, upon analyses of your
data, you find that, as you suspected, your precision is greater than required, it is easy to
pool measurements into larger categories. On the other hand, if you find that small differ-
ences in measurements are important to your study, for example, measuring to the nearest
whole milimeters and you measured only the nearest whole centimeters, additional costly
field data may have to be collected to accurately describe the growth of the species you are
studying. These data, of course, can never replace data that may have been lost for the
original samples and may not permit critical comparisons, and/or may extend the time of
your study.
The use of weight frequency distributions is shown for Hawaiian yellowfin (Fig. 5.12).
When drawing a growth curve from size frequencies, the time between measurements may
be (as in this case) on a monthly basis. The points that locate the center of the modal distri-
bution for any age group are marked on the abscissa for each month. Then all points are
connected by continuous lines, as is shown in this figure. That curve, in effect, estimates the
growth of the stock during the 1 year period (Fig. 5.13).
Chapter 5 Age and Growth of Resource Species 99


10 .. ·274

10 .. ·5040

10 '''10lI0

u .uc
~ 10 .. ·3036
C 0 AT

10 "'34'3


10 ... 1603

10 11·700

10 ... 4111


Figure 5.12. Monthly weight-frequency distributions, as percentages, of Hawaiian yellowfin tuna

grouped for the years 1948-1953. From Iversen (1956). U.s. Fish and Wildlife Service, Special Sci.
Report, Fisheries No. 184. 33 pp.
100 Part One Living Resources

.3 4 56? 8 9 10 11 12 ,~ 14 15 16 17 18 19 20 21aar.
1907 20

1909 20

1910 20



1914 40




12 13 14 15 16 17 18 19 20 21 oar

Figure 5.13. Progression of a prominent year class passing through the Norwegian herring fishery,
1907-1919 illustrates the flucutations in recruitment from year to year and that the rate of growth of a
year class can be calculated using the size frequency method (measuring lengths annually of the
prominent year classes). From Hjort (1926). J. Cons. Int Explor. Mer. 1(1):5-38.
Chapter 5 Age and Growth of Resource Species 101


Age Estimation of Fish and Shellfish

Everhart, W. H. and W. D. Youngs. 1982. Principles of fishery science. (second edition). Cornell Uni-
versity Press, Ithaca, NY.
Hurley, G. V., P. H. Odense, R. K. O'Dor and E. G. Dawe. 1985. Strontium labelling for verifying daily
growth increments in the satolith of the short-finned squid,Illex illecebrosus. Can. J. Fish. Aquat. Sci.
Jearld, A, Jr. 1983. Age determentation. In Fisheries techniques. L. A Nielsen and D. L. Johnson.
(eds.) 301-324. Am. Fish. Soc. Southern Printing Co., Blacksburg, VA
Lux. F. E. 1971. Age determination of fishes (Rev.). NOAA/NMFS Fishery Leaflet No. 637. 7 pp.
Prince, E. D. and L. M. Pulos (eds.). 1983. Proceedings of the international workshop on age determi-
nation of oceanic pelagic fishes: Tunas, billfishes, and sharks. NOAA Technical Report NMFS 8.
211 pp.
Rounsefell, G. A 1975. Ecology, utilization and management of marine fisheries. C. V. Mosby Co., St
Louis, MO.
Royce, W. F. 1984. Introduction to the practice of fisheries science. Academic Press, New York.

Growth Determination (Estimation)

Burnham, K. P., D. R. Anderson, G. C. White, C. Rrownie, and K. H. Pollock. 1987. Design and analysis
methods for fish survival experiments based on release-recapture. Am. Fish. Soc. Monograph No.
5. Bethesda, MD.
Buckley, L. J. 1979. Relationship between RNA-DNA ratio, prey density and growth rate in Atlantic
cod, Gadus morhua, larvae. J. Fish. Res. Bd. Can. 36:1497-1502.
Buckley, L. J. 1984. RNA-DNA ratio: An index of larval fish growth in the sea. Mar. BioI. 80:291-298.
Edwards, R. R. C. 1984. Comparisons of growth in weight of temperate and tropical marine fish coun-
terparts. Can. J. Fish. Aquat. Sci. 41:1381-1384.
Everhart, W. H. and W. D. Youngs. 1982. Principles of fishery science (Second Edition). Cornell Uni-
versity Press, Ithaca, NY.
Roff, D. A 1980. A motion for the retirement of the Von Bertalanffy function. Can. J. Fish. Aquat. Sci.
Summerfelt, R. C. 1987. Age and growth of fish. Iowa State University Press, Ames, IA
Weatherley, A H. and H. S. Gill. 1987. The biology of fish growth. Academic Press, Orlando, FL.

Chapter 6

Fish and Shellfish Behavior

In this chapter three aspects of fish behavior are discussed: the extent of migration of fishes,
feeding behavior, and schooling behavior as they relate to fisheries management.
Because an important aspect of fisheries management is knowing the size of the stock to
be managed, migration of fishes is considered. The adult billfishes outlined in Chapter 3 mi-
grate over great distances in the open sea and into waters of other countries, making their
management difficult. On the other hand, some of the nonmigratory fishes (such as reef
fishes) and shellfishes (such as bivalves), can be treated as a unit stock (single stock) for man-
agement purposes. Methods used to determine the extent of migration are discussed, using
as examples two species of pelagic fishes.
Understanding feeding habits of fish and shellfish helps fishery scientists understand pos-
sible fluctuations in population size, for instance, knowing that the lack of suitable foods dur-
ing early larval development can drastically reduce recruitment. Fishing techniques and gear
design are also determined by knowing the foods and feeding habits of many commercial
and recreational species.
Fish schooling behavior determines which fishing techniques should be used. Fishing
with purse seines is practical only on some species of schooling fishes. Schooling can reduce
the impact of predation on some fish stocks and enhance feeding and spawning success.


The concept of a unit stock means that, for management purposes, a group of aquatic ani-
mals can be treated as a single unit. As a rule, management in freshwater fisheries often
deals with a unit stock; for example, a population of fish in an isolated lake, in a river sys-
tem, or in a water shed. The extent of intermingling of stocks of marine fish is often compli-
cated, for example, in the North Pacific Ocean where there is no obvious physical barrier to
widespread migration and intermingling of salmon, Oncorhynchus sp., stocks over broad oce-
anic areas. Rational fisheries management requires knowledge of the extent to which ex-
ploited populations comprise a discrete, uniform, and self-sustaining stock (Fig. 6.1).

Recruits Natural mortality

Figure 6.1. A simple model of a hypothetical
fish stock showing gains and losses. A closed Immigration Stock Emigration
or unit stock would have no immigration or Growth Fishing mortality
emigration of fish affecting stock size.

106 Part Two Fisheries Biology

A difficulty that persists in scientific literature concerns proper names used for subpopula-
tions and acceptable definitions. In 1957, Marr defined some of these groups listed below,
with modifications.
Population includes all individuals of a given species when there are no subspecies or, if
there are subspecies, when their distributions are not discrete. It includes only all individuals
of a subspecies when the distributions of the subspecies are discrete. Obviously, there is gene
flow, or opportunity for such, throughout a population.
Subpopulation is a fraction of a population that is itself genetically self-sustaining. It is
the smallest natural self-perpetuating unit. Although differences between subpopulations
may be small, they are heritable.
Stock is a population or a portion of a population, all members of which are characterized
by similarities that are not heritable, but are induced by the environment. A stock mayor
may not include members of several different subpopulations. An important distinction can
be made between subpopulation and stock is that members of a subpopulation segregate at
spawning time, whereas members of a stock need not.
Race is a term sometimes used by fishery biologists, but it is not recommended because it
is often considered a synonym for subspecies. Different populations of a single species may
exhibit different structural peculiarities owing to differences in environment, or to differences
in heredity arising during long periods of isolation. The study of individuality of the popu-
lations is based largely on these structural differences; for example, numbers of vertebrae and
may be called subspecies, or races. The technique is called meristics (see below).


Biological aspects of fish and shellfish such as growth, mortality, fecundity, etc. are all neces-
sary kinds of information for wise management of fish populations directed at obtaining the
maximum sustainable yield. If the population that is to be managed is homogenous as to
these aspects, then it can be managed as a single unit. However, if the population consists of
a number of subpopulations with different biological characteristics, it may be necessary to
measure the rates for each subpopulation and if significant difference between groups exists,
to manage each of them separately. Hence, before management information is gathered, one
must determine the homogeneity of the population. If it is not homogenous, it is necessary
to find the most expedient means to identify the number of subpopulations and their geo-
graphic distributions.

Techniques Used to Study Subpopulations

Direct Methods. Fishery biologists use two general types of techniques to determine how
large a population unit they are attempting to manage is. These techniques may be direct or
indirect. Direct methods are marking and tagging of fish and shellfish to determine the ex-
tent of their movements; that is, when individual fish, or groups of fish are marked or tagged
in a specific location, at a particular time, and later some of them are recaptured: this sug-
gests the extent of movement. Indirect methods to study subpopulations include counts of
body parts (meristics), body proportions (morphometries), physiological attributes, parasite
fauna (natural marks), and genetics (growth characteristics) (Fig. 6.2).
Common sense can provide clues at the outset of a study to determine the extent of migra-
tion of a stock of fish, such as length of larval life and swimming ability of the species in
Chapter 6 Fish and Shellfish Behavior 107




~ CO .... ON (OVER IO~)

• RollIE (I-IO~)

lt3 1 ABSENT

Figure 6.2. Distribution of the myxosporean protozoan parasite, Kudoa clupeidae, in 1- and 2-year-old
herring off the coast of Maine illustrates little or no intermingling of herring from the southern part of the
fishery with those of the northern part. From Sindermann (1961). J. Wildlife Management. 25(1):41-47.

question. Species with long larval lives would have sufficient time to drift long distances
and intermingle with other members of the population, so it would be expected that they
might be homologous throughout their range. Large, powerful, streamlined, fast-swimming
fish like tunas and marlins would be expected to be far-ranging and hence comprise a single
large population with an extensive geographic range, such as the North Pacific Ocean, or as
with some marlin the North and South Atlantic Oceans. Tagging studies have demonstrated
this to be a fact.
Commercial and recreational fishermen can provide information on the extent of migration
in a population by changes in their catch rates as fish move from one place to another. Pop-
ulation homogeneity has been studied by tagging Atlantic menhaden, Brevoortia tyrannus, as
an example of the direct method.
In an experiment to determine the extent of movement between 1966 and 1969, over 1 mil-
lion menhaden were tagged. Of this number, over 200,000 were later recovered. Results
showed that although the menhaden migrate northward in the spring and southward in the
fall along the Atlantic Coast, there was only a single population of Atlantic menhaden from
New England to the east coast of Florida. Larger, older fish tended to migrate father north
than smaller, younger fish.
108 Part Two Fisheries Biology

Indirect Methods. The following examples are typical of some of the indirect methods of
determining population homogeneity.
Population homogeneity studied by growth characteristics-Pacific herring and Pacific
sardine (Table 6.1). Results from a variety of different racial studies suggest that the Pacific
sardine, Sardinops caerulea, population that extends from southeastern Alaska to Baja, Califor-
nia, consists of several subpopulations. By aging samples of fish collected during the same
year from four areas (Pacific Northwest, San Francisco, Monterrey, and San Pedro), evidence
for differential growth was found that suggest subpopulations.

Table 6.1 Comparison of Growth Curves of Pacific Herring, Clupea pallasii,

From Two Locations.

Length of Herring (mm)

Age Cook Prince William Difference

(yr) Inlet Sound (mm)

4 220.2 205.0 15.2

5 224.9 220.8 4.1
6 252.5 231.4 21.1
7 263.0 243.0 20.0
8 268.1 250.0 18.1
9 272.6 261.6 11.0
10 273.7 261.0 12.7
11 279.4 266.9 12.5
SX = 114.7
n = (n' - 1)
S2 = Sx 2 / (n' - 1) = 30.585
s = 5.53
t= "iN /s = 6.87
Sx 2 = 1,859.21
(SX)2 / n' = 1.644.51
Sx 2 = 214.70
±t for n of 7 = 3.499 at P of 0.01, t / 2 = 1.75, so that the probability of
a difference as great as 6.87 occurring by chance is extremely remote.
'Data from Rounsefell (1930). Bull. U.s. Bur. Comm. Fish. 45:227-320.

Analysis of subpopulations of spotted seatrout, Cynoscion nebulosus, were made using

growth and tagging data from several locations along the coast of Florida (Cocoa, Flamingo,
Fort Myers, Cedar Key, and Apalachicola). Growth rates from these areas showed differences
that suggested subpopulations of this species. These findings agree with tagging results
wherein 95% of recaptured seatrout moved less than 30 mi (48 km) from where they were
tagged. Results of two studies, growth characteristics and tagging, suggest a close associa-
tion of seatrout with the local environment.
Population homogeneity studied by morphometrics-yellowfin tuna. The study of possi-
ble subpopulations of yellowfin tuna, Thunnus albacares, in the equatorial Pacific was studied
by the use of morphometrics. Samples of yellowfin measured across the Pacific included
such measurements as head length and length of pectoral fin of fish of the same size in sam-
Chapter 6 Fish and Shellfish Behavior 109

pIes taken at different longitudes from the western Pacific Carolinas to waters off Panama
and Costa Rica. Orderly changes were found in measurements (clines) from west to east in
samples taken within 10° latitude of the Equator. The conclusion that the population of
North Pacific yellowfin tuna is at least semi-independent is somewhat at odds with what
would be expect based on the physical characteristics of the species and other scientific data
Size frequency data, for example, from the same general areas show that only small yel-
lowfin existed to the west and only large yellowfin occurred in the eastern Pacific that
strongly suggests that the species moves across the Pacific along the Equator from west to
east, growing along the way. Subsequent tagging and migration models confirmed that these
fish move freely over wide areas of the Pacific. The scientist who carried out the morpho-
metric study commented in his report, "There are numerous reasons to be skeptical of con-
clusions based on data of this kind. Foremost perhaps is the fact that racial studies which
have shown highly significant statistical differences between stocks have been contradicted
by tagging studies."
Population homogeneity studied by meristics (races)-Pacific herring. Another technique
used to study herring subpopulations was by meristics, in one study the number of vertebrae
of Pacific herring, Clupea pallasi, were counted (Fig. 6.3). From 1950 to 1958, herring from
northern Baja, California and southern California, showed that there are significantly differ-
ent numbers of vertebrae in sardines between the two areas, suggesting two racial groups.
Samples of Pacific herring were collected from five contiguous areas along the west coast of
Vancouver Island, Canada, to investigate the possibility of subpopulations. Over 22,000 ver-
tebral counts were made involving 12 consecutive year classes. The conclusion from the
study was that two subpopulations were present and that, with the exception of one area,
each had an essentially separate spawning run. Comparison with conclusions drawn from a
herring tagging program in this area showed agreement.
Population homogeneity studied by meristics and morphometrics-anchoveta (Fig. 6.4).
The anchoveta, Centengraulis mysticetus, a principal tuna bait species in the eastern tropical
Pacific Ocean, was studied to determine if there is more than one population of this species.
The anchoveta could not be tagged by the direct method because there was no suitable tag.
Also, the low level of fishing effort, short life span, and high natural mortality of the ancho-
veta suggested that recoveries of tagged anchoveta would be too low to make the tagging re-

53.0 020
019 023
025 021

g 52.6
Figure 6.3. Mean vertebral count of Prince c
William Sound (Alaska) herring of 9 year
classes and March through June air o
temperatures. Data from Rounsefell and 52.4:t::--,-----,-----r-----r----.---.---,--,-
Dahlgren (1932). Bull. U.S. Bur. Fish. 47: 38 40 42 44 46
263-291. March·June air temperature OF
110 Part Two Fisheries Biology

suIts conclusive. Therefore, indirect population studies using both morphometric and
meristic techniques were used. Samples collected from 10 stations on the Pacific Ocean, ex-
tending from Mexico to Colombia, were sampled using four meristic characteristics (numbers
of anal fin rays, scales in longitudinal series, vertebrae and gill rakers) plus three morpho-
metric characteristics (head length, body depth, and eye diameter) for anchoveta in their first
year of life. The characteristics used showed significant statistical differences among the 10
localities and suggested that the anchovetas belong to more than a single population. The
majority of areas differed in four or five of the seven characteristics tested. This strengthened
the conclusion that there are semi-independent, or independent subpopulations because these
exist from year to year, regardless of whether they are phenotypic or genotypic.

Figure 6.4. Distribution of three northern

anchovy subpopulations based on transferrin
allele frequencies. Note area of overlap
between the northern and central
subpopulations. Morphometric and meristic
studies supported the genetic findings. From
Vrooman, Paloma, and Zweifel (1981). Calif.
Fish Game 67(1):39-51.
Chapter 6 Fish and Shellfish Behavior 111

Population homogeneity studied by electrophoresis-striped bass. This technique causes

movement of colloidal particles suspended in a fluid when an electric field is applied (Fig.
6.5). The process displays the genetically controlled structure of proteins, thus permitting
fishery biologists to differentiate discrete fish populations. A study of five proteins in the
striped bass, Marone saxatilis, from three river systems in upper Chesapeake Bay was made.
The protein analyses showed that the populations were unrelated to age, sex, or time of col-
lection of the fish. The results of the study indicated that striped bass in major river systems
(except Patuxent and Potomac) form discrete subpopulations; hence, a certain level of fishing
pressure for one river may be unsuitable for another. The scientists reporting on this study
pointed out that their results should be checked by other types of techniques, preferably a di-
rect approach like tagging because of the degree of uncertainty inherent in the electrophoresis
method. If the results are confirmed, then different management plans may be required for
the three major rivers.


I 3
I 25
C[~ 26
I 16
Figure 6.5. Cluster tree diagram of sample 9
sites by proportion of northern anchovies 10

transferrin alleles from blood samples ""'-
(transferrin electrophoresis technique). 13

~ r-"" 14
Clustering sequence is determined by the ""'- .....
distance of the vertical bars from the solid L-r- 20
vertical line. Example: 23 and 26 sites are most 12
similar, then 14 and 20 are next. From 6
Vrooman, Paloma, and Zweifel (1981). Calif. 8
Fish Game 67(1):39-51. 7

Population homogeneity studied by fecundity-Pacific salmon. The fecundities of Pacific

salmon, Oncorhynchus sp., from several different stream systems were studied. Because the
salmon home to native streams, reasonable expectation that fecundities would vary among
home streams was supported. Higher fecundities could result from higher mortalities by
salmon in streams that have somewhat adverse environmental conditions, and lower fecun-
dities might result in salmon from more optimum conditions in other streams.
Population homogeneity studied by otolith/scale morphology (Fig. 6.6). In a study using
otolith morphology on the deep-slope red snapper, Etelis carbunculus, in the Pacific Ocean,
Fourier-shape analysis descriptors, and other shape indices, linear proportions and dry
weights were used. Collections of fish from Hawaii, Vanuatu, Fiji, French Polynesia, and the
112 Part Two Fisheries Biology






Figure 6.6. Typical size and shape differences betweens male and female halibut otoliths. Although
not accurate enough for studies of halibut subpopulations, they have been successfully used to
distinguish subpopulations of other fish species. From Int. Pacific Halibut Comm. Tech. Rept. 29.

Commonwealth of Northern Marianas Islands were compared and permitted recognition of

regional otolith shapes and weight differences. Using Fourier-shape analysis on Atlantic
salmon, Salmo salar, scales to distinguish and identify two stocks met with success of almost
Population homogeneity studied by indirect methods-summary. Generally, when the
value of the resource is high and financial resources permit, and further, when direct meth-
ods are not possible, several different indirect methods should be used concurrently. Using
more than one technique lends support to results, especially when genetic and environmental
differences are measured.

Underwater Radio Telemetry

During the late 1950s a new technique was developed to trace the movement of land and
aquatic animals by attaching to their bodies an electronic battery-powered device that emit-
ted signals that could be followed in their natural environment by a remote receiver. A prim-
itive form of telemetry is the cowbell (transmitter) and the cattle owner's ear (receiver) that
is used to follow the movements of cattle. Today, technology is evolving rapidly in a much
more complicated form of telemetry. Scientists find this technique to be useful for many rea-
Chapter 6 Fish and Shellfish Behavior 113

sons. Biologists who study movement of large land animals, at least, can follow the study
species if a visual mark or tag is placed on them. Fish, on the other hand, require other
means for following because after release they disappear from sight, and all the data on
where they were tagged, how far and how fast they have travelled is revealed only when the
tagged animal is caught and the proper agency notified.
Radio signals emitted by the electronic device are directional so when several receivers
pick up the signal its location can be determined. Fish equipped with small transmitters
(which do not alter their behavior) can provide valuable data on environment parameters as
to where the fish live, short term movement (daily), and migrations, and are also of value in
conservation programs.
The amount of dissolved minerals in water weakens signals sent by transmitters. Ultra-
sonic transmitters similar those to used by the U.S. Navy avoid this problem. Problems are
also encountered in attaching the transmitters to the animals so as not to bias the results;
some are attached internally (surgically or otherwise), others externally. Antenna systems are
needed to receive signals. Tracking can be done by a number of different techniques, in
boats, on foot, in aircraft, and by satellites.
Temperature and movement of fish in Louisiana lakes are studied using transmitters with
a probe that sampled the water temperature. Upon locating the fish, signals sent indicated
water temperature as well as location.
Biologists in Australia tried to study the foraging behavior of the spiny (western rock) lob-
ster, Panulirus cygnus. Transmitters glued to the carapace of the lobsters had a operating life
of about 3 weeks. Early studies were beset with technical problems; aerial wires on the trans-
mitters were bitten off by fishes or abraded by the reef, and receivers broke.
Odd results sometimes come from studies. When a loggerhead turtle, Caretta caretta, fitted
with a transmitter was traced by the Nimbus satellite, the National Oceanic and Atmospheric
Administration (NOAA) encountered a curious result. Over an 8 month period the 212 lb (96
kg) turtle traveled 800 mi (1,287 km) from near Gulfport, Mississippi to offshore Brownsville,
Texas. There the transmission stopped and mysteriously started anew. The transmitter was
traced 500 mi (804 km) to the north of Port Arthur, Texas to a small town in Kansas. The line
on the 7 lb (3.2 kg) transmitter holding it to the turtle had parted and the transmitter drifted
up on a beach where it was found and taken by the finder to his home for use as a doorstop.


The migration of fish is a purposeful, concurrent movement of a majority of the population,

not random movement. Knowing when and where exploited fish move is fundamental to
fishery biology. The mechanism(s) fishes use for orientation and navigation are intriguing
but poorly known. However, well-described fish migratory patterns can aid fishery biolo-
gists in their management planning regardless of the mechanisms involved.
Studies of animal migration began many years ago. The mystery that formerly cloaked
the periodic travel of animals has been largely dispelled, especially in the case of birds.
There are many gaps, however, in our knowledge of the subject. As in other scientific stud-
ies, the majority of work has been done on animals that lend themselves to experimentation.
Insects are such a group and have received considerable attention as a result. The work of
Von Frisch in 1950 in Germany, demonstrating the ability of honey bees to navigate and com-
municate positions to their coworkers, is a classic. Likewise, bird migration has received
much attention. Extensive studies have been made by Lincoln extending over about 30
114 Part Two Fisheries Biology

years. The yearly migrations of 25,000 mi (40,225 km) made by the Arctic tern are spectacu-
lar and well-known. Homing pigeon migration also has been extensively studied.
The periodic occurrence of fishes at certain locations in the sea has been observed for
years, yet their migrations have not received attention as have birds and insects. Roule (1933,
translated from the French), writing of the journeys and migration of fishes, summarized
knowledge up to that time. It accurately portrays the migration of many European species
and includes the classic work on eels. It describes migrations of shad, herring, sardines, and
tunny, and many other species.
With advanced methods of marking (removal of fins) and tagging fish, the literature blos-
somed with the results of many studies that give more detail on the time of migration, rates
of speed, and directness. Investigators must take into account the distribution of fishing ef-
fort or captured tagged fish will not show where fish are moving, but where the fishing ves-
sels are operating in relation to the point of tagging.
"Drift migration" in the sea is used by many marine species to disperse eggs and weak-
swimming larvae. Eggs of most marine species are pelagic. From spawning grounds eggs
and larvae usually drift to nursery grounds where conditions are suitable for survival. Tides
and currents carry eggs and early life stages by drift migration sometimes over thousands of
miles. Life requirements for larvae and juveniles may differ greatly from adults, for exam-
ples, salmon and eels. Despite enormous distances between spawning grounds and nursery
grounds, the location of the nursery is fixed.
At least four kinds of controlled migrations of juvenile and adult fish have been identified.

1. residentially restricted-territorial (reef fishes);

2. broad (temporary) residential range (menhaden);
3. restricted migratory pattern, such as a short inshore-offshore migration (mullet);
4. extensive migratory patterns, seen in fishes such as tunas, marlin, and sailfish with little affinity
to land.

Migration Patterns-Pacific Salmon

Some early studies on homing abilities of the Pacific salmon, Oncorhynchus spp., are not
entirely reliable. After marking downstream migrant young fish by removing combinations
of fins, the same streams were subsequently searched to find marked adults moving up-
stream. Little effort was devoted to adjacent streams to find out how much straying of
marked fish was taking place. Despite this bias, the home stream theory is well-established;
in addition, we now know that salmon do not remain off the mouths of home streams until
it is time for the spawning migration, but instead migrate widely over the North Central Pa-
cific before returning to their home streams to spawn (Fig. 6.7). The North Pacific Salmon
Commission, which resulted from the United States controversy with Japan over fishing
salmon in the aforementioned area, has advanced knowledge of salmon migration consider-
ably. Three methods to study the extent of Alaskan red salmon migration employing parasite
tags, plastic tags, and body measurements (morphometrics) suggest that a proportion of the
salmon found as far west as 180 W longitude are headed for Alaskan streams. The propor-

tion increases as samples are taken eastward.

Migration Patterns-Pacific Tuna, Yellowfin

The Pacific yellowfin tuna, Neothunnus macropterus, provides an interesting example of a
strong swimming species of which the extent of migration was unknown for many years.
Chapter 6 Fish and Shellfish Behavior 115

Figure 6.7. Diagram of ocean migrations of pink salmon stocks originating in southeastern Alaska and
British Columbia. From Royce, Smith, and Hartt (1968). U.S. Fish and Wildl. Servo Fish. Bull. 66(3):

Near the Equator, approximately in the center of the Pacific, large schools of yellowfin tuna
extend in a narrow north-south band, and as far as the eye can see, to the east and west.
Each fish found in the open sea may weigh up to 200 lb (91 kg), and those found close to the
islands are generally under 100 lb (45.4 kg). Yellowfin found near the surface close to islands
and atolls can be caught by trolling gear. In the open sea where they are found at consider-
able depths, down to about 600 ft (loa fathoms), they are captured by floating longline gear.
At first glance it may seem that the movement of these fishes is only of academic interest.
But it goes beyond this. An understanding of the migration of a commercially important pe-
lagic species of fish is of considerable value in finally determining the production from that
fishery. If fishermen do not know when and where to find schools, their catches can be quite
small; yet to expect fishermen to undertake extensive open ocean exploratory fishing is un-
The scientific staff of the U.s. Fish and Wildlife Service (now NMFS) stationed at Honolulu
has been studying this species since 1950. Part of their job is to describe the structure of the
population of yellowfin and their abundance. Attempts by these scientists, as well as some
in Japan, to determine whether yellowfin tuna along the Equator are one big population or
116 Part Two Fisheries Biology

consist of several independent or semi-independent populations (subpopulations), has pro-

duced conflicting results. Reliable information is difficult to come by because of the vastness
of the area of study and the rather expensive sampling in this area.
In an attempt to track movements of these giant fish directly, researchers used spaghetti
tags (a plastic loop threaded through the back muscles of the tuna) to provide an identifying
mark should the released fish be recaptured. The probability of tag recoveries in this vast
area is quite small. Although Japanese vessels fish all along the Equator and U.S. Fish and
Wildlife Service research vessels make exploratory catches there, the fishery is not steady and
intensive. The fish caught that are suitable for tagging limit how many can be released, again
decreasing the probability of recovery. A tagging program in this fishery is of considerable
potential value because sometimes it takes only the recovery of a few tags to increase the sci-
entists' knowledge of migratory habits of yellowfin. In a migration study during 1955 and
1956, more than 1,000 spaghetti tags were put on yellowfin in the Line Island area (Central
equatorial Pacific). As expected, the results were somewhat discouraging because only three
tags were subsequently recaptured. Two of those recovered were caught close to where they
were tagged. They were small fish; one was at liberty for about 200 days, the other approx-
imately 300 days. The most distant recovery of the three tagged at Christmas Island was
subsequently recaptured by Japanese longline about 800 mi (1,482 km) east about 13 months
after the fish was released. Because it was caught originally with surface trolling gear and
later by longline gear deep in the ocean suggests a change of habitat from surface to depths
as the fish grew. More importantly, it was found east of the tagging position.

Migration Patterns-Pacific Albacore

The long-range migration of another Pacific species is worth mentioning. Possible migra-
tions of the albacore, Germo alalunga, studied using morphometric methods, suggested there
is no intermingling between the two groups of albacore in various widely spaced locations in
the Pacific. Subsequently tagged albacore moved from off the coast of California to Tokyo
Bay. These tagging results showed not only that movement is in the opposite direction and
that the movement was not necessarily in a straight line, but rather that these albacore appar-
ently move northward along the Pacific Coast of the United States, across the Pacific well
north of Hawaii, then southward along the east coast of Asia to Japan (Fig. 6.8). Again,
movement was found generally along a major current system. The failure of morphometric
studies on albacore to agree with tagging results made scientists aware that indirect methods
can be misleading.
Marr (1957) discussed the use of anatomical characteristics in subpopulation studies:
The use of anatomical characteristics in defining and describing subpopulations has at least
two limitations. First, it is well known from both empirical and experimental evidence that
body form, numbers of vertebrae, etc., are influenced by environmental variables (such as
food and temperature, for example). Obviously, then, in using such characteristics the risk
exists of studying the effects of environmental conditions rather than the effects of genetic
isolation. Second, even if the characteristics used are genotypic, their frequency distribu-
tions generally overlap, often to a large degree. If significant differences are found between
the samples, these have usually been interpreted as indicating that the two samples were
drawn from two distinct subpopulations. Actually, this indicates only that they were not
both drawn from the identical population, the possibility of considerable intermixing still
Since that time numerous reports about racial study methods of fish indicates that tagging
methods are clearly favored to gain direct evidence of migration.
Chapter 6 Fish and Shellfish Behavior 117

Figure 6.8. Model of albacore migration in the North Pacific Ocean, by age groups. From Otsu and
Uchida (1963). U.S. Fish and Wildlife Service, Fish. Bull. 63(1):33-44.
118 Part Two Fisheries Biology

Migration Patterns-Billfishes
Billfishes are found in all warm seas. These large fish, some reaching about 1,500 lb (680
kg), are powerful swimmers and are eagerly sought by commercial and recreational fisher-
men. Recreational fishermen favor marlin and sailfish. The family Istiophoridae includes
four species of marlin: white, blue, black, and striped; and the longbill spearfish, the shortbill
spearfish, and sailfish. The swordfish is in a family by itself (Xiphiidae). It was formerly be-
lieved that billfish stunned their prey with a blow of the spear before devouring them. How-
ever, evidence indicates that billfish can exist quite readily without a spear in food getting.
The current theory holds that the spear may reduce the resistance of the water to the billfish's
body. Tagging and recovery data, from assistance by anglers, showed a migration of 3,450 mi
(6,389 km) by a striped marlin in the Pacific swimming west from the coast of Mexico. Blue
marlin are known to cross the Atlantic Ocean from the Virgin Islands to Africa.


Insect studies have demonstrated that the sun can be used as a reference point in migration.
In the case of marine animals, reference points are not so simple. Marine animals that are in-
timately associated with the bottom, such as shrimp, tend to move into deeper water and far-
ther from land with increasing age. In the Florida Tortugas fishery, confirming evidence of
this has been gathered by tagging experiments and measurements of sizes for shrimp from
all over the fishing grounds.
It is even more difficult to suggest a reference framework for pelagic animals. Although
yellowfin tuna, albacore, and Pacific salmon can follow current systems (as mentioned ear-
lier), it must be remembered that a fish in an ocean current might have as much difficulty
telling the direction of flow as a man in a small boat in heavy fog. There is no reference
point. Sight feeders have reference points available that can be used in the open ocean to
maintain their position in a current. Even though the fish are moving in a mass of water, like
the man in the rowboat, the movement of plankton and other visible objects in the water rel-
ative to their movement on different headings could suggest the direction of the current.
The idea that fish may seek certain temperatures has been proffered. It has also been sug-
gested that, while over the great expanse of the Pacific Ocean the temperature change is con-
siderable, the change over the length of this body may also be so small that the fish would
be unable to move continuously toward the temperature sought.
That the sun may be a reference point for fish has given rise to a study of the anatomy of
the so-called "pineal eye" of the Atlantic bluefin tuna, Thunnus thynnus. Although there is no
indication of a lens or retina in its eye, a structure filled with a clear jelly leads from its brain
to the top of its head. It is remarkable that the skin at this point, instead of being dark and
opaque, lacks pigment and admits light. Some Atlantic bluefin tuna follow the Gulf Stream
for a time, eventually leaving it and heading for Nova Scotia.
An animal needs two components for navigation: a compass component for travel on a
straight course, and a map component to relate its position to its destination. The map com-
ponent can be thought of as a representation of the earth's surface in order to select a correct
heading. Many fish, birds, and insects apparently navigate by the sun, and certain birds
seem to orient by stars. We have learned how lower animals guide their direction by posi-
tions of heavenly bodies, but how do they orient themselves? We can take a homing pigeon
inside a closed basket to a strange place and release it, which is equivalent to blindfolding a
Chapter 6 Fish and Shellfish Behavior 119

person and putting him in a strange town with no name or directional signs pointing the
way to his home. To return home, in say, New York, he has to know which direction New
York is from his present, lost, position. Map orientation remains a puzzle to scientists, who
still cannot explain what cues animals use.
Changes in polarized light, barometric pressure, low level sounds (infrasounds), and pres-
ence of magnetic particles in fishes (to sense the geomagnetic field) have been examined to
try to solve the navigational map and compass puzzle, but so far have not provided convinc-
ing evidence to the mechanism of orientation. Scientists, who have all this advanced technol-
ogy, continue studying these small animals yet confess they still cannot explain how fish
navigate over miles and miles of ocean.


Fishery biologists study foods eaten and feeding habits of fish and shellfish for a variety of
reasons. The kinds of foods eaten must be determined because their abundance and availa-
bility can affect growth and survival of a stock of fish. Certain specific kinds of food items
are critically important in early larval stages and must be available or enormous numbers of
larva die. When aquatic animals feed and do not feed influences their growth. For example,
during cold weather when many species reduce their feeding activity, their growth rate is re-
In Chapter 14, the urgency of determining total seafood production from the world's
oceans is discussed. Also, the method of examining the food web all the way from primary
production to harvestable fish. To do this, the approximate energy transferred from one
trophic level to the next must be known, as well as the proportionate amount of food in-
gested to increase the weight of the animal (called the conversion ratio). The difference be-
tween these two figures is the amount of food used to satisfy only the maintenance
requirements of fish (respiration, swimming, etc.). In aquaculture, this ratio of food eaten to
weight increase is highly important and, indeed, may be critical to the success of a commer-
cial fish farming operation. This is especially true if feed costs are high.
Understanding feeding behavior is vital to successful fishing for many species of commer-
cial/recreational fish. Many marine fish eat a diverse array of foods. Their feeding habits
and their process of searching for and capturing food are likewise diverse. When the kinds
of food fish eat and the way they feed are known, suitable hooks and baits (real and artificial)
can be presented to the fish in a way to invite them to bite.
Diet usually changes as fish grow larger. Some species shift from a plant to an animal diet
as they mature from larval to juvenile and adult life stages. The most pronounced change oc-
curs when metamorphosis from larval to juvenile stages takes place. In some species, juve-
niles may have a greatly different diet than adults, cod for example, but usually, as juveniles
and adults, fish are opportunistic feeders and consume whatever is most numerous and read-
ily available.
Seasonal changes may reflect food availability. Availability and selection should not be
confused. In order to say that fish "select" or "prefer" certain kinds of food in nature, a fish-
ery biologist must be aware of all foods available to the fish in its environment, then compare
them with what is actually eaten. If their stomachs contain only certain food when a wide
variety of organisms are abundant, this suggests a preference or selection (Fig. 6.9). Within
the restrictions of artificial conditions in the laboratory, experiments can be designed to offer
a variety of foods and determine if fish select one single food item, or several food items.
120 Part Two Fisheries Biology


Stomach contents


Figure 6.9. Particle-size distributions from

o ..... ~
1"'"0_--0 ___0 Sediments
stomach contents and sediments being fed
..... - - - 0 - - - 0 - - - 0 - _ 0 _ _ -0 _ _ - 0

0~--,1--~~1~~1=-6-~1~,-1---r-,~~,==~,~ upon by mullet, Mugil cephal us, at Sapelo

o 50 100 150 200 Island Beach Station, Georgia. From Odum
(1968). Limnology and Oceanography.
Diameter of Particles (microns) 13(1):92-98.

Because carnivorous fish must first detect the food it is to eat, success of food acquisition
depends upon the nature of each species' sense organs and behavior pattern. Visual, olfac-
tory, auditory and tactile senses are all used to a greater or lesser extent by different species,
but usually in combination.
Predatory species, especially those that actively seek prey, depend heavily on vision, some-
times coupled with the tactile (lateral line) system to locate prey. Such species may lie hid-
den and immobile to surprise their prey. Sensitive taste cells on lips or barbels may assist
bottom feeders that consume detritus, or slow-swimming or sedentary prey. Some species
have extremely elongate pelvic fin rays that appear to be tactile organs. Still other deep wa-
ter species possess luminescent lures that bring prey close enough for capture.
Adaptations such as streamlined shapes, rearward positioning of median fins, powerful
dorsal muscles, deep, often pronouncedly forked, caudal fins of fast-swimming predators,
and the effective camouflage of "passive" predators, are all clues to feeding habits. Jaws,
teeth, and intestines of fishes are well-adapted to dealing with certain food substances. Pred-
ator's jaws generally have a fairly wide gape with long, sharp, backwardly directed teeth to
prevent escape of the prey. Some fish, for instance piranhas, that tear pieces out of a prey or-
ganism larger than themselves, possess canine-type teeth adapted for a shearing motion.
Predators generally have comparatively short intestines: the ratio of intestinal to body
length of a ladyfish, Elops saurus, a carnivorous predator, is about 1:24; that of a similar-sized
milkfish, a detritus and algal feeder, is about 6.5:1. As a rule, herbivores possess a relatively
longer intestine than carnivores providing the necessary increase in adsorptive area. Small
stomachs and long intestines are characteristic of fishes that feed at frequent intervals on
small organisms. Conversely, bathypelagic fishes, which may encounter food only infre-
quently, have large, thin walled, distendable stomachs together (in some species) with a
hinged mouth. Some of these species are known to swallow prey two or three times their
own size.
Plankton feeders, such as herring, sardines, and basking sharks, strain food organisms
from the water by means of elaborate gill rakers. The degree of elaboration reflects the de-
gree to which fish depend on this mode of feeding. Thus, in contrast to plankton feeders,
predacious freshwater fishes such as pike have small, inconspicuous gill rakers.
Chapter 6 Fish and Shellfish Behavior 121

Certain stimuli trigger feeding in fishes, for example, in most larval fish, size and move-
ments of the prey organisms is sufficient to cause the larvae to strike it. What may appear to
be food of the proper size may not be consumed by fish larvae. If the movement of the po-
tential food item (even though it is abundant) does not attract their attention, then the larvae
will starve. The most publicized example of feeding stimuli is the so-called "feeding frenzy"
of sharks. When the blood of a wounded fish is placed in the water, sharks become highly
aggressive and bite or devour almost any object in the area, including each other. Sounds of
struggling fish also cause sharks to become aggressive.
Economics of feeding is critical to survival and growth of a species. Fish feeding takes
into consideration the amount of energy required to locate, capture, and ingest food, and the
subsequent benefits they obtain. Figure 6.10 shows that the net energy gained by eating var-
ious-sized food particles has a range for a particular predator species. At the lower limit of
the range, effort expended to find and eat tiny organisms is greater than the energy received
from their consumption. The same applies to very large organisms: effort to locate, capture,
and chew foods is greater than the energy derived. The food particles in the center of the
graph are of optimal size, giving the fish the greatest gain per unit energy cost.



Figure 6.10. A hypothetical graph showing UNIT COST
net energy gained per food item plotted against
food particle size. Average gain is marked by x
the horizontal line dividing the figure into the 8
upper section A and the lower section B. The
optimal diet includes all those items that yield
a net gain that is greater than the average gain. _--_OPTIMAL _ _ _.I
From Pitcher and Hart (1983). Figure 4.1 AVI
Publishing Co. SMALL LARGE

Some fish feed only at night, others only during the day. Some are more active feeders at
certain times of the day or night (twilight and or daybreak), and some are more active at cer-
tain times of the year. If the objective is to find the kinds and amounts of food eaten, obvi-
ously an understanding of feeding chronology is necessary. There is no substitute for in situ
studies of the feeding habits of fishes. As indicated above, clues as to feeding habits are of-
fered by the anatomical structures of fish and shellfish; but, to confirm suppositions and ob-
tain greater insight, underwater observations are useful.
Study of stomach contents must be done with caution to ensure collection of representa-
tive data on kinds and proportions of food eaten. To obtain representative data:

1. Samples of fish stomachs should be removed and preserved as soon as fish are caught to pre-
vent the further digestion that goes on even after death and hinders identification of individual
food items. This is especially critical in warm tropical and subtropical seas.
2. The stomach contents of fish with large stomachs should be placed in cheesecloth bags to keep
them together loosely, allowing close contact with the preservative to prevent deterioration.
3. Number each stomach bag and record fish size (length and weight), sex, time and day collected,
and location because the kinds and amount of food eaten may vary. The contents should be ex-
amined with these variables in mind.
122 Part Two Fisheries Biology

4. Remember that remains of hard-shelled organisms, that is, shrimp exoskeletons and fish bones
present in stomachs are digested slower than soft-bodied organisms. Frequent appearance of
the same items may indicate importance in the diet.

Researchers use a variety of methods of expressing results of food eaten by fishes. Some
use simple counts of organisms found, others weigh or use volumes of the various food
items in proportion to the total weight of all food items. Other investigators give ranking to
the various food items depending on their estimated food value to the fish, that is, a thick-
shelled animal would be heavy, but only a portion of its total weight is digestible by the fish.
Occasionally researchers will use more than one method to estimate the important food items
and compare the results of the methods. Analyses of stomach contents are usually presented
in pie charts or tabular form.
In summary, the diet and feeding habits of fish are extremely diverse, and representatives
of fishes exhibiting these characteristics may be found occupying most feeding niches in the
sea (Fig. 6.11). Anatomical structures and diet are intimately linked with one another and
with the feeding habits of the fish. Slight changes in morphology may be accompanied by
considerable alterations in diet and methods of securing food; for example, the practice of re-
moving one or both large claws from a stone crab can seriously affect its feeding behavior
and types of food eaten. That some fish select certain food items requires careful observation
on the range of foods available to them to ascertain that they indeed select foods rather than
consume large numbers, items simply because they are very abundant and available. Energy
required to obtain and devour food is related to energy obtained. Properly designed studies
of stomach contents can produce evidence of the most important foods required by fishes .


Eliii' L..... Juvanllel

• ~A:t:p Adulto


Figure 6.11. Pacific whiting prey fish eaten at various life history stages. (Asterisk indicates major
prey species.) Modified from Livingston and Bailey (1985). Mar. Fish. Rev. 42(2):16-22.


Many animals travel in groups or pods known by such various and colorful terms as coveys
(partridges), gaggles (geese), prides (lions), swarms (bees), herds (elephants, etc.), and shoals
or schools (fishes). Within any particular group of animals this behavioral patten may be
widespread. Yet not all fishes school. The fact that many species of fish of commercial or
recreational importance to humans form schools and are therefore subject to easier capture by
fishing gear permits great harvests from the sea. But it can also contribute to their depletion.
There has long been speculation as to why groups are formed; investigations of this be-
havior have been attempted for many years and recent studies have shed light on the me-
chanics and value of schooling. At first glance, it appears that survival depends on whether
Chapter 6 Fish and Shellfish Behavior 123

a prey fish is alone or swims with a school. To learn if some survival benefit is gained from
schooling, a theoretical study employing a mathematical model was made. Findings suggest
that if a fish is a member of a school, it has a better chance of living longer than if it swims
alone, because predatory fish searching for food have less chance of encountering a school
than of encountering individual prey fish scattered throughout wide areas of the oceans.
This study also suggests that the larger the number of fish in a school, the fewer times the
school would be detected by predators. On the other hand, schooled fish would seem to
have a better chance to detect predators.
The limited eating capacity of any predator is another factor that influences higher sur-
vival of the schooled fishes. A predator will encounter a school of prey fish only occasion-
ally; because it can eat to capacity only, the larger the school the less the probability of any
individual member has of being eaten. Let us assume that 10 fish is the most a certain pred-
ator can eat. In a school of 100 prey fish, the chance each has of being eaten is 1 in 10, but if
the school consists of 1,000 fish, the individual chance of being eaten is reduced to 1 in 100.
If, on the other hand, a predator encounters individual prey fish fairly regularly, at widely
spaced intervals, it would never become satiated, and assuming it is a highly efficient preda-
tor, it would consume every fish it encounters.
Schooled fish that prey on tiny swimming invertebrates have a decided advantage in feed-
ing. A tiny prey animal may be capable of darting quickly for a short distance to avoid a sin-
gle predator approaching them with an open mouth, but such tactics would not save it from
a school; in avoiding one open-mouthed predator, it may dart directly into the path of an-
other. Mackerel, Scomber sp., for example, have been observed to swim with their mouths
and gill covers open wide while feeding on copepods.
From the air, large fish schools look like giant amoebas pulsing through the ocean as they
move forward and side to side together. When viewed from a distance underwater, a fish
school might appear as a monster to a predatory fish, giving rise to speculation that a school
frightens away predators. Aerial photographs show menhaden, Brevoortia tyrannus, and her-
ring, Clupea sp., schools as large masses that to an individual predatory fish could look so
formidable as to discourage even the hungriest of predators from attacking.
A study of the effects of sound on menhaden noted that the species formed a tight school
by crowding together in frenzied swimming when a recording of porpoise calls was played.
This type of response may be interpreted as a defense against noisy predators.
Fish swimming in schools may require less effort for individual fish than if they swam
alone. The lead fish of the school exerts no more effort than their usual (solitary) effort to
swim; however, the other members in the school may receive the benefit of less exertion by
riding along in the wake of the lead fish, just as surfboarders can ride "free" behind a motor-
Beside survival value and feeding advantages, schools may make successful reproduction
more certain. During the spawning act in the sea most pelagic spawners release their ova or
sperm when the sexes are in close proximity (Fig. 6.12). Mass fertilization tends to ensure
that a superabundance of the milt comes in contact with the eggs. In schooling fishes, the
milt may be so dense that the water is white for miles around. When individual males and
females of nonschooling fishes pair off to spawn, the milt from a single male, although con-
taining a superabundance of sperm cells, may be carried away from the eggs by currents and
many eggs go unfertilized.
Because large fish swim faster than small ones, fish in any school are generally uniform in
size. Muscle weight increases as a cube of the length of the fish, but the surface area of the
fish increases as a square of the length. This means that there is less frictional resistance per
124 Part Two Fisheries Biology

Figure 6.12. Tight school of California

mackerel. Photo courtesy George Mattson,
U.S. Bureau of Commercial Fisheries.

gram of muscle; hence, larger fish are able to swim faster and will outswim the smaller fish
in a school. Experienced tuna fishermen use this knowledge to increase their catches. They
know that if they begin catching small fish from a school, small ones are all they will get, so
they discontinue fishing to conserve their hard-to-get live bait. Conversely, when they locate
a school of large fish, they get greater production from their bait.
Occasionally, schools of fish consist of more than one species, but this usually occurs only
when both the food habits and the size, shape, and color of both species are similar. In Ha-
waiian waters and off Central and South America, yellowfin tuna, Thunnus albacares, and
skipjack tuna, Euthynnus pelamis, are caught from the same schools. Porpoises and tunas will
school together as is pointed out in Chapter 8.
Although suggestions and theories have been advanced concerning the value of schooling,
the benefits of high survival rate seem to outweigh the disadvantages, but no firm answer
can yet be provided for the reason or reasons for such behavior. The subject is complex and
the experimental difficulties are many. Whatever it may be, marine fisheries benefit from the


Many decades ago scientists had to satisfy their curiosity and desire for underwater observa-
tions of fishes in their natural habitat using heavy helmets and air hoses that restricted their
vision and depths to which they could descend. Self-contained underwater breathing appa-
ratus (SCUBA) vastly increased man's range beneath the surface so he could observe marine
animals and plants in their natural surroundings with little effort (more details on the use of
spear fishing using SCUBA follows in the description of kinds of recreational fishing). Later
research vessels were modified with special "windows" in their hulls to observe near-surface
fish; other scientists used sea sleds towed by vessels to observe fishing gear performance at
moderate depths. Then came submersibles that greatly increased the dive depths and the
ease of making and recording observations on fish feeding behavior, schooling, spawning,
fishing gear performance, plankton distribution, etc. In contrast to submarines, submersibles
may be manned or not, but require surface support; submarines are always manned and
Chapter 6 Fish and Shellfish Behavior 125

need not have surface support. During the 1960s over $100 million was invested in a fleet of
about 60 submersible vehicles for possible scientific uses. In early 1970 they were left idle
due in part to the lack of federal support because of Viet Nam War expenditures. Some
manned but more unmanned submersibles are available today for scientific uses, but the de-
mand is not great. In a 1993 study a submersible was used to study the deep water spawn-
ing habits of lingcod, Ophiodon elongatus, an important commercial and sport species in the
northwest Pacific. Male lingcod are nest guarders and are highly susceptible to capture using
traditional gear during the spawning season. The study indicated that significant spawning
took place in depths between 90 and 300 ft (30 and 100 m) and that to prevent overfishing,
the fishing season should be closed during the spawning season. The ability of submersibles
to collect a variety of different kinds of samples using manipulator arms and to photograph
even in abyssal depths greatly expedites locating and exploring sunken vessels. The Harbor
Branch Oceanographic Institution currently has two manned submersibles, one of which is
certified to 3,000 ft (500 fathoms). In 1974 the Soviets had a large research submarine used
for fisheries research.


Unit Stock/Subpopulations
De Pontuel, H., and P. Prouzet. 1988. Numerical analysis of scale morphology to discriminate between
Atlantic salmon stocks. Aquat. Living Resour. 1:17-27.
Harden Jones, E R. 1986. Fish Migration. St. Martens Press, New York.
Marr, J. c. (ed.). 1957. Contributions to the study of subpopulations of fishes. The problem of defining
and recognizing subpopulations of fishes. U.S. Fish & Wildl. Serv., Spec. Sci. Rept. Fish. (208).
McCleave, J. D., G. P. Arnold, J. J. Dodson, and W. H. Neill (ed.). 1984. Mechanisms of migration in
fishes. NATO Conference Series IV: Marine Sciences. Vol. 14. Plenum, New York.
McDowall R. M. 1988. Diadromy in fishes. Migrations between freshwater and marine environments.
Timber Press, Portland, OR.
McKeown, B. A. 1985. Fish Migration. Timber Press, Beaverton, OR.
Royce, W. E 1984. Introduction to the practice of fisheries science. Academic Press, New York.
Rounsefell, G. A. 1975. Ecology, utilization and management of marine fisheries. C. V. Mosby Co., St
Smith, M. K. 1992. Regional differences in otolith morphology of the deep slope red snapper (£telis car-
bunculus). Can. J. Aquat. Sci.49:795-804.

Food and Feeding

Clepper, H. (ed.). 1979. Predator-prey systems in fisheries management. Sport Fish. Inst., Washington,
Cowey, C. B. , A. M. Mackie, and J. G. Bell. 1985. Nutrition and feeding in fish. Academic Press, Or-
lando, FL.
Grahame, J. 1987. Plankton and fisheries. Edward Arnold/Chapman Hall, New York.
Halver, J. E. 1989. Fish Nutrition. Academic Press, San Diego, CA.
Lagler, K. E, J. E. Bardach, and R. R. Miller. 1977. Ichthyology. John Wiley & Sons, Inc., New York.
Lovell, T. 1988. Nutrition and feeding of fish. Van Nostrand Reinhold, New York. 260 pp.
Royce, W. E 1984. Introduction to the practice of fisheries science. Academic Press, New York.
126 Part Two Fisheries Biology

Schooling Behavior of Fishes

Keenleysilde, M. H. A. 1975. Schooling behavior in fish. In Animal behavior in lab and field. E. O.
Price and A. W. Stokes (eds.). 35-38. W. H. Freeman, New York.
Moulton, J. M. 1963. Acoustic orientation of marine fishes and invertebrates. Ergeb. BioI. 26:27-39.
Moyle, P. B. and J. J. Cech, Jr. 1988. Fishes: An introduction to ichthyology. Prentice-Hall, Inc., Engle-
wood Cliffs, NJ. 2nd ed.
Shaw, E. 1970. Schooling in fishes: Critique and review. In R. L. Aronsen et al. (eds.). Development
and evolution of behavior. 453-480. W. Freeman, San Francisco.
Shaw, E. 1978. Schooling fishes. Am. Sci. 66:166-175.
Chapter 7

Population Size
and Fluctuations

To determine the effect of fishing on a stock, some estimate of the size of the stock that is be-
ing exploited is necessary. The weight or numbers of fish removed therefrom (fishing mor-
tality) can be obtained from landing statistics. The question then is what portion of the total
stock is not being harvested by the fishery? That is, how many are uncaught, or have es-
caped the fishery, remain to breed, and sustain the stock. Also, the remaining fish that sur-
vive predation and disease will grow to a larger size and thereby increase the biomass
available to fishermen in subsequent fishing seasons. The amount that can be taken without
harm to the fishery will be discussed in Chapter 12 and is generally called "surplus stock." It
is the amount that can be removed without harming the level of reproductive potential. At
an extremely low level of fishing, many fish are unavailable for use by humans and are con-
sidered "wasted"; alternately, at extremely high levels of exploitation, the reproductive poten-
tial of the population is greatly reduced and recruitment failures can occur. What is sought
in management plans is a suitable level of exploitation that is somewhere between the two
In addition to the numbers of fish known to have been removed from the population as re-
corded by fish sales, we need to know the size of that population. Where fish are aggregated
and visible it is possible to simply count them. Numbers of salmon moving up rivers and
streams to spawn can be counted through weirs when they enter freshwater or by counting
them on the spawning grounds on foot, by boats, or aircraft. Where there are numerous
streams in a spawning area, the total salmon spawning area is measured and counts are made
at important sample streams. The total numbers of spawners that have escaped fishermen and
reached the spawning grounds can be projected based on numbers in the samples.


In making population estimates occasionally the fish themselves cannot be seen; but, for ex-
ample, indicator organisms such as birds feeding on small prey fish driven to the surface by
tuna schools can be counted to estimate tuna abundance in an area. This is called "indirect
population estimation."

128 Part Two Fisheries Biology

Area Density
Population size of sessile species, such as oysters or clams, can be estimated by measuring
areas on a tidal flat, counting all individuals in each area, then expanding that portion of the
area the species covers. Bottom fish can be counted by the area density method in which a
trawl of known dimensions is dragged over the bottom for a certain time and at a certain
speed, thereby sampling a fixed area of the total fishing grounds. The total area of the fish-
ery is estimated before trawling begins. The numbers of various species in the resulting
trawl samples are totaled, then the size of the sampling area is multiplied by the fraction of
the samples of the total area to determine total population.

Correlated Populations
A technique called "correlated population" is used to estimate population size. It involves
estimating the total number of eggs and early larvae from spawning ground surveys that have
used fine mesh plankton nets to collect them (Chapter 4). If the average fecundity per female in
the spawning population and the sex ratio is known, it is possible to estimate the size of the
spawning population. This technique was used in the study of the Pacific sardine, Sardinops
sagax, in the California Cooperative Fisheries Investigation (CaICOFI) research program.

Population Size Estimation-Catch and Effort

Landings of fish or shellfish, and fishing effort required for these landings during certain
time periods (weeks, months, years), provides an index of population size. Because fisher-
men do not take all the fish from a population, estimation of population size by the catch and
effort method can serve as an index. Changing levels of fishing effort cause the size of land-
ings to fluctuate: more effort increases landings, less effort reduces landings. With constant
fishing pressure the landings will reflect the size of the total population.
Another method to determine population size, called the DeLury catch and effort tech-
nique (Fig. 7.1), will only work when the effort is constant and catch per effort is declining.
In this method, catch and effort data from a fishery must be obtained concurrently for short
time intervals (by days or weeks). Also, for each of these intervals, figures for the total land-
ing from the fishery are collected. A graph is made with accumulated catch on the abscissa
and catch per unit of effort on the ordinate. As fish are removed from the stock, the catch
per effort will decrease with time and the accumulated landings will increase. If little or no
immigration and emigration are taking place, or these rates are essentially balanced, the line
tracing the catch per unit of effort in be nearly linear and will tend downward. After suffi-
cient points are obtained, a best fitting line can be drawn through the points and extrapolated
down to the abscissa. An estimate of population size at the time the data were first collected
will be seen where the extrapolated line crosses the line of accumulated catches (Fig. 7.2). If
high rates of recruitment, emigration, or immigration are taking place during the period of
record gathering, the estimate of population size will be invalid because a straight line will
not result. The DeLury catch and effort technique can be modified by using a tagging exper-
iment that, if properly designed, will provide estimates of rates of immigration and emigra-
tion (Fig. 7.1).

Population Size Estimation-Tagging Method

One popular technique used to estimate population size of fish that cannot be estimated
directly by visual counts is the Petersen tagging method. It is very simple in principle, but
Chapter 7 Population Size and Fluctuations 129


o c o F E

Figure 7.1. Relationship between DeLury estimates of the whole population (above) and the marked
population (below). OE is the size of the apparent initial population. The slope of AE (the untagged
population) is affected by fishing, emigration, immigration, and natural mortality. The slope of line BC
(tagged population) is affected by the above changes to the population but is not affected by
immigration because no additional fish were tagged when the experiment was begun. See text for
additional details. Modified from Ricker (1975). Department of the Environment Fisheries and Marine
Service. Bull. 191.

in practice requires careful attention to detail. A subpopulation of fish is caught and tagged
that are in all respects the same as the untagged population except for the tag or mark. It is
extremely important to select a tag that best fits all the criteria for an "ideal" tag. The esti-
mate of population size is based on the equation:


where N = population size, M = the number of marked or tagged members in the popula-
tion, C = number of fish sampled from the population (caught by fishermen and examined
for tags), and R = number of tagged fish recaptured. Important assumptions are made when
using the tagging method for population estimation:
• no mortality is caused by the tagging procedure, tags will be retained by the fish, and
can be easily recognized;
• tags must not alter the behavior of the fish or make them more subject to capture;
• all tagged fish captured by fishermen will be returned;
• tagged fish will be randomly distributed, or over the fishing grounds, and/or fishing
is random.
Several Petersen population size estimates can be made during short periods of time. The
average of several size estimates thus obtained may be more accurate than a single estimate
made over a longer time. Also, statistical procedures giving upper and lower confidence in-
tervals for population size estimates around the population size are available.
130 Part Two Fisheries Biology

CCtl-622.8-0.002072 K(t)
600 C(t)=435.5-0.002806 K(t)

200 ---
--- --- --
o 1947 - 48 c(t)- 365.3-0.001795 Kef)

• ••
...:., .. --- --- --- ---

• ••• ••
en •
--- ---
u::: 200 1942 - 43·
--- --- --- ---
cet'): 541.7-0.001396 K(+)
Q. 0 1943 -44
600 • C(t)·492.5-00.001519 K(t)

--- -- ---
O~~.~~--~~~~--~~ 2 ----- ----
() •

--- 6 C(t)=389.8-0.0020M K(t)

~ 200 • 4

1944 - 45 C(t)- 589.6-0.001399 K(U

.... --- --
O~~~---~~---~~~ 200
I.l..I 1945- 46
I.l..I 600. • cCtl=U3.0-0.001150 O~~--~--~--~--~~------~--
1951 -52
•• • 400
--- ---
C(t):290.0-0.00t440 K(t)
•••• --- --- ---
50 100 ISO 50 100 150

Figure 7.2. Weekly accumulated catch of the sample fleet K(t) of the giant scallop, Placopecten
magellanicus, in the Digby area of the Bay of Fundy. From Dickie (1995). J. Fish. Res. Bd. Can. 12(6):
Chapter 7 Population Size and Fluctuations 131


Population Estimation-Aerial Surveys

Aircraft are frequently used to count individallarge organisms that swim near the surface,
such as porpoises, manatees, and whales. Fishes that form large schools, such as menhaden,
salmon, surface swimming tunas, and herring, are frequently counted by aircraft. Numbers
in these schools are estimated, then totaled for all schools seen. An estimated number of Pa-
cific salmon that return from the sea to freshwater lakes and streams to spawn can be
counted from the air to determine whether or not the seeding of spawning beds is suffi-
ciently heavy to provide enough young salmon for future runs. Aircraft have been helpful in
this connection, especially in Alaska when many salmon spawning grounds are not easily ac-
cessible on foot or in small boats.
The technique of population estimation used for herring in Canada is somewhat indirect.
Biologists in aircraft can locate areas of active herring spawning by clouds of white milt in
the water that last for a day or two. When spawning is over, concentrations of sea birds con-
tinue to mark the area. Accurate estimates of the miles of herring spawn can be translated
into estimates of the size of the spawning populations using fecundity per female and the ra-
tio of males to females. These figures, in tum, are useful indicators to determine the size of
the potential harvest that can be safely taken from this stock.

Population Estimation-Hydroacoustics
The echo sounder originally developed as a navigational aid and depth finder became use-
ful in detecting fish schools. Although the echo sounder was invented during the 1930s, and
has been widely used since by fishing industries, several decades passed before fishery biol-
ogists had sufficient background research to apply it to population estimation and manage-
ment of fish stocks.
There are serious flaws in most of the traditional methods of population estimation. Tag-
ging programs require considerable time in planning and executing and waiting for results to
accumulate before the population size can be estimated. The sales records from fishermen,
fish dealers, and processors used to calculate catch-per-unit-of-effort are costly to obtain and
involve a time lag. With echo sounders, obtaining population size data is much cheaper and
quicker than most other methods.
Echo sounders, as with any new electronic instrument designed for scientific use that
comes on the market, may have inherent limitations and disadvantages that might make it
unsuitable for certain research projects, even with the most recent innovations. A principle
concern when a researcher considers purchasing an echo sounder is the high price and cost
of installation on a vessel. Once installed, maintenance must be available when needed.
Electronic instruments do not perform as well as desired around saltwater. Ground-truthing
is required to be certain that species identification from the image on the screen is correct.
This requirement becomes less critical with experienced operators, but somewhat inaccurate
when working in habitats having species that might be confused with the target species.
Nonetheless, fish samples netted from targeted schools can provide species identification.
Assessing stock sizes of fish or shellfish near the surface like surface swimming tunas and
herring, or those very near or on the bottom, for example Penaeus shrimp, is usually imprac-
tical with echo sounders because these targets cannot be clearly detected. The target strength
is important to obtain accurate counts of fish schools. Echo sounders are manufactured with
frequencies and pulse lengths that vary to suit the needs of the particular fisheries.
132 Part Two Fisheries Biology


Fluctuations in abundance seem to lack clear patterns (Fig. 7.3); however, analysis of histori-
cal landing statistics for a few of the best known stocks (North Sea plaice, California sardine,
Pacific Halibut, Japanese sardine) has established four general categories according to natural
patterns of variation. Any classification of this type suffers from the very high overlap that
exists between categories. As pointed out above, living organisms evolve mechanisms that
apparently even out the effects of predators, competitors, disease, and the physical and
chemical environment. Numerous studies of land and aquatic animals demonstrate this rela-
tionship. Add to this the effects of varying levels of hunting or fishing pressure by man, and
the neat orderly pattern that may exist in a population of animals in nature can be obscured.
Four broad categories for stocks of exploited marine fish and shellfish have been suggested
by two FAO scientists, Drs. Caddy and Gulland, to assist in maximizing the sustained yield
from these fisheries.
Some stocks remain at about the same size, year in and year out, having variations of
about 20 to 30% of total stock size. A prime example is the North Sea turbot. This stable
stock phenomenon is simply called steady or predictable.
A second group, cyclical fisheries, shows cycles of high and low landings that can be
identified in regular repeated intervals. The category is illustrated in North America by the
California Dungeness crab and the Bay of Fundy scallop. Pink salmon in most areas in


Nlllion. pound.

110 _ _ _ _ _ _ _ _ _ _ _ _ _ _ 110

u.s. CAICH O. A!LANIIC COASI MACKE.llo 1900.65

MUlIon poIIndl

ISO _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 110

100 ___________________ 100

I... '910 IUO 1'4. Itl. •. . .

Figure 7.3. Fluctuations in catches of Atlantic coast mackerel, Scomber scombrus, United States, 1804-
1965. From Hoy and Clark (1967). U.s. Fish and Wildl. Servo Fish. Leaflet 603.
Chapter 7 Population Size and Fluctuations 133

Alaska run in larger numbers during one year and smaller runs during the next. Occasion-
ally the runs of successive years are approximately equal in numbers.
Significant changes in the landing from year to year may be caused by stock recruitment,
changes in the abundance of predatory species, and changes in the level of fishing (some-
times caused by changes in availability). Such irregular stocks include the Georges Bank
scallop and Norwegian juvenile herring.
Stocks in the last category are the hardest to manage. They are called spasmodic stocks,
and include Japanese sardines, California sardines and anchovies, and Gulf of Maine shrimp.
One approach to effectively fish these stocks is to increase landings in periods of high abun-
dance and tum to other species in periods of low abundance.
The above categories show that management plans offering the greatest chance of success
are reasonably those that apply classic theory to stocks having a steady pattern of abun-
dance. Once the pattern in a cyclical stock has been determined, fishery administrators and
scientists can estimate what point of the cycle they are in, then regulate fishing effort ac-
cordingly. In managing this cyclical stock, it is most important to be aware that when the
stock is approaching a peak, overfishing should be prohibited because heavy fishing pres-
sure would harm the stock as it declines to a natural trough. The last two categories, irreg-
ular and spasmodic stocks, are the hardest to manage even with a good background
knowledge of biology and ecology of the species, environmental parameters, and levels of
fishing pressure. This is true because of difficulty in sorting out the roles environmental
parameters and levels of fishing pressure play in reducing the stock size. In the past, regu-
lations were put into effect too late, after stocks had reached perilously low levels of abun-
dance. When levels reach this state, it complicates any understanding of reasons for
declines in stock size.
Karluk Lake (Kodiak Island, Alaska) pink salmon runs recorded as early as 1898 through
1944, when corrected for various physical and biological factors, showed that odd year re-
turns are significantly higher than the even years. These differences are quite consistent and
fairly constant. Fluctuations in the abundance of sockeye salmon in adjacent streams showed
survival from broods of odd-numbered years is very significant and consistently higher than
from broods of even-numbered years using records covering the years 1882 through 1944.
These cycles coincided with salmon runs in adjacent streams indicating that these fluctua-
tions have a common cause that was difficult to explain. Later sockeye salmon runs declined
and cyclical fluctuations became obscured. Earlier the wide variation in the number of
spawners resulted in wide oscillations in the numbers of young sockeye moving down river
to Karluk Lake and therefore available as food for predator fishes. Regular oscillations in the
supply of available prey may have acted as a control on the abundance of fish predators at
Karluk Lake where young juveniles reside before going to sea. These included young of
coho and king salmon, Dolly Varden chars, alpine chars, steelhead, and rainbow and fresh-
water sculpins.


Fish stocks are known to respond to the climatic conditions that can be measured: wind
speed and direction, air and sea temperatures, and current direction and velocities. How-
ever, stocks do not always show distinct changes in abundance in response to "obvious" var-
iables. In fact, they may be responding to other variables we cannot detect. Changes in
stock size may be the result of biological overfishing or climatic changes that affect successful
134 Part Two Fisheries Biology

recruitment. If climatic changes are severe prior to a drop in stock size, a correlation between
the two events is sometimes suggested. If, on the other hand, climate changes are too subtle
to be measured, or come from an undetectable variable(s), the decline in stock size may be
ascribed to overfishing.
A decade of warming ocean water between 1925 and 1935 was widespread along the U. S.
east coast and had dramatic effects on marine life. Southern living species appeared in
northern waters, for example. A more recent warming period that extended until 1950, with
peaks between 1945 and 1950, was apparently accompanied by stronger than normal ocean
and atmospheric circulation. Again, subtropical species moved or were carried by currents
outside their normal temperature (geographic) ranges. Because biological data suggest simi-
lar changes in distribution or abundance of animals across an ocean, approximately at the
same time, such as across the Atlantic Ocean, it seems fair to conclude that animals on both
sides of the ocean respond to widespread environmental and climatic changes.
North Sea stock abundance changes appear to be related to periods of warming and cool-
ing. Determination of relationships of stock size to climatic conditions is hindered when
heavy exploitation reduces herring and mackerel stocks; therefore the effects of climate are
less spectacular. Cold winters occur in the North Sea about once or twice in 20 years and are
evidenced by dead fish amassed offshore, obviously reducing the size of the population. In
nearshore waters, lagoons, and estuaries, the effects are much more visible as has been seen
in some areas of the southeastern United States.
The sharp decline of the once large fishery for the Peruvian anchoveta, Engraulis ringens,
caused by the El Nino current, is a recent and well-documented example of naturally caused
fluctuations and collapses. When normal along-shore currents, which upwell nutrient-rich
cold water to the surface and cause plankton blooms that feed the anchoveta, turn and blow
from offshore, the effects devastate anchoveta stocks and fishermen alike. When these stocks
became unavailable to fishermen, it was a severe economic blow to the anchoveta fishing in-
dustry although it may not harm anchoveta stock size or their ability to subsequently repro-
duce. El Nino also affects bird populations that feed on the anchoveta and similar species.
Christmas Island in the central equatorial pacific has a large breeding population of boobies
and other sea birds whose instinct to feed and protect their young is very strong, yet during
the 1982-1983 El Nino, parents abandoned nestlings because they could not find food for
them. Such behavior is indeed abnormal.
Processes affecting abundance of open sea fishes interact and include such perturbations
as rates of primary production (food for young) not synchronized with the hatching of a spe-
cies of fish, that is plankton blooms coming too soon or too late to serve as food for the
young. Also, the possibility is that with recruitment failures of a dominate species a compet-
ing species can replace a dominant species. This change in species abundance was suggested
earlier as a partial explanation of the decline of the Pacific sardine. Anchovies became abun-
dant when the competition for food was reduced by the decline of sardine stocks. The cli-
mate's influence on abundance of open sea fishes is imperfectly understood. Factors
affecting primary production in the sea, recruitment relationships, and the roles of predators
and competition, all need additional study by fishery biologists to adequately manage fisher-
ies (see Section on Chaos, Chapter 12).
In addition to widespread climatic conditions that affect fishing success, violent local
weather conditions can cause mass mortalities, as do hurricanes in the eastern United States
and Gulf of Mexico. Tsunamis and earthquakes also affect the abundance of fish and shell-
fish. Violent storms can also destroy fishing vessels and shoreside fish processing facilities
thereby reducing fishing effort for a time.
Chapter 7 Population Size and Fluctuations 135


Parasites and Diseases

Disease is a deranged condition of an organism that may be inherited, but it can also be
caused by dietary deficiencies, by physical and chemical factors in the environment, or by
parasites. In this section emphasis is placed on disease-producing parasites that are impor-
tant causes of mortality and income loss to fisheries. After a severe epizootic, American oys-
ter, Crassostrea virginicia, production in New Jersey declined about 65%, and 30 years later
still has not recovered.
What is a parasite? It is any organism that lives part or all of its life in or on another or-
ganism (called a host) and is dependent upon its host for metabolic requirements. Some par-
asites may do no more harm to the host than simply rob it of some of its food, but others can
be pathogenic. While some parasites do no harm to their hosts at certain times of the year,
they can cause considerable injury at other times. Sometimes a single parasite, or a species
by itself, may do no substantial harm, but in conjunction with other parasites of the same, or
other species, it may do great harm. It is difficult to determine whether a parasite is harmful
to a host and whether the harm is sufficiently severe to cause the animal to die (Fig. 7.4).
Fishery biologists need not be concerned about parasites that obtain food from the host if
their abundance is low, because usually the host can compensate by simply eating more food
if it is available. But he/she must be especially concerned and knowledgeable about para-
sites that directly or indirectly cause diseases that may contribute to mass mortality in wild
fish stocks.
Fish have become adapted to an environment of competition and encounters with large
predators and diseases. While not all parasites are disease producing and burden or ham-
per their hosts, anyone that does so makes the host lethargic and contributes to its early
death. This arrangement serves the needs of the parasite, for parasites must pass through
several hosts to complete their life cycle and reproduce. When parasites are in developmen-
tal stages requiring another host, it is to their advantage that the host fishes they are living
in be hampered in some way in their struggle for existence so that they will be eaten by
predatory fishes, birds, or mammals, and thereby pass into whichever organism is the nor-
mal host for the next developmental stage of the parasite. In this way, the parasites find
suitable requirements for growth and reproduction and thus are able to continue their life
cycle (Fig. 7.5).
How widespread are parasitic diseases in the sea? From the tiniest one-celled animal up to
giant whales, nearly every animal species has parasites of one kind or another. In some ani-
mals, parasites can be found inhabiting nearly every organ or surface area; some parasites are
microscopic and can live within the animals' cells. It is virtually impossible to find a marine
organism that does not harbor at least one parasite at some stage of its life history. The high
degree of parasitism is illustrated by one species of oyster and one of salmon. Approxi-
mately 30 parasites infect the American oyster, C. virginica, some place within its range; a to-
tal of 50 different parasite species parasitize the sockeye salmon, Oncorhynchus nerka, of the
eastern Pacific Ocean. As many as 45 species parasitize the mullet, Mugil spp., with fewer in
other geographic areas (Table 7.1). The true importance of parasites in the economy of the
sea cannot be appreciated because below the surface is a world strikingly inaccessible to di-
rect observation by humans. There parasitism is a common way of life, and because preda-
tory animals are known to more readily eat diseased animals, inroads by the combined
effects of predators and parasites into wild populations are continuous (Fig. 7.6).
136 Part Two Fisheries Biology


......,FIN RAYS
::::;"I:':~ 'U"OI
·.OTOIO ..
i~:~:':I! ".MATOOA .0110......
EYE "YD ••CU.Na DIGINIA _• .,••
co.... o ..

I I •

OPERCULUM • • I '.~<':'=::S:::58
NE • .,TODA I' I ' I





DIGINI .. I.,., ••
CESTODA la ....

Figure 7.4. Diagram of fishes showing sites of infestation by ectoparasites and infection by
endoparasites. From Fernando, Furtado, Gussev, Hanek, and Kakonge (1972). Methods for the study
of freshwater fish parasites. University of Waterloo, Waterloo, Ontario, Canada.

In recent years, rapid strides have been made in the study of parasites of commercial and
recreational fish stocks. It is becoming more apparent that the role of parasites in controlling
fish stock sizes is an important and imperfectly understood one.
Diseases are clearly important factors affecting losses of stock to fishermen, but in some ar-
eas, under certain environmental conditions, predators and competitors can be equally as im-
portant (Table 7.2).
Chapter 7 Population Size and Fluctuations 137



3rd stage larva

~ ::;::<
3rd stage larva

~ t
~ ~-.-

2nd stage larva

3rd stage larva

Figure 7.5. Life cycles of a parasitic roundworm (nematode) illustrating the variety of intermediate
hosts parasitized by a single species, Anisakis simplex. After Oshima (1972). Progress in medical
parasitology in Japan. Vol. 4, Meguro Par. Mus., Tokyo.

Figure 7.6. Myxosporean (protozoan) cysts in

fish flesh are not known to be harmful to
humans but as a group protozoans are believed
to play an important role in killing or disabling
their marine hosts. Photo courtesy W. M.
Stevens, University of Miami.
138 Part Two Fisheries Biology

Table 7.1 Numbers of Known Parasites from Mullets.

Eastern Northern Black Reports from
Parasites Mediterranean Red Sea Sea Mississippi Southeastern US
Blood protozoa 0 1 0 2 0
Gill protozoa 1 2 3 4 2
Microsporida 0 1 0 0 0
Myxosporida 2 4 2 3 0
Monogea 6 5 2 3 6
Digenea-adults 6 7 7 11 5
Metacercariae 12 2 2 6 3
Cestoda-larvae 2 0 1 3 0
Nematoda 4 0 1 3 4
Acanthocephalan 2 1 1 1 0
Copepoda 4 4 1 6 11
Branchiura 0 0 0 1 4
Isopoda 0 2 0 1 1
Hirudinea 0 0 0 1 1
Total 39 29 20 45 37
Parasite mix varies by areas. From Papema and Overstreet. In Oren (ed.) (1981). Aquaculture of grey mullets.
Reprinted with the permission of Cambridge University Press.


Predators are generally (but not always) larger than the animal on which they feed (Fig. 7.7).
They usually do not remain as closely associated with the prey animal as parasites (Fig. 7.8),
but move about until they find prey animals to attack and eat. Mortality rates of larval and
early stages of fish and shellfish are fantastically high. Only in recent years has the impor-
tance of the predator prey relationships in fisheries become understood. For example, seals
are estimated to eat almost half as many cod, Gadus morhua, salmon, Oncorhynchus sp., and
Salmo salar as commercial fishermen catch (Fig. 7.9). Concern for the inroads into fish stocks
made by predators has resulted in attempts to reduce the number of predators and hence
make more fish available to fishermen. This management technique, usually administered by
governmental agencies, frequently takes the form of a bounty program. These programs,
which have been notably unsuccessful, are discussed in the management section of this book.
Cannibalism, a special form of predation in which an animal eats its own kind, occurs
naturally in fish and shellfish populations. Shrimps and crabs become vulnerable to their
fellows when in the soft-shelled stage of their molting process. At this time other shrimps
or crabs of similar, or larger, size may very likely consume them. Cannibalism of young

.ll8llJLJIf...£_II.l;".IFIC COD ~.lJL.L""",,,,"" PREDATORS

SEA _~I~~~ ~__ ~~:I~~~_~~~~~~ ___________ _ Figure 7.7. Predators and competitors of
ALASKA PLAICE yellowfin sole, Limanda aspera. From
EELPOUTS COMPET !TORS Wilderbuer, Walters, and Bakkda (1992). Mar.
~ Fish. Rev. 54(4:)1-8.
Chapter 7 Population Size and Fluctuations 139

Table 7.2

American oyster (Crassostrea virginica)

Predators: Competitors:
Oyster drills Boring sponges
Urosalpinx cinerea Cliona spp. (seven specias)
Eupleura caudata Boring clams
Thais lamellosa Diplothyra smithii
T. haemostoma Mud wonns
Conchs Polydora websteri
Busycon contrarium P. ligni
B. canaliculatum Oyster crabs
Melongea corona Pinnotheres ostreum
Snails Mussels
Odostomia bisuturalis Mytilus edulis
o. impressa
Murex pomum Fouling organisms:
Starfish Slipper shell
Asterias forbesi Crepidula spp.
Flatwonns Tunicates
Stylochus ellipticus Molgula manhattensis
s. inimicus Sponges
Crabs Hydroids
Callinectes sapidus Bryzoans
Cancer irroratus Ascidians
Carcinides maenas Algae
Menippe mercenaria
Neopanopeus sp.
Black drum, Pogonias cromis
*From Iversen (1976). Farming the edge of the sea. Fishing News Books.

finfish by larger fish, including their parents, is widespread. Fish culturists have to separate
the parents and young of some species to prevent serious loss of their stock due to canni-
Competitors are organisms that may compete for space or food with another organism.
They can be of the same or different species. Competition for space is keenest among sessile
animals such as oysters, mussels, and clams because they cannot move from an area when it
becomes too crowded. In nature, animals with a high degree of mobility can spread out if
space is available to them. In some habitats, such as reefs, competition for limited space can
restrict the population size. Space competition is demonstrated also in some of the crabs,
such as the stone crab, Menippe mercenaria.

Red Tide
Red tide is a natural phenomenon that becomes visible when an unusually dense concen-
tration of microscopic plant like organisms color seawater reddish-brown (Fig. 7.10). Toxin
produced by these tiny cells causes massive fish and shellfish kills. Dead animals float to the
surface and eventually wash ashore to litter beaches and cause offensive odors.
140 Part Two Fisheries Biology

Figure 7.S. Starfish, a serious predator on

oyster beds, open the oyster and devour the
meat. Methods to reduce the populations of
starfish and other oyster predators are
described in Chapter 13. Photo courtesy of
Fisheries Research Board of Canada.

Figure 7.9. Sea lions are important predators

on Alaska salmon. Here one eats a salmon
caught in a fishermen's net. Photo courtesy T.
R. Merrell, Jr.

Red tides caused by the accumulation of

small marine algae called dinoflagellates occur
worldwide. In Florida, the red tide organism,
Ptychodiscus brevis, is only about one-thou-
sandth of an inch in diameter. Red tides usu-
ally appear form September through February
about 10 to 40 mi (16 to 64 km) offshore. Areas
of the ocean as large as 14,000 mi 2 (36,260 km 2)
may be affected by red tides. Poisonous, dis-
colored water may be carried inshore by cur-
rents and winds.
Slow-moving, bottom-dwelling fish are usu-
Figure 7.10. Sketch of a dinoflagllate
ally the first to die during a red tide. Death re- protozoan, Ptychodiscus brevis, the species that
sults from either a nerve toxin that inhibits causes red tide off the southwest coast of
respiration, or a blood toxin that interferes with Florida. From Torpey and Ingle (1966). State
normal oxygen transport within the body. The of Florida Board of Conservation Educational
extent of the kill depends on the density of the Series No 1. 27 pp.
Chapter 7 Population Size and Fluctuations 141

bloom and length of time fish and shellfish are exposed to the toxin. Shellfish seem to be as
severely affected by red tides as are finfish. Fish killed during red tides are usually not suit-
able for human consumption. Large kills can substantially reduce the size of a fish stock in
a local area.



Critical measurements in any fishery are how long an exploited fish species lives, and at
what rate it dies. As was indicated earlier, management schemes are quite different between
short-lived and long-lived species of fish and shellfish. The total mortality rate (natural and
fishing) for a certain level of fishing of a long-lived species is an indication of how long it
will be in the fishery.
Survival rates can be determined by using "catch curves" that are constructed by plotting
age of a fish species against the number of fish of each age in the catch of a trawl (Fig. 7.11).
Here we are not using cohorts (fish from a dominant year class), but numbers from different
year classes. Then, from a trawl haul of a fish species, the smallest sizes the trawl will retain
up to the largest present obtained. The fish are aged and numbers of each age counted.
When the frequency of each age is plotted against age on an arithmetic scale, a curve will re-
sult with a short ascending line on the left side, and on the right side a ski slope shaped line
will result. Converting these numbers to natural logs will generally straighten both of the
lines. The left line is of little interest because it shows numbers of fishes that are not fully re-
cruited into the fishery; because of their small size many escape through the meshes of the

sAs 925 1025 1225

Figure 7.11. Catch curve for the Pacific halibut population (southern grounds). Samples taken from
tagging in 1925 and 1926. Abscissa is mean length of successive 5 em length groups, in millimeters.
Ordinate logarithm of the numbers of fish taken at each length interval. From Ricker (1948). Indiana
University Publications. Science Series No. 15.
142 Part Two Fisheries Biology

net. The rate of decrease in the number of fish from one year to the next after the fish are
fully recruited is the total annual mortality rate.
Fishery biologists have long used catch curves to describe total mortality (natural and fish-
ing mortality) for many species of fish in both exploited and unexploited populations. Catch
curves generally show what fish are dying in the population in proportion to the numbers
present in each year class because a natural log plot of abundance of each group on age is
linear. This relationship holds despite variation in the size of recruitment from year to year
and means that natural mortality is approximately constant.
To use the catch curve method to estimate total mortaliy for a species, certain conditions
must exist:

1. there are no large changes in survival rate of fish with age;

2. fishing pressure is the same over all ages used in the analysis;
3. the trawl samples all age groups randomly;
4. the level of recruitment is approximately equal for each age group.

Predictably, when the level of fishing pressure on the stocks increases, the line of a catch
curve will drop faster than when fishing pressure is low. This means that few older large fish
remain in the fishery because they have been previously removed by fishing.
Total mortality also can be measured by estimating population size visually or by tagging
means, at one time, and again after a period of time. Reduction in the size of the stock is due
to total mortality if rates of immigration and emigration are low or absent.
Virtual population analysis employs data on the number of fish of a particular age (co-
horts) that are in the catch and fishing effort data. In contrast the parent-progeny relation-
ship (Chapter 4) is based on the number of recruits at some time after data on the catch of
adults is obtained.
Cohort analysis can also be used to measure total mortality. This means a cohort (a group
of fish spawned in the same year) can be followed through the fishery estimating the size of
the population during each year. The Jackson equation is suitable to calculate survival:

N4 +Ns +N6 +N7 +Ns

S = ---------
N3 +N4 +Ns +N6 +N7

where 5 = survival and N = the population size with subscripts used for the age of the fish,
hence, N6 is the size of the population of 6-year-old fish.
For example, starting with a population of 25 fish that are first recruited at age 3, the re-
duction would go like this and the annual total mortality rate of 0.5 is obtained (see below):

12 +6 +3 +2 +1
S= = 0.5.
25 + 12 + 6 + 3 + 2

When a dominant year class (cohorts) passes through the fishery, it is easy to follow and to
estimate the numbers of fish remaining as the years pass. (Chapter 5, Fig. 5.13).
Population of tagged fish recaptured over short periods of time is another way to estimate
total mortality. The short periods can be extrapolated to monthly or yearly mortality rates.
Up to this point we have been concerned with estimating total mortality. However, it is
important to go beyond this to find the role that each kind of mortality plays in reducing the
Chapter 7 Population Size and Fluctuations 143

size of the population. If fishing mortality is high and natural mortality is low in a declining
fish population, some way to reduce fishing pressure should be sought. If, on the other
hand, natural mortality is high because of a degradation of the environment, and fishing
mortality is low, effort should be focused on reducing the level of natural mortality to arrest
the decline of the stock. Generally fishing effort is high and by reducing it over time stocks
can be restored to productive levels.
Other ways to separate natural and fishing mortality may have more theoretical than prac-
tical value:

1. First, measure the mortality in a virgin population, then measure it later when the stock is
2. When fishing stops on a exploited stock for a period of time, the stock size will increase be-
cause only natural mortality is reducing it. During World War II fishing stopped for 6 years on
the 1932-1933 year class of North Sea plaice. Estimates of abundance were made before and af-
ter fishing stopped.
3. Plotting several different levels of fishing effort against total mortality (the rate of natural and
fishing mortality) results in an upward-trending straight line that, when carried back to the
point of no fishing effort, allows natural mortality to be read from the ordinate of the graph.
(Fig. 7.12). Catch curves with two different levels of fishing. The slope differences of the two
lines is an estimate of total mortality.
4. Using recapture data of tagged fish, formulas are available to separate fishing mortality from
"other losses." When emigration is very low other losses will be largely natural mortality
(Fig. 7.13).




...a::~ 1.00
j! .!SO

o 50 100 150 200 250


Figure 7.12. Instantaneous total mortality and fishing effort as a method to separate natural and
fishing mortality. From Widrig (1954). U.S. Fish and Wildl. Serv., Fish. Bull. 54:141-166.
144 Part Two Fisheries Biology

LOGe Y-4.9511- .1396 X
• •
I&J •



o~ _________ _________ _________ ______

~ ~ ~ ~~

I 5 10 15 20

Figure 7.13. Separating natural and fishing mortality using tagged spotted seatrout, Cynoscion
nebulosus. From Iversen and Moffett (1962). Trans. Am. Fish. Soc. 91(4):395-398.


Population Size Estimates (Fish Stock Assessment)

Clark, S. H., W. J. Overholtz, and R. C. Hennemuth. 1982. Review and assessment of the Georges Bank
and Gulf of Maine haddock fishery. J. Northwest Atl. Fish. Sci. 3(1):1-27.
DeLury, D. B. 1947. On the estimation of biological populations. Biometrics 3(4):145-167.
DeLury, D. B. 1951. On the planning of experiments for the estimation of fish populations. J. Fish.
Res. Bd. Can. 8(4):281-307.
Dickie, L. M. 1955. Fluctuations in abundance of the giant scallop, Placopecten magellanicus, (Gmelin) in
the Digby area of the Bay of Fundy. J. Fish. Res. Bd. Can. 12(6):797-857.
Evans. G. T. 1980. Remote sensing as a technique for synoptic inventories of fisheries related resources.
In Estuarine perspectives. Proc. Fifth Biennial Int. Est. Res. Conf. Oct. 7-12, 1979. V. Kennedy (ed).
Gulland, J. A. 1983. Fish stock assessment. A manual of basic methods. John Wiley & Sons, Inc. Som-
erset, NJ.
Hoag, S. H., G. H. Williams, R. J. Myhre, and I. R. McGregor. 1980. Halibut assessment data: Setline sur-
veys in the North Pacific Ocean. 1963-1966 and 1976-1979. Research Report IPHC No. 18. 1-42.
Holden, M. J. and D. F. S. Raitt. 1974. Manual of fisheries science. Part 2. Methods of resource investi-
gation and their application. FAa Fish. Tech. Pap. No. 115.
Jamieson, G. S., N. B. Witherspoon, and M. J. Lundy. 1981. Assessment of Northumberland Strait scallop
stocks, 1980. Can. Tech. Rep. Fish. Aq. Sci. No. 1017. .
Chapter 7 Population Size and Fluctuations 145

Ketchen, K. S. 1953. The use of catch-effort and tagging data in estimating a flatfish population. J. Fish.
Res. Bd. Can. 10(8):459-485.
Sissenwine, M. P. 1981. An overview of some methods of fish stock assessment. Fisheries 6(6):31-35.
Williams, K. 1981. Aerial survey of pelagic fish resources off southeastern Australia. 1973-77. Rep.
Div. Fish. Oceanogr. CSIRO No. 130.

Stock Size Fluctuations (Population Dynamics)

Beverton, R. J. H. and S. J. Holt. 1957. On the dynamics of exploited fish populations. Minist. of Agric.,
Fish., and Food. Fish. Invest. Ser. II, Vol. XIX.
Caddy, J. F. and J. A. Gulland. 1983. Historical patterns of fish stocks. Marine Policy. October
Cushing, D. H. 1982. Climate and fisheries. Academic Press, San Diego, CA.
Gulland, J. A. 1972. Population dynamics of world fisheries. University of Washington Sea Grant Publ.
WSG 72-1.
Gulland, J. A. 1983. Fish stock assessment: A manual of basic methods. John Wiley & Sons, Inc., New
Gulland, J. A. (ed.). 1988. Fish population dynamics: The implications for management. John Wiley &
Sons, Inc., London and New York. Second Edition.
Laevastu, T. 1993. Marine climate, weather and fisheries. John Wiley & Sons, Inc. Somerset, NJ.
Laevastu, T. and F. Favorite. 1989. Fishing and stock fluctuations. Fishing News Books, Ltd., Farnham,
Surrey, England.
Rothschild, B. J. 1986. Dynamics of marine fish populations. Harvard University Press, Cambridge,
MA and London.
Sherman, K. and L. M. Alexander (ed.). 1986. Variability and management of large marine ecosystems.
AAAS Selected Symposium Series 99.
Sinclair, M. 1988. Marine populations: An essay on population regulation and speciation. University
of Washington Press, Seattle.

Survival (Mortality)
Clepper, H. (ed.). 1979. Predator-prey systems in fisheries management. International Symposium on
Predator-Prey Systems in Fish Communities and Their Role in Fisheries Management. Atlanta, GA,
July 24-27.
Ellis, A. 1958. Fish and shellfish pathology. Academic Press, Orlando, FL.
Everhart, W. H. and W. D. Youngs. 1982. Principles of fishery science. Cornell University Press, Ithaca,
NY. (Second Edition).
Kerfoot, W. C. and A. Sih (eds.). 1987. Predation: Direct and indirect impacts on aquatic communities.
University Press of New England, Hanover and London.
Kinne, 0. (ed.). 1980-1985. Diseases of marine animals. Vols. 1-4 (Protozoa through Mammalia). Bio-
logische Anstalt Helgoland, Hamburg.
Kinne, O. and H. P. Bulnheim (eds.). 1984. Diseases of marine organisms. International Helgoland
Symposium 1983. Helgolander Meeresuntersuchungen Vol. 37 (Nos. 1-4). .
Royce, W. F. 1984. Introduction to the practice of fisheries science. Academic Press, New York.
Sindermann, C. J. 1990. Principal diseases of marine fish and shellfish. Second Edition. Volume 1 Dis-
eases of marine fish, Volume 2 Diseases of shellfish. Academic Press, New York.

Gear, Methods, and Landings
Chapter 8

Fishing Vessels, Gear,

and Methods

Fishermen throughout history have known about fluctuations in abundance of the stocks
they sought. Times of poor fishing were explained by superstitions and dogma because few
scientific facts were available. Statements suggesting that poor catches were evidence of
God's wrath for abusing "His gifts to man" were common. Fishermen even blamed the con-
struction of lighthouses for poor catches. At the beginning, of course, fishing pressure was
so low that its effect on large stocks of fish was undoubtedly minimal.
Early commercial fishermen were seriously handicapped because their sailing vessels were
reliant on favorable winds. Portuguese fishermen sailed to the Grand Banks in schooners to
fish cod. Once on the fishing grounds, individual fishermen used small sailing dories to
move about the grounds to and from the mother ship. Hand lining was the method of fish-
ing. Today, small sailing boats, still used in developing countries, are practical for the short
runs to and from the fishing grounds.
As the numbers of fishermen-and the sizes of their vessels-increased, the important
question of the effect of fishing on the stocks of fish began to nag. With competition, vessels
operated farther from home ports. Fears of depletion of stocks of fish were aroused, plus
questions whether these virgin stocks could withstand the tremendous drain.
Although more recent, the history of fishing methods and the development of fishing gear
in the United States essentially parallels that of other fishing nations (Table 8.1). Over 300
years ago in the United States, fishermen used primitive gear types close to shore because
productivity of these waters provided sufficiently high yields to afford fishermen a comfort-
able living. Using small skiffs and hand lines, rakes, or beach seines near estuaries and on
tidal flats, the market demand for seafood was satisfied. As America grew, both the demand
for seafood and competition for available seafood increased to a point where fishermen were
forced to go offshore to make reasonable catches.
Each new development in the fishing industry brought another. As fishing went farther
afield, more effective and larger, heavier fishing gear was designed and used, including gill
nets, longlines, traps, and purse seines. Offshore fishing with the new and larger gear re-
quired larger, and more expensive fishing vessels (Table 8.2). Vessel propulsion changed
from sails to steam engines, and finally to diesels that reduced running time to and from the
fishing grounds; and, these more efficient, powerful engines, enabled use of large nets like ot-
ter trawls that came into use about the turn of the century. The heavy gear eventually
brought about the invention of such deck gear as power blocks to retrieve and haul nets
heavy with fish from the sea into the boat's holds.

150 Part Three Fisheries

Table 8.1 Historic Developments in Fishing Gear and Methods

Year" Development

1825 Long-range beam trawlers increased in size, numbers, and length of trips
1850 Steam power introduced. Longer range and greater towing power permitting use of
more efficient gear types, e.g., newly introduced otter trawls
1920 Introduction of diesel and gas engine power permitted use of still more efficient gear,
plus ever longer fishing trips
1920 Locating fish schools by aircraft
1930 Synthetic fibers for fish nets and lines
1930 Use of freezers aboard vessels to preserve catches and improved processing
1940 Electronic gear for navigation and fish school location
1954 Mother ship (factory trawler operations) began
1955 Conversion from single net fishing to double-rigged shrimp trawls in southern waters
1958 Power blocks for salmon, menhaden, and tuna seining
1959 U.S. Pacific tuna fleet converts from live bait fish to purse seining for surface-
swimming tunas
1959 Escalator oyster dredge
1980 Searching for tuna schools by satellite
1980-1983 High diesel fuel prices caused concern among fishermen, feasibility of sail-assisted
fishing vessels was investigated
"Dates were obtained from a variety of sources and are approximate. Furthennore, new, technology may require
several years for acceptance by a majority of some fishing fleets.

Table 8.2 World Fishery and Fishing Support Fleets, by Type of Vessel, Number, and Gross
Registered Tonnage: For Vessels over 50O-GRT, 1975, 1980, and 1985-1992.
World fishing fleets. An analysis of distant-water fleet operations.
Vol. 1 National Marines Fisheries Service

Fishing Vessels Support Vessels Total High-seas Fleet

Year Number Tonnage Number Tonnage Number Tonnage

1975 18,217 7,830,244 723 3,508,374 18,940 11,338,618

1980 20,671 9,195,225 852 3,473,892 21,523 12,669,117
1985 21,251 9,446,935 852 3,538,451 22,103 12,985,386
1986 20,974 9,521,831 865 3,852,767 21,839 13,374,598
1987 21,267 9,666,065 875 3,831,468 22,142 13,497,533
1988 21,827 9,960,566 879 3,851,775 22,706 13,812,341
1989 22,149 10,139,102 881 3,984,871 23,030 14,123,973
1990 23,132 10,764,053 989 4,135,738 24,121 14,899,791
1991 23,581 11,069,085 1,032 4,151,741 24,613 15,220,826
1992 23,718 11,146,416 702 2,087,823 24,420 13,234,239
Sources: Lloyd's Register of Shipping. Statistical Tables. London, various years. The data are
completed as of June of each year, except for 1992 which is effective through December 31,
1992. World Fleet Statistics (as of 31 December 1992)., London, 1993; page 25.
Chapter 8 Fishing Vessels, Gear, and Methods 151

Large schools of fish are necessary for successful purse seining. To increase their success
fishermen began to use aircraft to locate large schools of migrating fish. The otter trawl, de-
veloped in New England and later adopted in Gulf of Mexico and southern Atlantic shrimp
fishermen, widened their fishing range. It replaced traditional shrimp haul seines (beach
seines) that could only be used in the shallow nearshore waters. Offshore, otter trawls could
make bigger catches of larger sized adult shrimp.
After World War II, electronic equipment for navigation, communication and depth indica-
tors and recorders, and automatic pilot devices developed for war use became available for
fishing vessels. What was formerly done by hand now had mechanical help, and fishing be-
came more efficient.
At about the same time, another improvement in fishing gear came with the introduction
of synthetic fibers for fishing lines and nets. While natural fibers are satisfactory for fishing,
they cannot match the greater strength, longer life, and lighter weight of synthetic fibers.
Even though present day fishing gear is expensive and has been modified in many ways,
much of it is remarkably similar to what fishermen used at the dawn of history. But this
more efficient gear, larger fleets of bigger vessels, and greater market demand created prob-
lems in biological and economic overfishing of fish stocks around the world. The situation
worsened in local fisheries with the influx from time to time of heavily government subsi-
dized foreign fishing vessels from many other countries. This will be discussed in subse-
quent chapters.
Some typical signs of biological overfishing are:

1. Catches drop with the same or increased fishing effort. An index such as numbers or weight of
fish caught per some standard unit and time period (boat day, boat week, etc.) is used to docu-
ment these changes in a fishery. These measures are usually called "catch per unit effort"
(CPUE) or simply "catch per effort" (C/E). With constant effort in the fishery, the total landings
per month, or year, decrease.
2. Fishermen work harder and sail farther to compensate for reduced catches in nearshore stocks.
3. Average size and age of fish in an exploited stock decrease. When the larger, older fish in a
stock are removed the younger stock cannot replace them because the heavy fishing pressure
removes small fish before they can grow to a large size.

Overfishing will be discussed in more detail in Chapters 12-14 on fisheries management.


Several general stages can be identified in the history of marine fisheries. The first of these
is the use of small sail-powered vessels to fish for food. As vessel sizes increased and gear
technology improved, so came a strong demand for fish as human food and fish meal for
farm animals. In countries with strong economies where there is a class of people who have
leisure time, a second kind of fishery developed: recreational fishing. Large recreational fish-
eries have developed primarily in tropical or subtropical areas such as Bahamas, Gulf of
Mexico, both coasts of Panama, the west coast of Mexico, the north coast of Australia, both
coasts of the United States, and Hawaii. Most recreational fisheries developed rapidly after
air transportation and tourism became popular. Countries having popular recreational fish-
eries do not necessarily have a large wealthy class of people, but the game fish attract fisher-
men from elsewhere. In the chapters on recreational fishes and commercial fishing, I discuss
152 Part Three Fisheries

the problems of allocation of resources and added fishing pressure on stocks of fish harvested
by both commercial and recreational fishermen. Culture fisheries (aquaculture or sea farm-
ing), well-established in developing countries, have became popular in developed countries
recently as a result of the high cost of capture fishing (Chapter 11).

Sail-Assisted Commercial Fishing Vessels

Because of high diesel fuel costs, and the fuel shortage at the time, commercial fishing ves-
sel owners in the United States sought ways to trim fuel expenses. In the early 1980s, the In-
ternational Conference on Sail-Assisted Commercial Fishing Vessels produced numerous
published articles on the subject. The return to sailing vessels was also suggested for ocean-
going cargo vessels and oil tankers to reduce fuel costs.
Sail trawling is a small operation that began on Lakes Kitaura and Kasumigaura in central
Japan about 1877 to catch whitebait and pond smelt. Narrow wooden vessels, which drift
sideways during the fishing operation, have booms extending from bow and stern that in-
crease the width of the net opening. Small otter boards fixed to the net wings further in-
crease the width. Fishing by sail there is possible only because the predominant wind blows
parallel to the long axis of the lake with sufficient strength and frequency during the fishing
season to justify the effort expended. To begin fishing on days when winds are favorable the
vessels run upwind under diesel power.
The possibility of sail trawling for shrimp in the southeastern United States is somewhat
remote because of adverse wind conditions, and there are few locations where it might be
carried out. Sailing assistance can be used more practically to go to and from fishing
grounds than for the actual fishing operation, because with anything larger than small trawls,
dredges, or light trolling gear, high horsepower engines are needed to pull the gear fast
enough to capture, or to attract the attention (trolling) of active swimming species (Fig. 8.1).

Figure 8.1. Baiting hooks on long line gear on

deck of a Gloucester haddock sailing schooner
"Mystic." From Goode (1887). U.S.
Commission of Fish and Fisheries. U.S.
Government Printing, Office, Washington, D.C.

The economics of sail-assisted fishing vessels is important to consider in evaluating this

method of propulsion. The purchase cost of a sail-assisted vessel, or retrofitting an old ves-
sel, is generally much higher than a motor vessel; therefore, if fuel costs are a relatively small
component of total operating costs, then such vessels are not cost efficient. However, if fuel
figures are about 30 to 50% of a total operating cost, then sail-assisted vessels could be a cost-
Chapter 8 Fishing Vessels, Gear, and Methods 153

efficient investment. In 1970 the average annual fuel bill for a 65 ft (20 m) shrimp boat was
$13,000. By 1980 it climbed to $70,000. Later with the reduction in the price of diesel fuel
and all the disadvantages and limitations of sail-assisted fishing, sail trawling as an alterna-
tive was abandoned.
In addition to commercial fishing by sailing vessels in an effort to reduce fuel costs, these
vessels are mandatory in some fisheries to reduce or prevent overfishing. Years ago Bristol
Bay, Alaska fishermen used what was basically a 28 ft (8.5 m) boat with a small sail to fish
for red salmon (Oncorhynchus nerka). A "monkey boat" powered by a small engine towed the
fishermen to and from the grounds. During fishing, sails were erected on the boats for move-
ment about the grounds. In 1950, this practice stopped because fishermen were lost during
sudden heavy storms. "Skipjack" sailing vessels are still in use today in the Chesapeake Bay
oyster fishery where they pull small dredges. At its peak, over 1,500 skipjacks were in used
in this fishery. A 1991 estimate places about 20 skipjacks still in use.


Vessels are not always necessary for certain commercial, artisan, and recreation fisheries, es-
pecially the nearshore shallow water species. The behavior and life history of many of these
species permits collection without specialized fishing gear. Some species of macroalgae are
collected along beaches by hand or rakes, such as Irish moss that is used for, among other
things, extraction of chemicals for food additives (Fig. 8.2).

Figure 8.2. A blood worm digger working the

intertidal zone in Maine. Photo courtesy of
Ivan FIye.
154 Part Three Fisheries

Invertebrate species that can be harvested without gear include species that attach to the
substrate, such as oysters, mussels, and sponges. Unattached species, capable of only limited
slow movement are easily caught by hand. Examples include: sea urchins, sea cucumbers,
top shells, abalone, and queen conch.
Vertebrate species often caught by hand or with primitive instruments are those that move
from the sea into freshwater streams to spawn; anadromous species like salmon, shad, and
eels are examples. Species that emerge from the sea onto beaches or rocky shores to repro-
duce include sea turtles and grunion (Leuresthes tenuis). Seals come out and form harems on
rocky shores and bear their young.


Boats, vessels, and ships are not interchangeable names. Boats are defined as commercial
fishing craft not powered by motor, for example rowboats or sailboats, and have a capacity
of less than 5 net tons. Commercial fishing vessels are crafts having a capacity of 5 net tons
or more. In the United States, these crafts are either enrolled or documented by the U.S.
Coast Guard and given an official number. Ships may be defined as any vessel of consider-
able size that is motorized and is capable of navigating deep waters.
While most fishing vessels have conventional hulls, they are modified in many different
ways depending on the kind of commercial fishing they are designed for. Large offshore ves-
sels include those designed for trawling, seining, gill netting, hook and lining, and miscellane-
ous uses. Within these groups are vessels of nonstandard design, but which fit the needs of
particular fisheries in different parts of the world where different species are fished in varying
sea conditions. In the discussion of fishing gear and methods that follows, many of these differ-
ent vessels are discussed and illustrated. Major differences include, size of vessel (offshore ver-
sus nearshore vessels), placement of the wheelhouse, height of wheelhouse and/or crow's
nests, amount and location of work space and heavy deck machinery, that is, winches, line
haulers, gallows and gantries, catch storage areas, etc. Estimates of the numbers of fishing ves-
sels worldwide are very difficult to obtain. In 1991, there were 85,800 fishing vessels and boats
in the United States, excluding Hawaii and the Great Lakes states (Fig. 8.3).


.J!l 600
~ 600 I'
.0 400
E 200
1975 1980 1985 1990 92
IGNUmber of vessels -Tonnage I
Figure 8.3. West European high-seas fishing fleet, by number of vessels and tonnage, 1975-1992. From
Beaudry and Folsom (1993). NOAA Tech. Memo. NMFS-F/SPO-9.
Chapter 8 Fishing Vessels, Gear, and Methods 155

Characteristics of Major Types of Fishing Vessels

Some general characteristics identify the larger fishing vessels designed for a particular
fishery. In most cases, the wheelhouses and trawler superstructure are located either near the
bow or stern to allow working space for handling gear and catches. In older live bait tuna
vessels the live bait holding tank and the racks where fishermen stand while fishing easily
identified them. Longline vessels fishing for deep-swimming tuna and ground line gear for
halibut could usually be identified as Japanese style line haulers (Fig. 8.4). Halibut vessels
characteristically have a stern chute used to payout (set out) their baited ground line. Power
blocks and seine tables are installed on vessels for purse seining salmon, menhaden, and

Figure 8.4. Fishermen setting longline using a

chute typical of halibut vessels. Photo by G. 1.
Murphy, u.s. Fish and Wildlife Service.

Otter trawlers, sometimes called side trawlers, have, the wheel house astern and the side
of vessel where the trawl is retrieved is reenforced to prevent damage to the hull by the otter
boards when the catch is hauled aboard. Shrimp trawlers usually have "gallows" to pull the
boards out of the water when the trawl is being retrieved.
Until the oil crisis of the late 1970s, the trend was toward larger vessels and more sophis-
ticated equipment. For many years, trawlers were of wood construction, but as early as the
1940s and since World War II, the number of steel boats increased. The proportion of steel
boats in the fleet continues to increase for several reasons: steel construction does not require
the highly skilled labor necessary for building wooden trawlers, and the life expectancy of
steel vessels is about double that of wooden ones. This greatly offsets the 25% or so higher
construction costs for steel vessels; also, repair, maintenance, and hazard insurance are
cheaper for steel vessels. Other materials for vessel construction include aluminum, ferro-
concrete, reinforced plastic and molded fiberglass hulls (up to 60 ft or 18 m).
A major innovation in trawler design made its appearance during the early 1960s, namely
stern trawlers (Fig. 8.5). Prior to the introduction of stern trawling, side trawling was the ac-
cepted method. However, more fishermen are needed in side trawling because setting the
trawl over the side of the vessel and retrieving it, loaded, with the vessel standing beam onto
the seas with a heavy load of fish is dangerous, especially in heavy seas, and fishermen can
lose their lives. Some stern trawlers haul the gear up and over the stern using a swinging
156 Part Three Fisheries

Figure B.S. British distant water stern trawler

with full freezing capacity. Photo courtesy of
Fishing News Books.

gantry. Others have a chute cut in the stern that extends from the water line to the working
deck. Stern trawling with a chute allows the gear to be set down the chute and retrieved up
the chute onto the deck. Whale factory vessels have found stern ramp vessels very efficient
for their uses.
Many large stern trawlers are called factory trawlers because they process the fish on
board. Worldwide use is made of the factory trawler concept. Stern trawlers are easily iden-
tified because the wheelhouse and crew's quarters are located arnidship to allow ample room
on the afterdeck for the fishing operation. The gallows and gantry on larger stern trawlers
have a trawl bridge located aft to expedite the trawling operation. Western Germany, Great
Britain, Spain, Russia, United States, and Japan are among the countries that use factory
trawlers (Fig. 8.6). Many are about 280 ft (85.3 m) in length with a displacement of about
3,700 tons. Fish are processed on board, with edible fish parts packaged and frozen and
waste converted into fish meal and oil.
The British developed the factory trawler concept late in 1940 when they refitted a mine-
sweeper. In 1954, they designed and built one. The Russians saw the advantages, borrowed
the plans, constructed vessels, and put them into use as fast as they were built.
Two small, 270 ft (82.3 m) long, stern trawlers were built for the United States in 1968, at
a cost of $5 million each, to compete with foreign vessels. Four years later they were both
laid up because inefficiency of operation ate the profits. Other reasons included difficulty in
keeping qualified crews, and labor union regulations. The 200-mile limit has made factory

(a) (b)

~ ~
Figure B.6. Examples of Soviet stern trawlers
and a factory ship that fished off the west coast
of the United States in the 1960s. Modified
from Hintz (1970). U.S. Fish and Wildlife
Service, Bureau of Commercial Fisheries.
Circular 332. pp. 53-75.
Chapter 8 Fishing Vessels, Gear, and Methods 157

trawlers obsolete because they could no longer compete in markets because of their high op-
erating costs.
In distant water fisheries, ships that provide services and supplies to catcher boats are col-
lectively termed "support ships" or "mother ships" (Fig. 8.7). Fish factory ships may be as
large as 600 ft (183 m) in length, but more often are smaller, about 370 ft (112.8 m) long.
These ships have hospital and recreational facilities for their own crew and for the fishermen
on catcher boats. In addition, tankers and transports have been used to deliver supplies in
these large fisheries where ships can be away for several years at a time.
Trolling vessels are generally small. They can be easily identified by long trolling poles
that are held upright when the vessels are running to or from the ground and lowered when
fishing. Vessels in search fisheries usually have a crow's nest and a high wheel house for lo-
cating schools of fish.
Vessels that must bring their catch to the canneries alive frequently have a large portion of
their holds converted to holding tanks with circulating seawater. Crabs can be maintained in
this way for up to about 3 weeks. Larger vessels, some up to 120 ft (36 m) long to operate
in heavy seas, carry the weight of the tanks and deliver large traps to and from the fishing

Trawlers and
aUKlllary craft

. .
Fish carriers
r-- Processing
ship ~

I t
Figure 8.7. Organization of Soviet offshore
+Fleet's bose
fishing operations in 1974. From Sealy (1974). port
Mar. Fish. Rev. 36(8):5-22.
158 Part Three Fisheries

grounds. They also relocate traps on the fishing grounds. Another characteristic afterdeck
feature are line haulers to pull the large traps (7 x 7 x 3 ft, 2.1 x 2.1 x 1 m) with frames of
welded steel from depths down to 900 ft (150 fathoms).
King crab vessels must be large and sturdy, with enough room aboard to carry about 30
crab traps, each about the size of a small grand piano, and the hydraulic gear to haul them
from considerable depths. Once on deck, crabs are loosened from the traps and dumped into
live holding tanks. Because these traps are heavy and awkward to handle, fishermen are
continually seeking a better crab trap. Tangle nets and trawls were used in the early king
crab fishery, but were unpopular not only because they damaged the catch, but, because of
the crabs' spines, it took a long time to untangle them from the webbing.

Multi-Fisheries Vessels (Combination Vessels)

Many fishermen seeking a single species use vessels designed to catch that species. Over-
fishing has resulted in more and more closed seasons; poor runs throw fishermen out of
work. Some take land jobs and return to fishing when it appears to be profitable again. Oth-
ers who are able to make changes in their vessels to shift to fishing in other areas or for other
species. Changes in market demand can trigger increased fishing effort on a certain species,
or find or create a market for underutilized species, either of which may require different
fishing gear and methods (Figs. 8.8, 8.9).

Figure 8.8. Forty-eight ft (15 m) vessel rigged

for net fishing (net reel over the stern working
area and trap fishing structure on stern lifts
traps). Note traps stored on dock during off
season. Photo courtesy National Marine
Fisheries Service. Photographer M. Bert.

Figure 8.9. Small Caribbean multipurpose

fishing schooner. Note arrowhead traps on
dock. Photo courtesy National Marine
Fisheries Service.
Chapter 8 Fishing Vessels, Gear, and Methods 159

Examples of vessels participating in more than one fishery can be found on the North At-
lantic coast. Before 1950, large otter trawls fished for offshore species, mainly haddock, cod,
gray sole, and other bottom fish, but are now also being used to catch ocean perch and yel-
low tail founder. Small trawlers specialized in silver hake and pollock. Medium trawlers,
able to fish offshore, dragged for sea scallops and lobsters in deep water. An example of an
important vessel on the Pacific coast of the United States is the Pacific, or western, combina-
tion seiner. Since its arrival around the end of World War I, many countries with coasts on
the eastern Pacific have adopted it, adding various gear configurations, to fish waters from
Alaska to Chile for species including anchoveta, tuna, mackerel, anchovy, cold water shrimp,
salmon, and ocean crabs.

Radical Vessel Designs-Catamarans

Catamarans as fishing vessels have been designed for experimental commercial fishing.
Catamarans provide a large very stable deck work platform, low rolling amplitudes, minimal
deck wetness, high maneuverability, improved accommodations and working conditions, and
the ability to fish from more than one stern ramp, all attractive features. On the down side,
there is a tendency to overload the large deck area.
In 1968, the Russians built a 120 ft (36.5 m) catamaran for experimental fishing (Fig. 8.10).
On a 59 day trial fishing trip, half the time spent in stormy seas, the vessel functioned well,
and outfished neighboring trawlers. Using both stern ramps, the crew was able to shoot the
second trawl within 3 minutes of retrieving the first. Changing the vessel from trawler to
purse seiner took less than 1 hour. Today catamarans are used for fishing in some areas of
the world, e.g., France and Norway.

Figure B.I0. Twin-hulled fishing vessels

(catamarans) have many advantages over
conventional single-hulled vessels. This 35 it
(11 m) catamaran built in Norway is a pioneer
vessel for use in coastal fishing. Proposals are
being considered for constructing catamarans
up to 130 ft (40 m) for use in purse seining and
longlining. Thirty fishing catamarans are
currently in use in France. Photo courtesy of
Marintek Sintef Group, Trondheim, Norway.


Tuna Purse Seines

After the tuna seine "revolution" between 1959 and 1961 (changeover from live-bait fish-
ing to purse seines of 1959-1961), an unforeseen development caused considerable consterna-
tion both to conservationists and the fishing industry, and still does (Fig. 8.11). Incidental to
purse seining for valuable yellowfin tunas, many porpoises are caught causing a high death
160 Part Three Fisheries

(a) (b)

Figure 8.11. (a) Pacific pole and line tuna clipper. The live bait tank is located on the stern and the
racks where fishermen stand while fishing are pulled up against the hull. From Sundstrom (1957). U.S.
Fish Wildl. Serv., Circ. 48. (b) Tuna clipper after conversion to a purse seiner. Racks and live bait tank
are removed and replaced with a turntable seine and seine skiff. Note hydraulic hauler and lines to
pull the seine aboard. From Yoshida (1966). In Proceedings of the Governor's Conference on Central
Pacific Fisheries Resources. pp. 67-89.

and injury rate among them. Furthermore porpoises in the tuna seines slow the fishing op-
eration and, in general, are a bother to fishermen (Table 8.3).
To prevent harm to the porpoises, two gear modifications, tried even before the 1972 Ma-
rine Mammal Protection Act came into effect, were "backdown" and placement of a Medina
panel (a safety panel named for the designers). Backdown means that after the seine is set
around a tuna school and about two-thirds of the net is recovered, the vessel is run in reverse
in a wide arc. The pressure caused by pulling the net through the water causes the porpoises
to accumulate at the sea surface at the apex of the net, and the corkline to submerge, thus al-
lowing porpoises to escape, but leaving the tuna in the net.
With standard nets of about 4 in (10 cm) stretched mesh, the porpoises tend to catch their
beaks or pectoral fins in the webbing during the backdown. To avoid this problem, webbing
in a special panel with small mesh (1 to 2 in, 1.3 to 5 cm stretched) prevents entanglement as
they swim over the apex of the net. These adjustments greatly reduced porpoise injury and
mortality, but still does not completely prevent it.

Menhaden Purse Seining

Menhaden fishing before 1958 was extremely labor intensive hauling the large nets by
hand. After 1958 the hydraulic power block became standard equipment on menhaden seine
vessels. The power block passes the purse seine net through an elevated free-swinging
power sheaf. Gravity forces the net into the "V" of the sheaf and gives traction that pulls the
net out of the water to the deck. Traditionally, gear pulled by hand required many men in
the seine skiffs, whose effort was limited by physical exhaustion. The power block offers nu-
Chapter 8 Fishing Vessels, Gear, and Methods 161

Table 8.3 Examples of Kinds of Fishing Gear Used in U.s. Marine Fisheries

Encircling or Encompassing
Haul-mullet/ seatrout/kingfish/ flatfish
Purse-salmon/ anchovy / sardine/ menhaden/herring/ mackerel
Lampara-Tuna bait
Beam-cold water shrimp/flatfish
Otter-flatfish/ shrimp (warm and cold water) / pollock/ rockfish/ snapper / grouper / flounder /
sea scallops/flat fish
Pound and trap nets-menhaden/reef fish/herring
Hoop nets-bait fish
Pots and traps-crabs (Dungeness)/long legged crabs/blue crabs/grouper
Slat traps-northen lobster/spiny lobster
Gill nets
Anchor-sea bass
Drift-Spanish mackerel/ocean perch/flying fish/ sharks/ flatfish/ croaker /prawn/lobster /
squid/ salmon/ cod (some of this is Japanese)
Runaround-bluefish/king macherel!Spanish mackerel
Trammel nets-mullet/ seatrout
Hand-snapper / grouper / red fish
Reeled lines-bottom fish
Troll-albacore/ salmon/king macherel! dolphin
Trot with baits-blue crabs
Bottom (ground) lines-halibut
Longline (flag line)-tuna/ rock fish/ swordfish/ sailfish/ marlin/ cod
Dip nets-blue crabs
Lift (blanket) nets-bait fish/ saury
Reef nets-salmon
Push nets-shrimp
Cast nets-shrimp mullet
Shellfish gears
Dredges-oysters / scalps
Tongs and oyster grabs-oysters
Rakes-mussels/ clams/bait worms
Hoes and forks-clams
Crowfoot bars-abalone
Diving gear-abalone / conch/ sea urchins / stone crabs
Wading by hand-clams/ oysters/ mussels
Modified after Dumont, W. H. and G. T. Sundstrom. (1957). Commercial fishing gear of the United States. U.S. Fish
and Wild. Servo Circular 109.
162 Part Three Fisheries

merous advantages to the menhaden fishery:

1. fewer men required (maybe half as many as needed for hand-hauling nets);
2. fishing can be carried out in rough weather because seine haulers avoid danger to fishermen;
3. net wear is reduced;
4. quick retrieval of nets prevents gear loss in emergencies;
5. because of the hauling speed, more sets per day are possible.



In the early days, cast nets and beach seines, also called haul or drag seines, were used. The
disadvantages of these types of fishing gear include restricted operation in shallow water
nearshore and the need for considerable manpower, plus extra hands when shrimp occur
seasonally nearshore. Catches were limited by the density of shrimp and endurance of the
fisherman. Following the introduction of the otter trawl at Fernandina, Florida, during the
early 1900s, the haul seine gradually declined in use. Fishermen in Louisiana were the last
to discontinue them.
The shrimp trawler in general use until about 1938 was a small, shallow-draft boat called
a "shrimp lugger," designed for inside and nearshore waters. These vessels, popular in Lou-
isiana, ranged from 25 to 50 ft (7.6 to 15.2 m) in length with the engine and wheel house in
the stern and the shrimp hold forward. They fished butterfly nets (wing nets mounted in the
bow of the vessel) or a single otter trawl off the stern.
The Florida-type trawler, which has replaced the lugger, has the wheel house and engine
forward and the shrimp hold in the stern, and is larger, about 70 ft (21.3 m) in length. Today,
nearly all offshore shrimp vessels fish two otter trawls. A more recent innovation rearranges
the winch configuration and reduces net sizes so four smaller trawls can be towed simulta-
Otter trawls have advantages over the haul (beach) seine; they can be used yearround in
deep water and operate with less manpower. The otter trawl is a bottom trawl using a cone-
shaped mesh bag that tapers from front to back with two lateral wings. The wings guide the

Figure 8.12. Florida style offshore double-

rigged trawler widely used in the southeastern
United States and Gulf of Mexico warm water
shrimp fisheries. Photo courtesy Texas Parks
and Wildlife Department.
Chapter 8 Fishing Vessels, Gear, and Methods 163

shrimp into the large central net opening (called a mouth). The small end of the cone called
a cod end is tied to retain the catch during fishing. At the wide end, each wing is attached
to the top and bottom ends of a flat, weighted board (called a door). Bridle-attached cables
tilt the doors to make them tow upright on the bottom and run to a winch on the vessel's
deck. It is powered by the main engine. As the trawl is towed under the water, the doors
act as kites to hold it close to the bottom and keep the mouth open. Floats on the top line
buoy the head rope. Shrimp jump when disturbed, and the overhang projecting forward and
above the bottom meshes prevents them from escaping over the top of the net. To further in-
crease catches, a "tickler chain" is fastened between the wings on the bottom to roust shrimp
from the bottom to be caught as the net passes by. Otter trawls are widely used in many fin-
fish fisheries to catch bottom fish, including cod, turbot, haddock, flatfish, etc.

Beam Trawl
Still in use, the old-fashioned beam trawl was developed by Europeans for use in the bot-
tom fisheries of the North Sea. The beam trawl, a large net bag that has a structural member
or beam that holds the net mouth open from side to side, eliminating the need for trawl
doors, thereby reducing water resistance and requiring less power to tow than for an otter
Beam trawls are also used to catch cold water shrimp in southeastern Alaska. Experi-
ments have been conducted to see if this trawl will catch penaeid shrimp in the southeastern
United States and the Gulf of Mexico. Early results show that beam trawls not only may
have higher catch efficiencies than otter trawls of the same size, but save fuel as well. How-
ever, the long, rigid beam that may be as long as 40 feet, is difficult and dangerous to handle
at sea, especially during heavy weather. This disadvantage alone may prevent its wide adop-
tion in the southeastern U.S. shrimp fishery.

Mid-Water (Pelagic) Trawls

During the 1960s a West German net maker developed mid-water trawling methods to
catch herring in the northeast Atlantic by utilizing earlier improvements in fish capture tech-
niques, such as the use of hydraulic power for winches instead of electrical power, and the
information that synthetic fibers reduced water resistance compared with natural fibers. The
name is descriptive. A mid-water trawl operates between the surface and the bottom. As
with any new fishing technique that shows promise of better landings, in due course mid-
water trawling was tried by fishermen in other countries and for capture of schooling, de-
mersal species other than herring, including cod and haddock.
Mid-water trawls can be fished from a single boat, or between two boats fishing side by
side. In a single-boat operation, net doors are used to hold the net mouth open horizontally.
Floats are attached to the head rope and weights to the foot rope to hold the mouth open ver-
tically. In two-vessel fishing, the distance between the vessels holds the net open from side
to side. To be effective, mid-water trawling requires rather tightly schooled fish, knowledge
about the depth off the bottom where the fish are swimming, and the depth at which the net
is fishing. Transducers and pressure capsules are used to obtain depth measurements. Con-
siderable time is used to locate desirable schools, determine the direction they are moving,
and set the net to pass through the target school. Experienced fishermen using mid-water
trawls can make a larger catch per tow than otter trawls, but searching time and net-setting
procedures limit the frequency of tows per day of fishing. However, tows made by mid-wa-
ter trawls are usually of much shorter duration than for bottom-fishing otter trawls.
164 Part Three Fisheries

Tuna Live Bait-A Search Fishery

For many years surface-swimming tuna were caught off the coast of Mexico south to Ec-
uador using live bait techniques from large vessels up to about 150 ft (46 m) in length. A
prominent feature of these vessels is the crow's nest, or lookout post, on the mast where a
crewman is stationed to look for bird flocks that fly low over the ocean's surface eating small
prey fish. Under the bird flocks, schools of tuna are eating the same prey as the birds. Once
sighted, the captain positions the vessel in front of the fish school and moves in the same di-
rection as the school, holding the school close to the vesseL Live bait held in a large tank at
the stern where all fishing activity was entered, is thrown into the water by a chummer.
When the bait fish cause the tuna go into a feeding frenzy, they can be quickly caught.
The first step in this three-step fishing operation requires catching a supply of bait (Fig.
8.13). Using a lampara seine, the bait is caught, transferred into a small live-well bait barge,
and delivered to the bait tank on the stern off the vesseL The second step is the search for a
sizable tuna school, then to position the vessel just ahead of the schooL The third and last
step is the catching process. Fishermen standing in racks fish on the stern with pole and
lines using feathered jigs with barbless hooks that resemble the bait fish. They may fish sin-
gle poles, or share a single line attached to more than one pole. The size of the individual

Figure 8.13. Pole and line tuna fishing.

Chummer using small dip net removes small
bait fish from tank behind him and throws
them behind the boat to hold the school in
reach of the fishermen. Photo by the author.
Chapter 8 Fishing Vessels, Gear, and Methods 165

fish in the school determines the number of poles used to catch individual fish. It is highly
labor-intensive method. This kind of tuna fishing is not done to any extent off the United
States mainland and Central America, but continues in Hawaii and by Japanese in their home
waters (Fig. 8.14). The live bait fishery that once operated out of California ports has largely
been replaced by a purse-seine fishery to catch surface-swimming tunas (Fig. 8.11).

Figure 8.14. Hawaiian skipjack pole and line

vessel (sampan) measuring between 50 and 80
ft (15 to 24 m) Live bait is used to fish for
surface schools. Photo by the author.

Halibut Longline (Ground line)

Pacific halibut are caught by hook and line gear off Canada and Alaska. The halibut lon-
gline, also called a ground line, is anchored to the bottom. Buoys with flags and sometimes
lights help fishermen to relocate the gear after it is set. From the long mainline, many branch
lines or gangling lines are attached with hooks baited with dead fish, played out over a stern
chute while the vessel is rapidly running; each unit of gear is called a "skate," or basket.
When it has "soaked" (fished) sufficiently long, the gear is retrieved to the boat by a small
power winch called a line hauler.
Halibut caught on ground lines are damaged by sea lions which concerned the halibut
fishermen of both Canada and United States who believed that large qualities of halibut were
destroyed by Steller's sea lions. Between 1958 and 1960 only about 8 percent by weight of
the halibut catch was estimated to damaged and unsellable. Halibut are not believed to be a
significant food for sea lions.

Another type of commercial hook and line fishing is called trolling. In south Florida wa-
ters it is also done in recreational fisheries for dolphin fish, mackerel, and tunas. In commer-
cial salmon trolling, artificial plugs (lures) are towed at slow to moderate speeds near the
surface. Long poles with lure-bearing lines attached extend from either side of the trolling
vessel. This type of fishery is practiced in Canada, Washington, Oregon, and Alaska, gener-
ally by a single fisherman on each boat, or sometimes a husband and wife team.

Snapper Hook and Line Fishing

Snapper fishing is also a hook and line fishery done from an anchored or drifting vessel
with fishermen standing around the rails of the vessel. Formerly, only hand lines were used,
then fishing reels with wire lines appeared near Campeche, Mexico and around the Gulf of
166 Part Three Fisheries

Mexico. More recently, power reels have come into use. A sounding lead has been used by
fishermen to determine the depth of reef areas where snappers are known to live, but depth
is now generally determined by electronic depth finders.

Tuna, Marlin, and Swordfish Longline

The last important type of hook and line gear is a pelagic gear called longline. It is fished
in the open ocean at great depths. Unanchored, it drifts with the current and catches sword-
fish, tuna, and marlin (Fig. 8.15). Japanese fishermen characteristically set about 70 mi (113
km) of line and adjust the droppers having hooks and bait attached to the depth most suita-
ble for catching the species of fish they are after. Gear is usually set soon after sunrise and
retrieved during the evening hours. Droppers are hung at intervals along the main line kept
afloat by buoys. Flags or radar screens to pick up signals are attached to buoys to help locate
the gear in heavy weather. Much the same as halibut ground line, this gear is paid out with
the vessel under way and also brought back aboard the vessel by a line hauler. It is truly an
impressive sight to watch the speed and skill of Japanese fishermen setting and retrieving
longline gear. Large fish are caught on the droppers and they are removed from the main
line and pulled aboard by fishermen while the gear continues to come aboard.

Figure 8.15. Floating longline gear for catching deep swimming tunas. From Umali (1950). U.S. Fish
Wildl. Servo Res. Rept. No 17.

Longline gear, pioneered by the Japanese over 200 years ago, is still widely used today,
with some modifications. Dead sauries, herring, and anchovies are used as bait. The method
is not trouble free. There may be considerable shark damage to the fishes caught in the Pa-
cific on longline gear, and occasionally large sections of gear may be tom loose and lost.

Salmon Traps
About 1885 in Cook Inlet, Alaska, the first salmon trap was constructed in the style of the
Great Lakes pound net (a fish trap with a wing to direct fish into an enclosure). Another
Chapter 8 Fishing Vessels, Gear, and Methods 167

type of salmon trap (the pile trap) uses logs or piles that are driven into the bottom to sup-
port walls of webbing and wire netting that guide the salmon into the trap and confine them
until the fishermen remove them (Fig. 8.16). Shallow water with a soft bottom is desirable to
construct these traps, but in southeastern Alaska many locations suitable for trapping salmon
are too deep or have rocky bottoms. To remedy this, floating traps were conceived about
1890. Similar to pile traps, floaters are buoyed by large logs with the walls of webbing an-
chored to the bottom by boat anchors and rocks. Both pile traps and floating traps consist of
a lead, or fence, that runs perpendicular from the shore to the heart of the trap to intercept
salmon migrating along shore, directing them into the hearts and spillers of the trap. Spillers
hold the fish and are arranged so that salmon can be netted or brailed out and loaded onto
a scow when needed, then taken to the cannery. Trap efficiency (a good location could catch
up to 100,000 salmon per day) is blamed for declines in salmon stocks. By 1958, all traps
were phased out of Alaska except those used by American Indian tribes.

Figure 8.16. Floating Salmon trap. From

Sundstrom (1957). U.S. Fish Wildl. Serv., Cire.

Another problem concerned trap ownership and fishing gear competition. Large packing
companies from outside Alaska owned traps and were disliked intensely by local purse sein-
ers, gill netters, and trollers. When eventually the trap issue came to a vote in 1956, the votes
of many small fishermen overwhelmingly outnumbered those of relatively few cannery own-
ers for abolishment of salmon traps (20,872 to 3,946).

Cast Net Fishing

The cast net is said to date back several thousand years to the Malayan Peninsula. During
part of its history it was used for catching birds. A simple device, a cast net consists of a cir-
cular piece of netting with lead weights secured around the perimeter that is tucked under to
form a pocket. A rope is rigged through the center or apex of the net for retrieval. The di-
ameter varies according to the net maker. Cast nets are used to catch schooling and migrat-
ing fish and shrimp. Mullet and shrimp fishermen use cast nets in shallow waters. A variety
of bait and food fish can be taken as well.
168 Part Three Fisheries

To make the cast, a fisherman holds some of the leads between his teeth while holding the
perimeter of the net with his arms extended, body leaning backward and turned to the right.
The net is swung about 180 0 and released so that the lead weights open the net as wide as
possible before it hits the water over a school of fish. Considerable practice is required to
cause the net to open fully and have it land over the target species, and also to avoid losing
teeth. The leads must be heavy to carry the thrown net through the air and cause it to sink
rapidly over the school of fish to entrap them before they escape. When the retrieval line is
pulled, the lead line purses so the fisherman can haul in his catch. The cast net can be used
by wading, from the shore, from sea walls, and from docks or boats.

Fish Wheels
Fish wheels have been described as the most ingenious and efficient fish catching devices
ever invented (Fig. 8.17). They are erected in rivers to catch migrating fish. Shad were taken
in them in the Pee Dee and Roanoke Rivers in North Carolina as early as about 1827, and
gear was introduced in 1879 to the Columbia River in Washington to catch salmon. By 1899
as many as 76 were in use. Fish wheels were outlawed in Oregon in 1926 and in 1934 in
Washington State. Anadramous fishes, swimming upstream into the path of the wheel, are
scooped up as the stream currents move the wheel and lifted into the center toward the axle,
then slide out to be deposited in a nearby scow or fish holding box. A fence or barrier made
of logs keeps fish from swimming between the wheel and shore.

Figure 8.17. Fishwheels were expensive to

construct on the large rivers, such as the
Columbia where 76 operated at the tum of the
century, but were efficient for harvesting
salmon. Photo courtesy of Instructional Media
services, University of Washington.

Lift or Blanket Nets

These nets are used to catch saury and other bait fish that form dense schools at the sur-
face and are attracted to light. A vessel fishing with a lift net stops the engine and lowers its
outriggers when a concentration of fish is located. The net is attached to the outboard end of
the outriggers and hangs vertically in the water. Lines are attached so the net can be pulled
to the side of the vessel to a ''blanket'' a short distance beneath the water. Lights placed over
the net attract the fish, but because the school may be frightened by the net, the fish may
swim to the side of the vessel away from the net. By extinguishing some lights and lighting
others, the school is maneuvered around the vessel until it is over the net. At this point, the
outriggers are raised and the trapped fish are herded close to the vessel where brailing (tak-
ing fish from the blanket net to the vessels hold) begins. Still, dark nights, and clear water
provide ideal lift-net conditions.
Chapter 8 Fishing Vessels, Gear, and Methods 169


Students of fishery science should recognize that the weight and number of fish that encoun-
ter fishing gear, and the weight of fish actually landed at the dock may differ greatly. This
apparent disparity is important in calculating population size and estimates of maximum
sustained yield and can partly affect estimates of production from the sea.
Fish encountering fishing gear mayor may not be captured, but, if captured, still might
not be brought aboard the vessel. If taken aboard the vessel, they mayor may not reach the
dock to be recorded as a landing for a variety of reasons as will be described. When landed
at the dock, they may be substantially lower in weight and numbers than what was caught
at sea due to processing (deheading, gutting, etc.) aboard the vessel.
Some fish encountering fishing gear can escape. In otter trawls, for example, fish entering
the body of the net often can swim fast enough to pass out of the mouth of the net to free-
dom. Also, fish encountering gill nets (gear which is highly selective for certain fish sizes)
may escape either through the net, or, if very large, will not become gilled.
Of the total live weight of fish caught, some may be destroyed or damaged (some 10ngline
caught fish are damaged by sharks and, seals Figs. 8.18, 8.19), and others through loss of fish-
ing gear (such as traps with buoy lines that are torn free). Next, the gross catch may be min-
imally reduced either by consumption by the crew, or by discarding. Undersized shrimp that
bring a very low market price have little demand, and are often discarded at sea. Also, many
of the species of fish in some trawl catches are unsellable. Called by-catch, it, is also dis-
carded. The retained catch can be diminished for any or all of a number of reasons such as
the consumption of caught fish by the crew, or use of fish for bait by the crew. Also, not all
weight changes of the catch are negative; for example, placing live spiny lobsters in freshwa-
ter results in a weight increase, as does salting of fish. Even the gross catch aboard the vessel
can be reduced before the catch is landed at the dock and emphasizes again that a fishery bi-
ologist must understand fishing techniques and behavior at sea before using statistics to de-
termine population parameters.

Figure S.lS. A shark attack on a yellowfin

tuna caught on longline gear in the central
equatorial Pacific. An example of precatch
losses that occur as a result of the fishing
operation. Photo by the author.

Net Selectivity of Commercial Fishing Gear

Many factors affect fish catches in commercial trawl nets, including gear avoidance by fast-
swimming species and the tendency of strong-swimming fish to escape after coming in con-
170 Part Three Fisheries

40 •
Poi red Observotions
• 1958(296)
<> 1959 (516) •
« •
a • •

... <> •
~ 20 ·0

~ 10

0 Moximum Numbers
185 Seo Lions & 48 Domoged Halibut-

10 20 30 40 50 60+


Figure 8.19. Sea lion damage to hooked halibut or those released by fishermen during 1958 and 1959.
Estimate of about 8 percent of halibut catches by weight were damaged and unsellable. From Bell
(1981). The Pacific halibut. The resource and the fishery. Alaska Northwest Publishing Co., Anchorage,

tact with the gear. Some of these losses of fish from gear were explained earlier in the gear
section. The usual measure of net selectivity of trawls is to determine for a particular mesh
size what percentage of fish of various lengths will be retained by the net covering the range
of a to 100 percent. The selectivity of the net is considered to be the fish length at which 50
percent of the fish are retained in the cod end. Relationship between fish length and percent-
age of fish retained in the cod end of a trawl is discussed in Chapter 14.
Escape of caught fish through the meshes of a net can be measured by several methods.
One method is to place a cover of mesh over the net that is smaller than at the cod end. It
will catch and retain whatever slips through the meshes of the inside trawl being tested for
the extent of escape. Trawls with two cod ends {"trouser trawls"}, each with a different mesh
size, document trawl escape from the side with the larger mesh. Similarly, two trawls with
different mesh sizes can be towed side by side from two vessels operating at the same speed,
and the catches compared.
Underwater television has been immensely helpful in this respect, by showing small fish
coming in contact with the larger mesh and escaping through the webbing. It provides proof
that fish too small for market can escape unharmed from commercial fishermen's nets with
large mesh, and convinced fishermen to use larger mesh so that the escaped fish might be
caught later at a desirable market size and/or have a chance to spawn, instead of being de-
Chapter 8 Fishing Vessels, Gear, and Methods 171

stroyed. The role of mesh size as a regulatory tool will be discussed under management
technique (minimum size limits) in Chapter 14.
Gill nets selective as to sizes of fish and their catch differs from trawls in that they have
both an upper and lower size limit selection. The size of fish caught depends on the net
mesh size and on the girth of the fish. Large fish cannot gill themselves in the mesh and
small fish can slip through the mesh. Several methods are used for measuring gill net selec-
tion; comparison of gill net catches with those of other fish gears, comparison catches of gill
nets of various mesh sizes, and theoretical considerations based on observations of fish girth
and mesh size.
The examples of gear selectivity given here are just a few of the many ways that fishing
gear is made selective. Essentially all commercial/recreational fish gear is selective to a
greater or lesser degree. Recreational fishing gear can also be selective by size of bait (artifi-
cial or natural) and/or hook sizes that are used. Again, fishery biologists who use fisheries
data in studies directed at stock management must realize that commercial fish catches are
far from random and understand how, where, and when fishermen fish in order to make use
of these samples.


It is a long known reality that certain bottom trawls and purse seines have killed nontarget
species, species unwanted by fishermen but that may be valuable on their own or to other
fisheries when they reach market sizes. These incidentally caught fish and shellfish have
been termed "by-catch" or "trash" fish, and, because of their large numbers, have been a
source of irritation to commercial fishermen, recreational fishermen, and environmentalists.
Each group has justifiable reasons for wanting this destruction of juvenile fish stopped. De-
tails on the by-catch problems and management strategies are discussed in Chapter 14, Fish-
eries Management-Laws and Regulations.


Fish Pumps
Fisheries for small species that are caught in large schools, such as herring, saury, men-
haden, and squid, make use of pumps for moving fish from nets into the holds of vessels.
Until recently, the menhaden fishery in the Gulf of Mexico transferred fish from the purse
seine held alongside the vessel by dip nets or brail nets (powered dip nets). This old method
has finally undergone modernization (Fig. 8.20). Now, a reinforced rubber hose of 12 in (0.3
m) diameter is lowered into the catch, through which water and fish are pumped to the deck
of the vessel. Fish travel by chute to the hold, while the water, by another chute, flows back
into the Gulf. When the loaded boat returns to the dock, the reverse of loading at sea takes
place. Water pumped into the hold of the vessel allow pumps to transport the catch into
boxes or onto an automatic scale on the dock, where they are measured, or weighed in units
of 1,000 fish. They are then conveyed to the plant for processing. These units determine the
crew's pay.
172 Part Three Fisheries

Figure 8.20. Today's fishing vessels use hydraulics in a wide range of applications including steering
cylinders, anchor winches, bow thrusters, deck machinery, and fishing machinery. From Modern
Marine Hydraulics advertisement.

Remote Sensing Technology

In oceanography, remote sensing simply means perceiving the presence or characteristics
of something (fish schools, plankton, measures of water temperature) at some distance and
transmitting a message to a receiver that the sensor perceives. There are two categories in
fisheries remote sensing, one where the message originates from underwater (hydroacous-
tics), and another from above the water surface.

Airborne Sensors
The crow's nest on fishing vessels is a long-used aid for sighting fish schools. In pelagic
fisheries, searching time can vary greatly, but some estimates suggest as much as 85% of a
vessel's operating time may be spent locating fish. Reduction of searching time is important,
considering the high costs of fishing. As early as the 1920s, the fishing industry began to use
aircraft for pelagic searches. Those early fliers who sat in open cockpits of seaplanes often
got soaked on landings and takeoffs, located schools, then flew low over the fishing vessels
and shouted directions to the fishermen. The advantages of spotter aircraft are readily ap-
parent; pilot observers can scan large areas quickly, can locate schools at greater depth, and
can provide accurate information on school size and their movements. Salmon on the Pacific
Chapter 8 Fishing Vessels, Gear, and Methods 173

coast and menhaden and mackerel on the Atlantic coast are examples of fisheries that have
increased their landing by the use of aircraft.
Helicopters are also used in addition to conventional aircraft. They possess qualities that
make them very efficient for searching. Helicopters can hover over fish schools, and can be
carried on the decks of large vessels such as tuna clippers, extending the search range.
Qualified observers can detect color differences that identify some species of pelagic fish.
Other features that indicate schools are fish oil films on the water's surface, the presence of
diving birds, ripples, and sometimes churning of surface waters.
Fishery scientists also use aircraft to make stock assessments of salmon and herring. The
number of salmon as they return from the sea to freshwater lakes and streams to spawn is
counted to determine whether or not the seeding of spawning beds is sufficient to insure
good runs of salmon in future years. Aircraft have done important work in this connection,
especially in Alaska.

Spaceborne Sensors
Tuna fishermen now use space age technology. They receive information on computer
terminals from satellites equipped with infrared sensors that measure surface water temper-
ature and water color variation (Fig. 8.21). The latter reflects the amount of chlorophyll
and plankton present in the water. Crabs are attracted to certain concentrations of plank-
ton, which in turn attracts concentrations of tuna in suitable temperatures. Fishermen are
able to use this service to locate schools of tuna. Cameras mounted on the boat scan the
area in the vicinity of the vessel, and the data are transmitted to NASA in Alaska where
they are processed digitally and retransmitted to the vessel. If the vessel is equipped with
telephone and facsimile equipment, a chart of the area also can be provided to the boat
captain, enabling him to locate possible concentrations of tuna and thereby cut down on
time spent searching. This is expensive, but remunerative. It is estimated that the average
cost of a tuna fishing trip can be cut by about 10% by using satellites to reduce searching

Underwater Acoustics
The basic principle of the echo sounder or "fish-finder" uses sound waves travelling in the
sea at 4,900 ft/sec. (1,493 m). By comparison radio, radar, and light waves are inefficient in
water. The echo sounder comprises four main components, transmitter, transducer, receiver,
and recorder. The transmitter sends an electric current to the transducer, that converts it to a
sound pulse. The sound is not radiated from the transducer uniformly in all directions but
is most intense in the direction at right angles to the transducer face. The transducer behaves
like a wide-angle search light. Sounds are beamed to the ocean floor and the returning ech-
oes are picked up by the same transducer, acting now as a microphone. The sounds are then
converted back into electronic signals that are directed to the receiver where they are ampli-
fied and passed on to the recorder. The recorder is the ''brain'' of the echo sounder. It starts
the transmitter, then records the echo after it has been magnified approximately 1 million
times by a pen passing over special recording paper. The speeds of both pen and paper can
be adjusted to alter the picture. When the recorder instructs the transmitter to send out a
sound pulse from the transducer, the pen in the recording unit marks the paper. This is
known as the zero" mark and is made the moment the sound leaves the transducer. Thus
the profile of the sea floor is traced from this point.
This information can greatly enhance the ability of the fishermen to catch bottom fish and
174 Part Three Fisheries

~.... NOAA-7 Nlmbu.-7

Satelllt. Oceanography 810
Facility VI.lblllt Laboratory
Data Received CZCS Data
and Archived
Data Processing
AVHRR Data Ocean Color
Data Proca •• lng Imagery

Sea Surface T emparature

and Chlorophyl Imagery Comm.rclal
V•••• I.

Fish Catch Data

Ocean Environment/Fisheries Relationships

Figure 8.21. Satellite data collection and processing network utilized by the National Marine Fisheries
Service on the west coast of the United States. Form Fiedler, Smith, and Laurs (1984). Mar. Fish. Rev.

shellfish and also can prevent the loss of expensive fishing gear that might be caught and de-
stroyed on a rocky or uneven bottom.


Benson, N. G. 1970. A century of fisheries in North America. Am. Fish. Soc. Spec. Publ. 7. Washington,
Browning, R. J. 1974. Fisheries of the north Pacific. History, species, gear and processing. Alaska
Northwest Publ. Co., Anchorage, AK.
Captiva, F. J. 1979. The American fisheries 1620--1979. Fishing Gazette. November 1979:51-90.
Donaldson, 1. J. and F. K. Cramer. 1971. Fishwheels of the Columbia. Binfords & Mort, Publ., Portland,
Jensen, A. C. 1967. A brief history of the New England offshore fisheries. U.S. Fish and Wildl. Servo
Fishery Leaflet 594. 14 pp.
Samples, K. C. 1983. An economic appraisal of sail-assisted commercial fishing vessels in Hawaiian
waters. Mar. Fish. Rev. 45(7-9):50--55.
Seki, H., A. Hamada, T. Iwami, and R. J. LeBrasseur. 1981. Hobikiami sail trawling in Japan. Fisheries
Smith, C. L. 1984. Evolution of the Pacific canned salmon fishery. ORESU-R-84-035. 15 pp.
Tunstall, J. 1962. The fishermen. Macgibbon and Kee, Ltd., London.
Chapter 8 Fishing Vessels, Gear, and Methods 175

Commercial Fishing Gear and Methods

Browning, R. J. 1974. Fisheries of the north Pacific: History, species, gear and processing. Alaska
Northwest Publ. Co., Anchorage, AK.
Gusey, W. F. 1978. The fish and wildlife resources of the Gulf of Alaska. Shell Oil Co.
Fowle, S. and R. Bierce (eds.). 1991. Proceedings of the shrimp trawl bycatch workshop, November 22-
23,1991. Center for Marine Conservation. Washington, OC. 183 pp.
Hintz, C. R. 1969. Fishing vessels and support ships. In F. E. Firth (ed.). The encyclopedia of marine
resources. 250-260. Van Nostrand Reinhold Co., New York.
Kristjonsson, H. (ed.). 1965. Modem fishing gear of the world. Fishing News (Books) Ltd., London.
Pike, D. 1988. Fishing boats and their equipment. Fishing News (Books) Ltd., London. (Third Edi-
Provenzano, A. J. (ed.). 1985. The biology of Crustacea. Vol. 10 Fisheries and culture. Academic Press.,
Orlando, FL.
Sainsbury, J. C. 1986. Commercial fishing methods-an introduction to vessels and gear. Fishing News
(Books) Ltd., Farnham, Surrey, England (Second Edition).
Shaw, S. and J. F. Muir. 1987. Salmon: Economics and marketing. Timber Press, Portland, OR.
Spence, A. 1989. Crab and lobster fishing. Fishing News (Books) Ltd., Farnham, Surrey, England.
Sundstrom, G. T. 1965. Commercial fishing vessels and gear. U.S. Fish and Wildlife Service Circular
48. 48 pp.
Traung, J-O. Fishing boats of the world. Vol. 1, 1955, Vol. 2, 1960, Vol. 3, 1967. Fishing News (Books)
Ltd., London.
von Brandt, A. 1964. Fish catching methods of the world. Fishing News (Books) Ltd., London.
Chapter 9

Food and Nonfood Fisheries

Most fish species contain less than 5% oil. This is of importance to calorie watchers who
know that lean meat contains usually 5 to 15%. Seafoods supply amino acids needed by hu-
mans, in balanced amounts. Studies sponsored by the American Heart Association show that
diets high in fish result in lower rates of coronary attacks in middle-aged men. This reflects
the beneficial polyunsaturated oils in seafoods that also have a high content of certain essen-
tial minerals and water-soluble vitamins.
Some shellfish suffered the consequences of an erroneous measurement made by early re-
searchers on cholesterol levels in shellfish. Improved analytical techniques that enable scien-
tists to distinguish cholesterol from its chemical relatives, show that cholesterol content in
shrimp, for example, is much lower than previously thought. In addition, shrimp contain
beneficial omega-3 fatty acids that may actually decrease the risk of heart disease. New
measuring techniques show cholesterol levels in shellfish to be lOw. Cholesterol content in a
single species of fish can vary widely at different times of the year, at different growth stages,
and in different reproductive stages. These variables indicate the need for additional study
to determine basic cholesterol levels, and, hence, the possible benefits of fish in the diet to
help control vascular diseases.
Fish meal is used worldwide as a protein supplement in animal feeds, and is known to cause
faster growth in poultry and other farm animals than vegetable protein alone. The fish species
used to make fish meal are generally the small schooling fish called "industrial fish" or "reduc-
tion fish" that are not used for food for example, menhaden, Brevoortia sp. (Table 9.1).

Table 9.1 Breakdown of Processing Methods in the U.s. Fisheries

Percent of total
by value

Fresh and frozen 65
Canned 27
Cured 3
Total edible 95
Bait and animal food (canned) 2
Fish meal, oil, and Solubles 3
Total Industrial 5
Grand Total 100
From National Marine Fisheries Service.

Chapter 9 Food and Nonfood Fisheries 177


Fish landed on vessels may not be completely fresh. Gill-netted fish and longline-caught
tuna may hang dead in the gear for some time before the catch is hauled aboard the vessel.
When catches are large, trawl-caught fish are subjected to considerable pressure from the
weight of the catch, and die in the net. Even when catches are small, fish are tumbled and
die in the gear where spoilage begins.

Preventing Spoilage
Spoilage is related to temperature. It is retarded by freezing or icing the fish, a short-term
means of preserving quality (Fig. 9.1). Quality can be preserved for longer periods by freez-
ing. Because the speed of chemical reactions doubles for every lOoF (5.5 °C ) increase, spoil-
age in tropical and subtropical areas is a greater problem than in temperate waters.
Unfortunately, ice to cool fish can be hard to find, or costly, as in most developing countries.
Freezers and refrigerators are often too expensive for small vessel owners, so short trips are
made to preserve the quality of the catch. In some small fisheries, part of the hull is sealed
off as a "live well," with water exchanged from outside the vessel to keep fish alive and in
good condition until they can be sold.
Because the mucus layer on fish protects it from invasion by certain bacteria, it is desirable
to keep it intact even after death. Gutting the fish destroys this natural barrier.

Figure 9.1. Icing processed fish prior to

shipment. Photo courtesy of Snake River Trout
farm, Buhl, ID.
178 Part Three Fisheries

To reduce effects of spoilage at sea:

1. keep fishing trips short;

2. using holding facilities to keep catch alive;
3. keep dead fish wet and protected from the sun (wet burlap bags or in tropical areas palm
fronds can be used);
4. bathe fish frequently with seawater;
5. provide good air circulation;
6. decks and fish bins should be scrubbed with a chlorine-water mixture.

Fresh fish can be identified by: firm, elastic feel of the flesh; belly not bulging; no "fish" odor;
eyes bright, transparent, and not shrunken; gills bright red; and skin not faded. Shrimp held on
ice for extended periods develop black spotting. To improve the appearance of the shrimp for
market, the spots can be removed by a sodium bisulfite dip approved by the U.S. Food and
Drug Administration (FDA) that has standards on traces of this chemical in the shrimp flesh.
Concentrated sodium bisulfite dips can leave an undesirable off-taste. Good handling practices
start aboard the vessel, and should not wait until fish reach the processing plant.

Processing Fish and Shellfish

Fish and shellfish are processed several ways before sale. Fresh fish are usually cleaned,
or cut into fillets or steaks. The National Marine Fisheries Service (NMFS) recommends these
additional methods (Fig. 9.2):
Batter-coated fish products. Sticks and portions or other forms of fish or shellfish coated
with a batter containing a leavening agent and mixture of cereal products, flavoring, and
other ingredients, and partially cooked in hot oil a short time to expand and set the batter.
Breaded fish products. Sticks and portions or other forms of fish or shellfish coated with
nonleavened mixture containing cereal products, flavoring, and other ingredients. Breaded
products are sold raw or partially cooked.
Canned fishery products. Fish, shellfish, or other aquatic animals packed in cans, or
other containers, that are hermetically sealed and heat-sterilized. Canned fishery products
may include milk, vegetable, or other products. Most, but not all, canned fishery products
can be stored at room temperature for an indefinite time without spoiling. Canning, used
for a variety of fishes including salmon, tuna, herring and sardines, requires expensive,
complicated machinery and high technology. (Fig. 9.3).
Cured fishery products. Products preserved by drying, pickling, salting, or smoking; not
including canned, frozen irradiated, or pasteurized products. Dried products are cured by
sun or air-drying; pickled or salted products are those products preserved by applying salt,
or by pickling (immersing in brine or in a vinegar or other preservative solution); smoked
products are cured with smoke or a combination of smoking and drying or salting (Fig. 9.4).
Curing reduces the moisture content necessary for bacteria to thrive. Smoking usually re-
quires that salt be used first to remove some of the moisture. Only certain kinds of woods
are suitable for smoking and for prescribed periods of time for the smoke to penetrate into
the fish flesh.

Surimi is a Japanese fish cake. The product, which looks like meat loaf or sausage meat,
is believed to have originated in Japan over 400 years ago (Fig. 9.5.). Fishes used are croak-
ers, lizard fishes, sharks, and walleye pollock, Theragra chalcogramma. It is made by removing
the head and viscera from the fish to keep the final product from darkening, then washed
several times before passing through a meat-bone separator. Excess water is removed and
the flesh is ground, and salt is added to bind the protein and starch and improve its elasticity.
Chapter 9 Food and Nonfood Fisheries 179


Fresh and frozen fish may be bought in .. variety of cuts, the more
important of which are shown here. Knowing the cuts and their
particular uses is important in buying or selling fish. The edible
portion.varies with the type of cut, from 100 percent for fillets to about
45 percent Cor whole fish.

Whole 01' roud fiah are thOll8 marketed

just sa the,. come from. the water. ID
this form, the edible portion iI about 45 Stem are crou-eection alicel of the larger
percent of the whole, but varies with aiM lilel of dresIed fish, uaually about", of an
and klnd of Alh. To prepare for cooking, Incb thick.. In this form the edible por-
fish should be scaled and eviscerated And, tion is about Me percont. Steak, are

if desired, head, tail, and flo! should be rady to cook as purchaled.
removed. Fish then may be taed for
baking, or may be aliced, &leted, or cut I
into ateaka or chunb. Small fiah, like
omelt, an often cooked wilh the eo-
traila removed. are the sides of fish cut a,..ay from
They an practically
Dra.... filh are thole marketed with onl,. and have little or no Waite.
the eotraila removed. In this form, the .... ready for oookillg. The Ikin
edible portion is about 48 percent, but be left on or may be removed. A
varies with aim and kind of fish. To cut from ODe side 01 .. fish iJ caJled
prepare for cooking, they are generally fillet. ThIa is the type moat
acaled~ Head, tail, and fin, may be Jeen in the mar.ltet.
removed, if deoired, and the fiah epllt,
fllleted.. or cut into .teaks or chunb.
Dreu. flab are acrJed aod en.cerated,
usually with the bead, tail, and fioa re-
moved. Edible porUon in tbit form it
about 67 pereent. but varies with aile aod
kind of flab. The smaller lIiHl are ready
tor cooking .. purchased (pan dreaaed.).
The larger Bisel may be baked .. pur.. are piecee of fish cut lengthwise Of
chued or may be cut into fU1etll, steak&, from fillet. in to portions of uni-
aDd le.gth, Ulu&lly about I
or chunD.
and 3 inchll long,


Some shell.6sh are marketed alive. Other market forms, depending
on the variety, include cooked whole in the shell, freahmeat (shucked),
headl_, and cooked meat.

ID Ihell: Shelllllh, luch .. hard and IIOfI Beadl_: ODly \he tall pari of Ihrimp II
blue crabe, lobaten, clams, and O)"llterw commonly marketed. Spiny-Iobater tails
should be alive if bought fresh in tbe IheU. aN aIMI a common market form. About
Crabs and loblten may &lao be cooked in 86 percent is edible.
the ahell. Edible portion .ariea widely.

CMtod aeat: The edible portio. II

plcked from \he eooked ohelllllh. Crab,
shrimp, and lobater meat is marketed tn
Shucked: Clam, oyater. and acallop meate t.hit way. Cooked meat is perishable,
may be bought free of the shell, commouly althouch packaged lD eont.&inen. since it.
known .. ahucked. In this. form the. por- II Dot further procoued 1>7 beat. It II
tion it 100 percent edible. 100 peree.1 edible.

Figure 9.2. Market forms of fish. From Anon (1954). Fresh and Frozen Fish Buying Manual. Circular 20
U.S. Fish and Wildlife Service.

Other materials such as monosodium glutamate and sugar provide flavoring. At this stage
the product has become a paste called surimi. It is an odorless block of white fish flesh that,
frozen, has a shelf life of about 7 months.
Drastic reduction in catches of king crab, Paralithodes spp., sent the fishing industry search-
ing for a substitute product. Artificial crab made from surimi, with similar color, shape, and
texture, and imported from Japan has become popular in the United States. Increased fishing
on Gulf of Alaska walleye pollock by U.S. fishermen (only) has caused construction of
180 Part Three Fisheries

Figure 9.3. Drawing from an early

photograph of children at a sardine cannery
preparing small herring (called sardines) for
canning. From Goode (1884). U.s.
Commission of Fish and Fisheries.
Washington, D.C.

Figure 9.4. Numerous drying and salting

plants appeared late in the 1800s. Salmon and
cod were important species processed by these
techniques. This photograph taken between
1907 and 1911 shows a west Seattle cod drying
plant. Photograph courtesy of University of
Washington Special Collections.

processing plants in Alaska and Washington State to produce imitation or simulated seafood
called surimi, or "analog products." Other species that are fabricated from processed fish
meat include lobster, shrimp, scallop, and some finfish.

Processing Machinery and Personnel

The high cost of catching, handling, and processing certain fish and shellfish is a signifi-
cant problem facing fisheries throughout the world. Expansion of certain fisheries is delayed
Chapter 9 Food and Nonfood Fisheries 181


DRESSING (Removal of Bone. Skin. Guts)








Figure 9.5. Steps in the production of

surimi. From Sonu (1986). NOAA Technical FROZEN SURIMI
Memorandum NMF5-SWR-013. 122 pp.

due to high processing costs requiring hand labor despite an abundance and market. Exam-
ples of species requiring costly hand labor during processing are given below (Fig. 9.6). The
seafood industry employs substantial labor on fishing vessels (Chapter 8), unloading,
processing, and packaging the finished product. For example, a large breaded-shrimp plant,
with a production of 3,000 to 4,000 lb (1,361 to 1,814 kg) of shrimp per hour, may employ 200
to 300 people to sort and package the product. The work is routine and messy, in damp and
sometimes cold conditions, and is generally low pay. It can also be seasonal, depending on
the availability of the fish/shellfish and market demand. This presents a hardship on work-
ers, many of whom live hand to mouth. Some oyster plants rent company housing to work-
ers, and others provide transportation to the plant if they live some distance away.
Prior to World War II, oyster plant workers along the eastern seaboard and Gulf of Mexico
employed mostly African Americans as shrimp workers, and/or undocumented aliens who
could not qualify for other types of employment. Typically these workers were at the bottom
of the hierarchy. Owners fought attempts to unionize their plants, leaving the workers who
had very limited skills in the hopeless situation of "owing their soul to the company store."
Called "piece work," this system continues to be used in some plants. For example, each
shrimp header and oyster shucker is given a container that, when filled with the shrimp tails
or oyster meats, is tallied and at the end of the work day settlement is made based on their
production. It offers flexible working hours and is an advantage for working mothers, for in-
stance. Large plants with production lines for species like salmon and tuna do not operate
on the piece work arrangement.
182 Part Three Fisheries

Figure 9.6. Trout are hand boned by

removing backbone and then rib bones. They
can be packed this way or processed through
the breading machine (in background) that
gives them a coat of batter and cracker meal.
Strictly sanitary conditions in processing plants
are high priority. Photo courtesy of Snake
River Trout Farm, Buhl, rD.

The cold-water shrimp (pandalid) found in northern waters is a good example of a species
that supported a small fishery in Alaska with the potential for expanded production, except
for expensive hand labor. Heading and removing the exoskeleton (called "peeling") was an
extremely tedious job on these small shrimp (120 to 140 heads-on shrimp per pound). In the
years after a peeling machine was introduced in Alaska in 1957, landings rose from 7.9 to
16.9 million lb (3.6 to 7.7 million kg) of heads-on shrimp by 1962.
Because large warm-water (penaeid) shrimp are more valuable than small, processors pay
different prices according to the size landed. Formerly hand graded, machine graders
speeded up the process of grading different sizes of shrimp. Heading (deheading) is done al-
most completely by hand aboard the vessel when large shrimp are caught (Fig. 9.7). Higher
prices are paid at the dock if headed shrimp are landed. Large catches of small shrimp are
delivered to heading houses ashore.

Oyster Processing
Oyster shucking always has been a labor intensive operation and a continuous problem in
hiring, training, and keeping good hand shuckers. Expensive mechanical devices that assist
in processing many species of fish! shellfish greatly expedite production and reduce high
processing costs. There is a great need for mechanical handling and shucking equipment to
replace hand labor at a reasonable cost. Many avenues have been explored, but none has
found commercial application. Oyster meats are usually graded according to size, with
smalls going for soups and the like, 300 or more meats to a gal (3.8 1), and large ones, 160 or
fewer per gal (3.8 1) are breaded or used for raw bars.

Salmon Canning
Crude crosses mark the graves near the old salmon canneries in Alaska of cannery work-
ers recruited from South America and China to process salmon landings. Many markers
show no date of birth or age of the deceased, and often, the only home address is their home
country. During years of heavy salmon runs the need for labor was critical to carry out the
Chapter 9 Food and Nonfood Fisheries 183

Figure 9.7. Shrimp require considerable

processing for different markets. Now a
processing machine automatically peels, tail-on
round, peeled, and deveined, tail-on butterfly,
and tail-on western (split tail). Photo courtesy
of Gregor Jonsson, Inc.

An account of the Chinese cannery labor in Bristol Bay, Alaska was reported in the Alaska
Salmon Investigations 1900 and 1901, U.S. Fish Commission for 1902 by J. F. Moser.
The arrangement for the employment of Chinese is made through the labor agencies of the
large cities, principally in San Francisco. They work under a boss of their own, who guar-
antees each man a certain amount for the season. They do all the work in connection with
the pack. They receive the uncleaned fish at the bins and deliver them canned, lacquered,
and labeled, in cases, at the other end of the cannery. The packing company transports the
Chinese to the field of work and carries them to the home port at the end of the season; it
provides them with a bunk house and furnishes fuel, water, and salt. The boss supplies the
Chinese chow but during the height of the season they also receive some food from the
white mess.
The Chinese contract this year (1900), in Bristol Bay, was 45 cents per case for machine-
filled and 60 cents per case for hand-filled pack. A certain number of cases are guaranteed
in the contract, which must be paid for whether packed or not, and if the pack runs over the
guaranty the extras are paid for at the same rate.
The most arduous work in a cannery falls upon the butchers and fish-cleaners and the
bathroom men. The former are on their feet during the long hours of each day, standing in
slush and gurry, and suffer much with swelled feet and ankles, while the latter are on the
constant move and are the last to clean up at night. The men that work at the lye tanks [af-
ter the cans leave the retorts, where they are cooked under pressure, grease and dirt are re-
moved by a lye bath, and after cooling are ready for lacquering], with the hot sputtering lye
splashing over them, have no sinecure.
184 Part Three Fisheries

At about the turn of the century a machine called the "Iron Chink" was introduced to
dress salmon (Fig. 9.8). In Bristol Bay, Alaska, results with the new machine were so success-
ful and labor saving that cannery managers hurried to order more. The machine is still in
use. Salmon pass along a belt and are deheaded, fins removed, viscera including kidneys are
removed, then brushed to remove blood and membranes. Next, the fish are cut into can-size
lengths for delivery to the filling machines. The machinery can be adjusted to accommodate
different species and different sizes of salmon. The small size variation within a species of
mature salmon is a distinct advantage in machine processing.

Figure 9.8. Salmon canning line. (a) Off-loaded

salmon placed on production line. (b) Salmon
processing machine (Iron Chink) removes heads,
fins, and entrails. (c) Retorting freshly canned
salmon. All photos believed to be taken around the
turn of the century in Alaska and/or Washington
State. Courtesy of University of Washington
Photograph Collections. (c)

Pacific Tuna Canning

The number of workers in a large tuna processing plant is staggering. Because no machin-
ery is available to process these large fish, every step prior to canning must be done by hand.
Although hand processing is expensive, it is justified on an economic basis because of value
and high demand of the product.
Chapter 9 Food and Nonfood Fisheries 185

Crab Processing
Crabs, unlike many other species, must come to the processing plant alive. Crabs are gen-
erally cooked as soon as they enter the processing plant (with the exception of the large spi-
dercrab species). After cooling, the back or carapace is removed enabling viscera, gills, and
mouth parts to be removed and discarded. After the tail flap (that part of the carapace
folded under the crab's body) has been removed, the crab is broken in half. Some crab spe-
cies have very little body meat, but if it is worth the effort, it can be removed with a pick or
by hitting the animal upon a hard surface to break the shell. The desirable leg and claw meat
is removed by cracking the hard exoskeleton with a hammer or by breaking it with pliers.
Care must be taken to prevent sharp exoskeleton splinters from remaining in the meat as
they may harm consumers (Fig. 9.9).

Figure 9.9. One step deboning machine can

be used for any fresh protein, including fish
and shellfish (lobsters, crabs or shrimp). With
the power on, fish/shellfish are simply fed into
the large hopper and wastes, bones, scales, etc.,
are expelled out the opening to the right of the
hopper. Directly below the hopper the cleaned
fish flesh is dropped out of the hopper
machine into a container (not shown). Easy
disassembly and cleaning are important
advantages of fish processing machinery.
Photos courtesy of Stephen Paoli
Manufacturing Corp.

Processing Plant Sanitation

A 1991 U.S. FDA report estimates that there are more than 4,100 seafood processors and
handlers in the United States. Much of the processing industry is old-fashioned compared to
other major food industries. Federal and state agencies exercise limited control over the han-
dling of seafood by seafood wholesalers and processors. Although most seafood is free from
chemical and biological contaminants and disease when caught, spoilage can occur as it
moves from harvest to the consumer's table. Increases in sales in recent years indicates pub-
lic interest in seafoods, by taste and by concern about cholesterol. Seafood quality should be
regulated to provide a fresh-tasting and desirable product; if violations of plant and market
handling of fish and cases of seafood poisoning continue, the market will suffer greatly.
Well-known criteria for good sanitation pertain to plant design, construction, and mainte-
nance: water and waste systems, cleaning and sanitizing methods and materials; and per-
sonal hygiene (Fig. 9.10). Processing plants are required to have stainless steel processing
tables and equipment that must be cleaned on a strict regular schedule. For many years te-
dious hand scrubbing and high pressure water and steam hoses with chlorinated water were
186 Part Three Fisheries



Waste TANK





Filine ....chine

Figure 9.10. Example of equipment arrangement in land-based surimi plant. From Sonu (1986).
NOAA Technical Memorandum NMFS-SWR-OI3. 122 pp. Courtesy of Yanagiya Machinery Works, LTD.
Chapter 9 Food and Nonfood Fisheries 187

the only way to reduce bacterial contamination. Today, chlorinated foam, said to be more ef-
fective than chlorinated water, is sprayed on walls, ceilings, and all equipment in the process-
ing rooms, let dry for about half an hour, then removed with water and wiping. Important
new regulations have been promulgated to improve fish handling; but unfortunately, the
FDA does not have the authority to enforce them, and the industry is left to police itself.


Cases of illness and death resulting from eating seafood are always widely publicized in the
news media, and results in reduced demand that may be temporary or long term, depending
on the severity of the problem. It causes a severe negative effect on the economy of fisheries.
Consumers who experience illness from seafoods may never again consume any seafood
products. Ciguatoxin, prevalent in some areas of the Caribbean (where only 44 to 50% of the
species are safe to eat) can cause serious harm to humans.
At present, demand for seafood far outweighs present catch levels and potential harvesting
levels. Reduced availability of seafood, as described above, reduces the nutritional intake
and can affect balance of payments in countries where seafood is an important export prod-

Mollusks like clams, cockles, oysters, and mussels that live in sewage-polluted waters may
be infected with bacteria and viruses pathogenic to man. Improper handling (again, unclean
equipment, poor personal habits, or inadequate refrigeration) can cause very high bacterial
counts. If no pathogenic bacteria are present, most people will find that recovery is usually
rapid after bouts of abdominal pain, diarrhea, and vomiting. Fish are the most perishable of
all foods and are particularly susceptible to spoilage by bacteria, autolysis (enzymatic action),
and oxidation (rancidity of fatty acids). The most important groups of bacteria that cause
spoilage of seafood and sickness in humans are: Pseudomonos spp., Salmonella sp., and Chro-
mobacter spp. Iced fish can have all three groups present because these bacteria survive and
grow at near-freezing levels of 20° to 32°F (-6.7° to OoC).

Enzymatic Spoilage
An enzyme is a protein produced by fish that functions as a catalyst in living organisms.
The action does not stop upon death of the fish: flavor, tissue color, and texture of the fish are
changed. In herring, the color change caused by this is called "rusting."
Oxidation gives a varnishlike odor. Oily fish such as herring, mackerel, and salmon are
more subject to oxidation than lean fish. Fish generally have more unsaturated oil than land

Scombroid Poisoning
Scombroid poisoning is a disease caused by enteric bacteria, and is found in mackerels and
tunas. It is different from ciguatera (discussed below) in that it is caused by bacterial action
in muscle tissue of the fish after its death, which produces a poisonous substance called
scombrotoxin. According to recent findings, it can be avoided by icing the catch immediately.
The name of this enteric disease came about because originally only tuna and tunalike
188 Part Three Fisheries

fishes, especially mackerel, bonito, and saury, were linked to the disease. Additional studies
demonstrated that at least eight other fish species groups, including such popular species as
dolphin fish (mahi mahO, Coryphaena spp., blue fish, Pomatomus saltatrix, and salmon, Onco-
rhynchus spp., are also vehicles for this disease. Bacteria in the fish flesh use an enzyme, his-
tidine decarboxylase, that produces a histamine. Certain conditions usually are required to
produce high levels of histamine, including improper shipboard handling of fish, poor stor-
age facilities, and temperatures over S9°F OsoC). The extent of poisoning can be much more
severe if the fish is also spoiled before it is eaten. Symptoms usually appear a short time af-
ter ingestion and may include some or all of these symptoms: cutaneous rash, edema, inflam-
mation, gastrointestinal disturbance (nausea, vomiting, diarrhea), hypotension, headaches,
palpitation, and itching.


Natural poisons should not be confused with ptomaine poisons produced by bacteria in de-
caying flesh.
Ciguatera usually occurs in reef or shore fish
and is apparently caused when fish feed regularly
on a dinoflagellate (protozoan), Gambierdiscus toxi-
cus, that accumulates the poison in their flesh (Fig.
9.11). When these fish are eaten by other fish the
poison becomes more concentrated. It is quite
possible that fish from one area of a reef are poi-
sonous, but not from another nearby area. About
300 species of fish have been implicated in this
type of poisoning, most of them popular food fish. Figure 9.11. Humans dining on fishes who
This form of fish poisoning is treacherous because have consumed Gambierdiscus toxicus, a
there are no outward signs or symptoms by which dinoflagellate (protozoan) suffer with cigua-
to distinguish poisonous from nonpoisonous indi- tera poisoning. From Steidinger and Joyce
viduals (Fig. 9.12). The toxin appears to have a (1973). FL. Dep. Nat. Resour. Mar. Res. Lab.
cumulative effect on patients receiving repeated Educ. Ser. 17.
Puffer fish poisoning. Fish causing puffer fish poisoning are in the family Tetraodontidae
(blow fish, balloon fish, or tetraodon; Fig. 9.13). They are small, and lack pelvic fins. Their
most obvious characteristic is an ability, when disturbed, to inflate themselves to a spheroid
form almost twice their size. The toxin, tetraodotoxin, is concentrated in certain organs and
tissues; ovaries are most always toxic, and to a somewhat lesser extent, the liver, bile, skin,
and flesh. Despite the danger of eating these fish, called Fugu, the Japanese are fond of them.
Specially trained cooks who know the proper species to serve and how to butcher them re-
duces the chance of poisonings, yet Japanese deaths annually attributed to eating puffers var-
ies from a few to in the hundreds. Eating it is a gamble. The toxin, it is said, in mild doses
provides a feeling of happiness, warmth, and well-being; and, as one person reported to his
wife shortly before his death, he felt like he was "walking on a cloud." For many years it was
believed that puffer fish manufactured their own poison. Now species of commensal bacteria
are thought to be the cause. Because it is a very strong poison known to cause fatalities, eat-
ing puffers should be avoided unless the consumer is absolutely certain of the possible con-
Chapter 9 Food and Nonfood Fisheries 189

(a) (b)

Figure 9.12. Marine reef associated fish that

cause ciguatera in certain locations. (a) Great
barracuda, Sphyraena barracuda, usually more
toxic over 4 lb (1.8 kg) From Goode (1884).
U.s. Commission of Fish and Fisheries.
Washington 840 pp. (b) Greater amberjack,
Seriola dumerili. (c) yellowfin grouper,
Mycteroperca venenosa. (b and c) From Fischer
(1978). FAO Identification sheets for fishery
purposes. Western Atlantic (fishing area 31).
Rome. Vol. 5

Figure 9.13. Species of puffer fish, Genus Fugu, from Japanese waters are hazardous to eat. From
Halstead (1965). Poisonous and hazardous marine animals. Vol. 2. U.S. Printing Office.


Filter-feeding mollusks may accumulate toxins from some phytoplankton species and, when
eaten by man, can cause serious illness. Paralytic shellfish poison (PSP), also called mussel
or clam poisoning, paresthetic poisoning, or mytilitoxification is caused by eating shellfish
that have been feeding on certain species of dinoflagellates. Symptoms usually occur in less
than an hour: tingling lips and tongue, numbness in legs, arms, and neck, lack of muscular
coordination; weakness, vomiting, diarrhea, dizziness, light headedness, incoherence, head-
aches, respiratory distress, and muscular paralysis. Death may occur in 3 to 12 hours.
Other diseases caused by eating toxic shellfish are called, variously, neurotoxic shellfish
poisoning (NSP), diarrhetic shellfish poisoning (DSP), and amnestic shellfish poisoning
190 Part Three Fisheries

(ASP), and are all caused by shellfish eating toxic phytoplankton. In 1987, an outbreak of
ASP in mussels from near Prince Edward Island, Canada poisoned 129 people and caused
two deaths. Among the neurologic responses
are memory loss and disorientation, and, in se-
vere cases, death.
Many cases of poisonous marine fish and
shellfish have been recorded from tropicalloca-
tions other than the southern United States, the
Gulf of Mexico, the West Indies, and the Baha-
mas. For some families of poisonous fishes in
these waters there are no valid records show-
ing that they have caused human sickness or
death, but because fishes in these families are Eriphia sebana
poisonous elsewhere, they are presumed to be
poisonous in the United States. They occur
worldwide in both temperate and tropical wa-
Mollusks may not be the only shellfish
group to contain naturally produced toxins.
Recently, the viscera of Dungeness crabs was
found to contain accumulations of domoic acid
that was also responsible for the 1987 outbreak
of ASP in Canadian mussels referred to above.
Poisonous crabs inhabit coral reefs and rocky
substrates and range widely throughout the Platypodia granulosa
tropical Indo-West Pacific, from the Red Sea
and East Africa to Hawaii and Tahiti (Fig. 9.14).
Although not usually marketed, deaths from
eating them have been documented.
It is important to note that the lack of au-
thentic records simply does not, of itself, mean
that an unfamiliar marine species is safe to
handle or eat, for the following reasons:

1. The relationship between eating poisonous

Demania aicala;
marine organisms and the illness is not
always recognized. Figure 9.14. Highly toxic crabs from the
2. Identification of the sea life causing an ill- tropical Indo-West Pacific reefs. From Garth
ness might be incorrect. and Alcala. (1977). In Proceedings of the
3. Illness caused by marine animals is fre- Third International Coral Reef Symposium.
quently not reported to health authorities. Vol. 1. pp. 646--651. University of Miami.


Fortunately for humans, most parasites that infect fish and shellfish are harmless because the
microenvironment in humans, the digestive tract, does not provide the essential environmen-
tal requirements for parasites to excyst, metamorphose, and reproduce. For parasites to in-
fect and live in a host, they must be able not only to overcome the physical and chemical
Chapter 9 Food and Nonfood Fisheries 191

defenses of the host, but under specific conditions derive substances from the host for sur-
vival and development. Health damage to the host depends on the species of parasite, its
size, numbers present and impact on its microhabitat (human flesh and organs).
Unfortunately for humans, several marine parasite species, if given the chance, will infect
them with possible pathological consequences. Those that infect and harm us are usually
parasites that have fish as an intermediate host, and an adult marine mammal (such as por-
poises or seals) as a final host. The importance here is that the environment in the intestinal
tract in large aquatic mammals, or land animals, such as bears, parasitized by worms from
salmon in freshwater rivers and streams, are roughly similar to those in man; whereas, larval
tapeworms found in fish such as sea trout, have sharks (not man) as final hosts, where they
find adequate conditions for transformation into adults.
Parasitism is a widespread, natural phenomenon in nature. We can prevent and control
the level of human infection by parasites from domestic animals used for human food; but
there is little that can be done to prevent wild fish from getting parasites, so public health
protection must come after capture. Cultured marine fish and shellfish food (aquaculture)
normally will not contain as many parasites as their wild relatives because the ecosystem
they depend on, including the presence of intermediate hosts found in nature, is not present
in controlled aquaculture facilities. Artificial feeds in these facilities substantially reduce
chances of infection.
Taste preferences for fish in North America range to extremes, from ''blackened'' to raw.
The latter may be partly due to the popularity of Japanese restaurants that serve fish raw,
lightly cooked, or smoked. Japanese have eaten sushi and sashimi for a long time, but until
a few years ago, it was new to Americans. This change in dietary habits has its down side;
in the United States there were 50 reported cases of parasitic infections from eating improp-
erly prepared marine fish since 1980. Seafood is excellent fare for humans, but some simple
precautions must be taken to avoid possible health dangers. Cooking and freezing usually
destroys parasites. It is important to note that microwaving for brief periods may not kill
parasites of possible harm to man, and can contribute to the increase in this disease.
Because only certain marine fish have parasites potentially harmful to man, the health haz-
ard can be avoided by cooking, not eating it raw. Some of these fish are popular, such as
salmon, herring, and cod, and this is to the worm's advantage. Parasitized fish that may be
potentially dangerous to eat may be misidentified, while others may completely lose their
identity after filleting and passing through market channels. Many species eaten raw in such
dishes as sushi and sashimi are pelagic fish like tuna. When served in quality restaurants,
they are known to be safe. It has been suggested that more parasitic worms are now present
in fish, and therefore infect more humans. Doctors from the University of California and the
U.S. FDA speculate that this increase may be due to the protection given dolphins, sea lions,
and seals by the 1972 Marine Mammal Protection Act that brought increases in abundance of
mammals that serve as final hosts. Marine fish parasites important from a human health
standpoint include the "larger" or macroparasites commonly called "worms," round worms
(nematodes), and one type of flat worm, a tapeworm (cestode). Another type of flatworm
(trematode) is also present in marine fish and may infect humans.
The parasites discussed in this section are normally found encysted in the flesh of fish.
Parasites, of course, are found in fish in all organs and organ systems but because humans
normally eat only the flesh, many parasites in other organs are of no concern.
All of these worms have complicated, indirect life cycles. This means they parasitize fish
and invertebrates that serve as intermediate hosts from the egg stage until they transform
into adult worms and move into a final host, where they spend the remainder of their lives.
192 Part Three Fisheries

Humans become infected when they eat larval parasites encysted in the intermediate host
flesh that is either uncooked, lightly cooked or smoked, or has never been frozen.


Roundworms (Nematodes)

Marine finfish are frequently infected with larval round worms (Fig. 9.15). They are elon-
gate, cylindrical worms tapering at each end, and they may be colored white, yellow, brown,
or red. Not all worms in this group are harmful to man, but because knowledge of minor
morphological differences between species is required for identification, it is wise to assume
that any round worm found encysted or free in a fish may be a potential human health haz-
ard. They are most commonly found in gadoid fish (cod and haddock), and herrings, but in-
fect other fish as well. A case of a live fish nematode in a human was recorded as early as

@ Figure 9.15. Life cycle of a roundworm
(nematode). In addition to the final host cod,
roundworms infect many other finfish
including herring. Roundworms in uncooked
herring eaten by man can cause diseases in
man. From Uspenskaya (1955). Zoo1. Zh.

The green herring in The Netherlands is an example of potential risk from nematodes.
People there are fond of this fish that is sold lightly salted. If gutting and salting is delayed
until the fishing vessel reaches port, human infections with roundworms can result. It has
been found that roundworms can move from the viscera to the musculature during storage
on ice, and the light salting that follows does not harm the worms. Such movement does not
occur in all species infected by this roundworm, but it is known to also occur in the mack-
erel. Patients who have unknowingly ingested these live roundworms usually complain of
gastrointestinal pains and stomach disorders similar to those from food poisoning and ulcers.
However, the cause of distress can be identified when the patient coughs up or vomits a live
nematode. Sudden stomach pains, nausea, and vomiting occur when the worms penetrate,
or attempt to penetrate, stomach tissue. If the pains and distress occur within a few hours
after eating raw seafood, nematode infection in the stomach should be suspected. Symptoms
of roundworm infection in the intestines are similar to other intestinal disorders, but may
take much longer to affect the patient, sometimes up to several weeks after the meal of in-
Chapter 9 Food and Nonfood Fisheries 193

fected improperly prepared fish is eaten, which may obscure the cause and effect relationship
between the fish eaten and the disease.

Tapeworms (Cestodes)

Tapeworms are more familiar to people than are round worms; they have a structure
called a scolex, used for attachment to the host, and a long chain of segments (Fig. 9.16). Fish
tapeworms, long recognized as human pathogens in northern Europe, recently have become
common in North America. Humans in western Canada and Alaska become infected by eat-
ing raw or undercooked salmon where larval worms are encysted in salmon flesh, usually
sockeye salmon, Oncorhynchus nerka, that as young fish become infected during their long pe-
riod in freshwater. The segments, called proglottids, are passed in the feces of the final host.
The first intermediate host is a small freshwater crustacean, a fish eats the crustacean: moves

= _.",
_ _ DlllllB_@Iiloo


Eaa (UId .aolHtl1DU
a.HI puaed L.a. ,.'"
Man ,. lafectld .",
ofI.l1. raw G, I mflra~ rlJ'
..... .-.
coo);tcI (fiil!! 'WM<:h .,.



URVA In. ralltd ... MATURI I.OQ
Itoal.c: ...... nl......
a coree 1l • •

to Illl.U(:lu .lIoen it boK:omes ..

IrCfll.\ , .aalmr;,n. pln

mleroc:raatacan I ..eattd bf 1I1h.


Figure 9.16. Life cycle of Diphyllobothrium latum, tapeworm (Cestoda). When mature, they infect
intestines of humans. From U.s. Naval Medical School.
194 Part Three Fisheries

to sea, where the final host in the ocean is a mammal, seal, or sea lion. Bears become in-
fected when eating spawning salmon in rivers and streams, then defecating proglottids con-
taining tapeworm eggs back into the stream. These are then eaten by crustaceans that, in
turn, are eaten by young salmon. These worms may become adults in humans and can reach
a length of 30 ft (9 m)!

flatworms (Trematodes)
Another flatworm, the trematode (also called "fluke"), lives in salmon and causes human
infections when the fish is insufficiently cooked (Fig. 9.17). Cases have been recorded in Rus-
sia and the west coast of the United States, from northern California to Washington. Abdom-
inal pains and diarrhea are possible symptoms, but occasionally none will appear. As with
cestodes, eggs in the feces of a patient signal the presence of the parasite.

-:...- Worms migrate to

tissues (usually the
Jlf~n is infected by ~.~:7J~1-"m::-:- lungs) and mature.
eating uncooked crab.

in the


" ...
" . t" .~,.t!

~ ~-~ . ~'V: -
Snail •• .

~ Figure 9.17. Example of the kinds of fish and
shellfish parasites of harm to man. Life cycle
which infect fresh·water crab. ~ ~ --4!Jl.-- of a trematode (Digenea). From Noble and

lata emerges as cercariae ••• I Noble (1982). Parasitology: The biology of

animal parasites. Lea & Febiger, Philadelphia,


There are records of parasitic infections in people who have eaten raw crustaceans, shrimp,
prawns, or crabs, animals most Americans do not eat raw. Crustaceans are hosts for worms
similar to those in finfish and could become potential human health problems; for example,
a larval nematode present in warm water shrimp (penaeid shrimp) from the northern Gulf of
Mexico can infect such laboratory animals as mice or monkeys. This suggests that man could
be a possible host, but research results in this area are skimpy.
Chapter 9 Food and Nonfood Fisheries 195

The incidence of infection of humans by fish-borne parasites is extremely low in the

United States population, where cooking is a traditional preparation method. Yet, a threat
does exist. Care is strongly recommended to avoid infection.


Some people have an allergy to mollusks called "erythematous shellfish poisoning." This is
manifested by rashes (diffuse erythema), urticaria, swelling, and stomach or intestinal disor-


Occasionally the news media report human deaths resulting from eating certain canned
foods, including some seafood. The cause of deaths (in some cases) is a spore-forming bac-
terium, Clostridium botulinum, that thrives in low oxygen levels and releases a deadly toxin in
smoked and canned foods. It normally does not cause an off-taste. The toxin formed by the
botulinum, the direct cause of botulism, is called botulin.
While deaths caused by botulism have not been many, 978 deaths in United States from
1899 to 1973, sensational news coverage occasions concern and loss of confidence in the food
canning industries. Reestablishing public confidence may take 2 years or more, which can in-
flict serious hardships on the industry. Most historic outbreaks come from home-canned
foods; only about 9% are from commercial products. This is small considering the output of
the canning industry in this country. The worst offenders are brine packed, smoked, pickled,
and pasteurized products (semipreserved) wherein the spores of C. botulinum could be unaf-
fected by processing, and, under suitable conditions, grow. Unfortunately, there is usually no
obvious sign of spoilage in the canned product to warn the consumer. Of the seven strains
or types of C. botulinum, only one is seafood related, and that is found in products that can
be eaten directly from the can at room temperature, including clams, clam juice, crabs,
salmon, sardines, sprat, tuna, and smoked white fish.


Certain diseases associated with seafood may seriously affect the industry. Cholera, a bacte-
rial disease, Vibrio cholerae, is such a disease. Infected humans vomit and experience violent
diarrhea that dehydrates victims usually within hours of infection. Untreated, cholera may
cause death through shock, kidney collapse, and circulatory failure. The disease is transmit-
ted through contaminated foods, usually seafood and unwashed vegetables. It is a mass
An outbreak near Lima, Peru, that began late in January 1991, affected 70,000 people along
1,200 mi (1930 km) of coastline. Health ministers in Peru advised against eating coastal
fishes and drinking unboiled water. Imports of fish from Peru dropped dramatically when
countries that normally import Peruvian fish and other goods cancelled orders. Some of
them that had received Peruvian imports burned them.
196 Part Three Fisheries


The objective of Fish Protein Concentrate (FPC) development is to provide needed protein to
the world's hungry. FPC is manufactured by removing moisture from fish by a solvent ex-
traction process. Since 1900, scientists have attempted to concentrate the protein in fish to
supplement low protein diets. As world hunger and human populations increased in devel-
oping countries, the need to provide better diets has becomes more critical. FPC research ac-
celerated in the 1930s and continued into the 1970s.
The process begins with locating large populations of fish or shellfish safe for human con-
sumption yet not presently utilized, that are inexpensive to harvest and available for ex-
tended periods of time. Liquid is removed, then the remainder is dried and ground up to
make a product that does not need refrigeration, can be easily stored, and is convenient to
use. Because of its appearance and use, it has been called fish flour.
FPC never achieved the success envisioned in the United States. Fishery scientists were una-
ble to locate fishery resources that fitted the criteria mentioned above, and problems of extract-
ing liquids from the fish proved complex and were never fully solved. As to the future role of
FPC in reducing international malnutrition, suggestions have been made that it would be better
to encourage persons in countries where protein is lacking to use fish that can be processed
cheaply by local technology and be acceptable to local consumers. Today a type of FPC is pro-
duced in Norway and India that avoids the problem of extracting fish liquids.


Another type of production from fish and shellfish fisheries is feed for farmed animals that
man eats. Products from this type of industry are considered "feeds," and should not be con-
fused with food fisheries. Fisheries that process these species are referred to as industrial or
reduction fisheries. Some fisheries also produce for recreational fishermen (Fig. 9.18).

Figure 9.18. Packing sand worms in Maine

for shipment to bait dealers catering to
recreational fishermen. Photo courtesy Ivan
Chapter 9 Food and Nonfood Fisheries 197

A wide array of diverse products come from the sea and enrich our lives. Some uses are,
or have been, practical and others simply luxuries. N. Karrick (1971) pointed out the variety
of nonfood products that come from water bodies and the animals that live there.
Resources from the water world have fed man's pets; have illuminated and heated his
home; helped to paint his factories; furnished strong adhesives and also fine glues for deli-
cate photoengraving; and helped to produce plastics that are in themselves used for many
Aquatic resources have furnished man with pearl, coral, and tortoise-shell jewelry for
adornment; shark, alligator, and salmon skins to make leather; sepia for his favorite oil
painting; isinglass to clarify his favorite wine; mother-of-pearl to add beauty to inlaid art
objects or to such practical things as trays or cutlery; walrus ivory for carved objects; amber-
gris from whales to make the best perfumes; fish scales to make artificial pearls; mussel
shells to make buttons and buckles or to use in the culture of pearls; luxury furs; salt for his
food; sponges for cleaning; bromine to make ethyl gasoline; shells ground for roadbeds or
railroad beds or for lime; and lubricating oils for fine watches and instruments.

Some of the products have been synthesized, or substitutes found when the wild caught
animals became scarce and too expensive. A good example is the fishery for the sperm
whale, sought by whale fishermen, that produced oil for the lamps of London and New York.
Demand for whale oil declined in the late 1850s when the world's first successful oil well
was drilled in Pennsylvania. Around 1940, the soupfin shark (Galeorhinus zyopterus) fishery
in California was producing large and oily livers containing vitamin A that were used to for-
tify poultry feeds. The production of vitamin A in 1950 developed after shark stocks were
threatened with extinction, and the fishery ceased operations.
A few examples of non-food products from the sea are discussed briefly below.

Jewelry Industry
Marine fisheries for jewelry are small in terms of weight landed, but have a very high
value per unit of weight. Shell jewelry is made from the inner mollusk shell layers called na-
cre; it is that hard, iridescent material, often called mother-of-pearl. Nacre is found in conch,
Strombus gigas, and abalone, Haliotis sp., and some other mollusks. Jewelry and other items
also have been made from shells (carapaces) of sea turtles, including green sea turtle, Chelonia
mydas, now an endangered species.
Pearls from pearl oysters, Pinctada martensii, formerly were obtained only in Japan for the
natural pearls they might contain. In 1894, Kokichi Mikimoto developed techniques for cul-
turing pearls and dominated the world pearl culture industry.
Precious corals used for the commercial jewelry market can be pink or red, Corallium spp.,
and some of lesser market value are gold, black, and bamboo. Beginning in the early 1980s
about half of the world's supply of Corallium spp. was harvested from the Emperor Seamount
in the northwestern Hawaiian Islands, at depths of 1,312 and 15,533 ft (400 and 4,734 m). To-
day, in U.S. Pacific Island possessions, fishing for corals is permitted only by regular or ex-
perimental permits. In the Mediterrean Sea, and off Hokkaido, Japan, precious corals also
have been found on seamounts.


Production of fish meal and fish oil began in North Europe early in the 19th century. Herring
fisheries occasionally harvested surplus catches, and is believed to have been the basis of an
198 Part Three Fisheries

early industry that eventually spread worldwide. The largest producers are Japan, Chile, and
Peru. A trend has emerged where production units are larger and are concentrated in fewer
plants. Prices of fish meal and fish oil vary greatly and depend on prices of soya protein and
other oils, and on the price and supply of raw fish.
The meals used in diets of poultry and pigs has a vitamin content of about 70 percent.
More recently, farmed salmon, trout, and mink have been fed large quantities of fish meal.
Fish oils are used by humans in margarines, shortenings, and compound fats.
Large numbers of small oily schooling fish are the raw material for these industries. Oily
fish tend to deteriorate rapidly after capture and the yield of meal and oil is reduced, so low
temperatures must be maintained in the tanks when rapid delivery to the reduction plant is
not possible.
At the reduction plant, oil and solids from the fish are separated, and the water removed
by evaporation and drying. The fish meal and fish oil are stabilized. The basic process has
changed little over the years except for modernization of equipment to reduce energy con-
sumption, improve quality of the product, and satisfy environmental guidelines. Fish meal
plants, notorious because they emit fish odors in enormous volume, have been required to
use scrubbers or afterburners to reduce this nuisance as much as possible. Plants are usually
located close to shore to expedite handling and are generally located in areas of low human
population density. Reducing odors and polluted wastewater becomes especially important
when the plants are in higher density areas.
A product called "stickwater" or "press water" released from cooked fish pulp contains a
rich source of the B complex vitamins and other growth-promotion factors, and is now recov-
ered from the process and preserved.

Four species of menhaden are found in North American marine waters: Atlantic men-
haden, Brevoortia tyrannus; Gulf menhaden, B. patronus; yellowfin menhaden, B. smithi; and
fine scale menhaden, B. gunteri. Members of the genus range from coastal waters of Ver-
acruz, Mexico, to Nova Scotia. Those in the Atlantic and Gulf are very abundant and sup-
port large purse-seine reduction fisheries. Menhaden are filter feeders and are estuarine
dependent. Their life span is about six years.
When the Colonists arrived in New England, the Indians were using Atlantic menhaden
for fertilizer. They called the fish "Munnawhatteaug," meaning, sensibly, "that which ma-
nures." Elementary school children learn that the Indians taught Pilgrim colonists to fertilize
their crops with fish, and by 1612, beach seines were being used to catch menhaden from the
large nearshore schools.
Small-scale crude oil industries using menhaden are believed to have started along the
coast in Rhode Island in 1812, and in Maine about 1850. Operated part-time with crude ma-
chinery, this shore-based industry graduated to a factory-type operation when screw presses
were invented. By 1870, steam was used to cook the fish and power the machinery for re-
moving the oil. The 10 year period from 1865 to 1875 brought rapid growth of the menhaden
industry. Processing machinery was sent out on barges about 1875 to avoid the long, slow
runs by fishing vessels to the shore-side plants. Later, higher powered vessels made delivery
to land-based plants quick and easy, and signaled the decline of the floating processing
plants. When companies began using fish oils in paint in 1812, the menhaden market im-
proved considerably.
The New England states had the highest menhaden production until 1875, when stocks
Chapter 9 Food and Nonfood Fisheries 199

disappeared from north of Cape Cod. The industry followed the fish south to New Jersey
and Virginia. The collapse of the New England fishery stimulated harvesting the menhaden
stocks in the Gulf of Mexico. First landings from this new fishery were made about 1900 on
the west coast of Florida and in Texas.
Following World War II, the menhaden industry became more firmly established and vita-
min B 12 was found in fish meal. Between 1950 and 1959, catches for both the Gulf and At-
lantic menhaden increased, with the greatest from Atlantic stocks. By 1959, an annual record
catch of over 1 million tons was landed.
Menhaden purse seining formerly involved backbreaking handhauling of nets by teams of
fishermen in seine skiffs. About 1958, hydraulic power blocks for hauling seines became
available (see Chapter 8), and lighter, more durable synthetic webbing replaced cotton in the
seines. Fish pumps sped up loading and unloading, and refrigeration ensured that better
quality fish were delivered to the plant. Locating menhaden schools by spotter aircraft began
in 1950 and greatly increased the fishing pressure on the stocks; setting the seine is directed
from the air by radio communication.
Uses of menhaden have increased as have the catches. Fish solubles and fish meal for
poultry and livestock feeds are the primary use, but additional uses in the manufacture of oil,
for curing leather and tempering steel, in soaps, paints, and varnishes, caused the menhaden
industry to flourish. Attempts to sell fresh and canned menhaden as human food have


Nonfood fish in pet foods has become an important industry since 1955. Canned fishery
products used as pet food are valued at about $50 million annually, yet probably no more
than half the potential market has been tapped. The industry is constantly looking for a sta-
ble, large, and inexpensive supply of fish. Many marine and freshwater species are being
Animal feeds (pet food) are canned in New England and California, with production
about equally divided between the two areas. On both the Atlantic and Pacific Coasts, ani-
mal food is canned as a by-product, utilizing material that otherwise would be wasted. In
New England, pet food consists of filleting waste. In California, whole sardines (pilchards)
or mackerel, fresh and of good quality, unsuitable for canning because of size, are used.
Whale and seal meat were also utilized in pet food there. Most of these early pet foods were
purchased in large sized containers by kennel operators, fish hatcheries, and "fur farms."
The first commercially prepared dog food, a biscuit, was introduced in England about 1860.
Dry dog foods were subsequently developed, with formulas based on the nutritional knowl-
edge of the day. Horsemeat was canned for dogs after World War I; improvements came over
the years. The 1960s were marked with great diversification in the types of dog and cat food
available to pet owners, offering more varieties of canned and dry products. With the growth
of the industry has come a greatly expanded use of by-products from the meat, poultry, and
seafood processing industry using approximately 1.1 million tons of these by-products annu-
ally in pet foods, providing an important source of income to the processors.
Pet food production and sales increased about 73% between 1952 and 1963, and at the con-
sumer level, sales amount to about $500 million per year. In 1958, $485 million was spent for
pet foods, of which $220 million was for canned dog food, $192 million for dry dog food, $42
million for canned cat food, and $31 million for bird food. This volume involved the utiliza-
200 Part Three Fisheries

tion of about 12.5 billion lbs of varied food products, much of which is often regarded as
waste. The future growth of this industry may be even more rapid than that of the past 10
years, because it is estimated that only 23% of the food presently consumed by the canine
population is commercially produced pet food. The dog population, furthermore, is increas-
ing at a more rapid rate than the human population, so the need for products high in pro-
teins and amino acids will undoubtedly become greater. Trimmed red meat from tuna, also
called "blood meat," saved in early operations, along with cleaning scrap, was formerly used
for reduction. It is now being canned for pet food with the occasional addition of vitamins,
minerals, and smoke flavoring. Canned cat foods contain nearly 100%, while dog foods con-
tain substantially less. When the United States was still a whaling nation in 1963, whale
meat was sold as mink feed in fur farms, for zoo animals, and was also used in canned pet
The practice of using certain portions of fish waste to process hatchery fish food would not
have become as important as it has had not the pellets of dry hatchery feeds been developed.
About 1970, when the nutritional requirements of rainbow trout were determined by the U.S.
Fish and Wildlife Service, trout feeds were improved greatly.


Macroalgae are the large plants growing in aquatic environments, freshwater, saltwater, and
brackish water. The term "seaweed" is often used for the large marine algae, many of which
are called "kelp." These plants are classified by botanists according to several characteristics,
including color; some are blue-green, some brown, and some red, the latter two marine algae
being the most valuable commercially.
Many species of algae occur along the sea coasts, mostly in the intertidal zones. In Can-
ada, about 300 species grow along the Atlantic coast.
Commercial algae, with few exceptions, is gathered in public areas. Several species are
used as thickening agents in many processed foods, such as soups and ice cream, and are nu-
tritious foods in their own right. One species of red algae, dulse, consists of about 25% pro-
tein, about 44% carbohydrates, and 27% mineral salts. Agar, algin, and carrageenan are
important algae products. Agar, a gelling agent, is useful in the home and in the laboratory.
Algin, used in ice cream and chocolate milk drinks, has other uses in manufacturing and in
Carrageenan is an emulsifier used in food items such as dairy products, puddings, and a
number of nonfood items such as toothpastes, beer, pharmaceuticals, water-based paints, in
drugs such as cough syrups, dental impression material, in cosmetics such as shampoos, and
industries such as paper, paint and rubber. These are but a few examples. This industry is
valued at $30 million in the United States. As much as seven times the production in the
United States of some seaweeds are imported to meet the demands.
Most algae contain important minerals and vitamins valuable to the human body. Irish
Moss, Chondrus crispus, is a small perennial marine red algae about 10 in (25 cm) high that forms
a thick carpet over rocks and ledges. It is found along the east coast of North America from
Nova Scotia to Rhode Island, and the coasts of Scotland and Ireland. Growth is slow, only a
tiny fraction of an inch per day during the summer. Growth in suitable conditions where tem-
peratures are 41 ° to 59°F (5° to 15°C) is considerably faster than in the wild. Weight increases as
high as sixfold per month have been achieved with low density trials. Growth results from the
increase in the number of plants, rather than in the production of large plants. Irish moss grows
Chapter 9 Food and Nonfood Fisheries 201

by photosynthesis, extracting nutrients and minerals from shallow water. Plants can absorb
enough nitrogen in a few hours to carry them for several days.
An extract from Irish moss, carrageenan, is widely used in food processing and other in-
dustries. Harvesting is done by raking attached plants loose from underwater beds, or by
simply collecting plants that wash up on the beach. Irish moss will also grow detached;
there are special strains that lack holdfasts. Labor demands are generally low.
Another important algae is the kelp, Macrocystis pyrifera, a large, red algae that may attain
a height of 100 to 150 ft (30 to 46 m). It ranges from central Baja, California to Alaska in cold
nearshore waters, down to 150 ft (46 m). The large fronds have flotation devices to keep
them near the surface of the water that enable them to photosynthesize. A giant algae, kelp
has special reproductive leaves that produce spores that are released into the water where
they swim about before settling to the bottom. The spores grow into tiny plants that produce
eggs and sperm that unite to grow into adult plants. They extract nutrients from the water
for photosynthesis. A full-grown plant may be 6 years old. Kelp are eaten by sea urchins,
other echinoids, gastropods, and crustaceans.
When the fronds near the surface of the sea are harvested, they regenerate so that contin-
uous production can be realized. The plants yield several commodities: additives for human
food, fertilizers, stabilizers in plants and drinks, and alginic acid. Years ago kelp was an im-
portant source of both potash and acetone.


Medicinal properties of ocean animals have been

known for thousands of years, yet little systematic
research has been done. Suggested reasons for
this are that until the 1960s this was a new field
where no trial and error had been done, as on ter-
restrial plants, to reduce human diseases. Asian
countries have identified some marine species that
they claim alleviate human illness or suffering,

and these found a place in their folklore. The
ocean's hostile environment prevents gathering of
different species living at different depths that
might produce useful drugs. The advent of un-
derwater breathing devices and manned submers- C.
ibles brought opportunities for easier collecting;
but sampling the enormous variety of species in
the ocean with no clue as to which might be use-
ful made collecting and testing expensive. Dis-
tance of testing laboratories from collection sites
cause additional problems when assays have to be
conducted on partially decomposed specimens.
In the late 1960s, interest by pharmaceutical Figure 9.19. Examples of the variety of
companies promoted research and testing of ma- aquatic organisms that been tested for pos-
rine plants and animals to develop new therapeu- sible production of drugs useful to man.
tic agents (Fig. 9.19). The work required obtaining From Storr (1957). Florida Board of Conser-
extracts to be tested for biological activity, antibac- vation Educational Series No.9.
202 Part Three Fisheries

terial properties, and potency against a wide spectrum of human diseases including cancer,
viral infections, fungal disease, and disorders of the immune system. Active components of
the extracts were identified chemically to determine their structure for possible further test-
ing and synthesis. Testing was expensive because potential drugs had to be first tested on
laboratory animals to be certain that there were no adverse side effects. If the extracts passed
this stage, they could be tested on humans. Development of a new drug can cost millions of
dollars before a drug company can apply to the U.S. FDA for marketing approval. Some
early trials that encouraged more research included sponges, common quahog clam, Merce-
naria mercenaria, extracts that showed antibiotic capabilities, and sea cucumber extracts that
inhibited tumors in mice. The list of animals that can be tested for use by pharmaceutical
companies for extracts is lengthy.
There have been many blind alleys in the
search for valuable drugs from the sea. Some-
times their value becomes apparent during pre-
liminary testing; others take much longer to
evaluate and have a higher research price tag.
One success story comes from the lowly Jamai-
can gorgonian, Plexaura homomalla, that produces
prostoglandins, a relative of the human hor-
mones involved in regulating many body func-
tions at the cellular level. The blood of
horseshoe crabs, Limulus polyphemus, gave rise to
a test whereby their reconstituted dried blood,
when exposed to bacterial endotoxin, forms a
gel. This test is used worldwide (Fig. 9.20).
Near the end of the 1970s, interest in drugs
from the sea diminished substantially, although
now and then new claims are made for marine
pharmaceuticals. A new possible cure for cancer
that has been promoted is made from shark car-
tilage. Cartilage is sterilized in vats of boiling
chemicals and ground into chips, dried, and put Figure 9.20. Horseshoe crab, not a crab
on the market in pill form. Preliminary tests but a relative of spiders and scorpions.
made on 15 cancer patients in Cuba resulted in From Stewart (1980). Geojourney pp. 14-15.
signs of reduction in the size of cancer tumors
for three patients. Shark fishing is being observed vigorously by conservationists in Costa
Rica who are concerned that shark stocks will be driven to extremely low levels if cartilage is
found to be an effective cancer treatment. The concern that sharks stocks will be overfished
is justified, because sharks mature late in life, are slow growing, and bear few offspring.


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Food and Agriculture Organization of the United Nations. FAO Yearbook of fishery statistics. Com-
modities (Published annually).
Gulland, J. A. (ed.). 1970. The fish resources of the ocean. FAO Fish. Tech. Pap. (97) 425 pp.
Gutherz, E. J., G. M. Russell, A. F. Serra, and Bennie A. Rohr. 1975. Synopsis of the northern Gulf of
Mexico industrial and foodfish industries. Mar. Fish. Rev. 37(7):1-11.
Harvey, M. J. and J. McLachlan (eds.). 1973. Chondrus crispus. Halifax: Nova Scotian Institute of Sci-
Idyll, C. P. 1973. Fish meal and FPc. Sea Frontiers. Int. Oceanogr. Fndn. 19(2):83-91.
Kent, G. 1987. Fish, food and hunger: The potential of fisheries for alleviating malnutrition. Westview
Press, Boulder and London.
Mitsui, A. and C. C. Black (eds.). 1982. CRC handbook of biosolar resources. Vol. 1, Parts 1 & 2, Basic
principles. CRC Press, Boca Raton, FL
Shapiro, S. (ed.). 1971. Our changing fisheries. U.S. Department Commerce, NOAA/NMFS.
Stansby, M. E. (ed.). Fish oils in nutrition. Van Nostrand Reinhold, New York.
204 Part Three Fisheries

Stansby, M. E. (ed.). 1976. Industrial fishery technology. Robert E. Krieger Pub!. Co., Huntington, New
York. (Second Edition).
Walford, 1. A. 1958. Living resources of the oceans: Opportunities for research and expansion. The Ro-
nald Press Co., New York.
Wheaton, F. W. and T. B. Lawson. 1985. Processing aquatic food products. John Wiley & Sons, Inc.,
New York.
Yarish, c., C. A. Penniman, and P. Van Patten (eds.). 1990. Economically important plants of the Atlan-
tic: Their biology and cultivation. Connecticut Sea Grant College Program, Groton, CT
Youngken, H. w., Jr. 1970. Food-drugs from the sea. Proceedings 1969 Marine Technology Society.
Chapter 10

Major World Fishing Nations

It is important in our studies to have firmly in mind a concept of the relative value of the
world fisheries. This concept is especially important in today's world of rapid communica-
tion and transport of goods, as is understanding how large imports and exports of fish affect
market demand and price. It is the kind of information fishery biologists and fishery man-
agers need for determining the status of fish stocks when they design plans to manage these
resources for maximum yield. The United Nations Food and Agriculture Organization (FAO)
has the arduous task of collecting and publishing these figures for fisheries worldwide. Total
world fish catch has increased from about 22 mmt in 1938 to 96.9 mmt in 1991 exclusive of
marine mammals and aquatic plants. These FAO statistics show a fourfold increase in pro-
duction during this time (Table 10.1).


The fishery statistics collected for use in stock management must reflect, as accurately as pos-
sible, the fishery fishing effort and catches. Any carelessness or irresponsible data collection
will produce information that may be misleading, and hence worthless. A cardinal rule in
sampling is that the quality of the final results of any study is directly related to the quality
of the original basic data that goes into the study. Fishery biologists must judiciously select
personnel to collect fishery statistics. Their recruitment, training, and supervision are essen-
tial. The collectors must be kept informed of the actual use of the data and of its potential,
as it is collected, so they can be sure to obtain the exact kind of statistics to be used by the
biologists. If the staff collecting data feels that what they labor to obtain is placed in file cab-
inets or computer banks and is not being used, the quality of their samples is almost certain
to drop.
For many years and in many countries, the sole objective of fishery statistics was for econ-
omists to determine the worth of fisheries to their countries, and how the value of their fish-
eries compared to that of other industries. Unfortunately, the collection procedures for this
type of data usually do not fill the same needs of the fishery biologists.
The sampled fish should be identified as carefully as possible. In some fisheries there are
a variety of species that may be difficult to separate and identify on the species level. Ob-
taining truthful landing and effort data from fishermen and fish processors is a touchy prob-
lem for the collection staff. Fishermen tend to be suspicious of anyone wanting information
on their activities, so it behooves the collection agent to discuss their reasons for collecting
data and encourage them to talk about their problems and views about the present status

206 Part Three Fisheries

Table 10.1 World Commercial Catch of Fish, Crustaceans, and Mollusks, by Species Groups,

Species Group 1987 (1) 1988 (1) 1989 (1) 1990 (1) 1991

Thousand Metric Tons

Live Weight
Carps, barbels, cyprinidis 4,430 4,877 5,004 5,307 5,394
Cods, hakes, haddocks 13,786 13,636 12,905 11,827 10,476
Flatfish 1,292 1,342 1,204 1,223 1,113
Herrings, sardines, anchovies 22,375 24,387 24,800 22,183 21,406
Jacks, mullets, sauries 8,296 9,128 9,350 9,728 10,077
Mackerel, snoeks, cutlassfishes 3,644 3,862 3,771 3,505 3,480
Redfish, basses, congers 5,713 5,705 5,917 5,656 5,742
River eels 103 116 110 120 121
Salmons, trouts, smelts 1,103 1,174 1,455 1,455 1,638
Shads 808 658 739 646 663
Sharks, rays, chimeras 668 689 678 690 698
Sturgeons, paddlefish 24 21 19 18 15
Tilapias 658 708 756 852 865
Tunas, bonitos, billfishes 3,644 4,065 4,082 4,373 4,478
Other fishes 15,506 15,956 16,295 17,074 17,523
Crabs 974 1,055 1,167 1,143 1,348
Krill 376 371 396 375 233
Lobsters 229 226 211 216 222
Shrimp 2,365 2,497 2,508 2,575 2,647
Other crustaceans 327 312 320 323 340
Abalones, winkles, conchs 1,101 100 87 71 68
Clams, cockles, arkshells 1,490 1,465 1,459 1,463 1,540
Mussels 1,135 1,287 1,311 1,321 1,332
Oysters 1,112 1,093 1,041 1,005 1,007
Scallops 739 868 838 877 816
Squid, cuttlefish, octopus 2,318 2,290 2,722 2,328 2,560
Other mollusks 799 847 751 797 791
Miscellaneous 364 281 312 283 342
Total 94,379 99,016 100,208 97,434 96,926
Source: Food and Agriculture Organization of the United Nations. Yearbook of Fishery Statistics, 1991,
Vol. 72. Rome.

and future of the fishery. If they feel the motives are sincere, the quality of the data obtained
will improve.
Effort data (the labor input, vessels, skill, and technology used to catch fish) is difficult to
collect because, in many fisheries, time on the fishing grounds in actual fishing may be insep-
arable from the time running to and from the grounds, or from restricted fishing due to
heavy weather, etc. The amount of time spent searching for schools to make sets on can in-
dicate the abundance of the stock. Effort data should be collected from fishermen at the dock
when they land their catch so that the information will be fresh in their minds and more ac-
curate. An index of the measure of effort that is related as nearly as possible to the stock
abundance should be selected for each fishery and each type of gear. Information that is as
Chapter 10 Major World Fishing Nations 207

complete as possible on the number of fishermen, the kind of gear used, and how and where
it is used is desirable. In most developed countries, vessels used in marine fisheries are reg-
istered with a government agency, so there is no need to bother the skipper or crew for a de-
scription of their vessel. Such records are kept by the Coast Guard in the United States.
Fishery biologists interested in the effects of fishing on fish stocks need data on the gross
catch or nominal catch from a fishery, not just what is landed. Often landings have to be
converted to nominal landings for each way the catch is landed, e. g., frozen whole, frozen
filleted, unfrozen gutted, headed. It follows that the collector-biologist must also obtain es-
timates of fish caught but not brought to the dock for whatever reason (Chapter 8) in order
to determine total fish removed from a stock. It cannot be overemphasized that the collec-
tion of all manner of fishery statistics must be done in a careful, detailed, and standard
Questions frequently arise about the recentness of statistics. It is only natural to want
the most up to date information available when working on a problem. However, obtain-
ing fishery statistics of catch and effort in just one single country can be a enormous task.
For a developed country like the United States, and any country using computers, the gath-
ering, assembling, and distributing statistics from a wide geographic area for many different
kinds of fish and shellfish, as well as getting details on types of gear and, types of process-
ing takes considerable time (Fig. 10.1). This is a formidable undertaking; and, as a result,
the statistics that fishery biologists and managers work with are frequently 2 to 3 years old
before they become available (Fig. 10.2). Some governmental agencies mitigate this problem
to a degree by issuing preliminary figures suitable for some analyses until the final figures
become available.

Figure 10.1 Distribution of world fisheries by oceanic zones. From National Science Board (1972)
Patterns and perspectives in environmental science. Washington, D. C.
208 Part Three Fisheries

18 Artic Sea 47 Atlantic, Southeast 71 Pacific, Western Central

21 Atlantic, Northwest 48 Atlantic, Antarctic 77 Pacific, Eastern Central
27 Atlantic, Northeast 51 Indian Ocean, Western 81 Pacific, Southwest
31 Atlantic, Western Central 57 Indian Ocean, Eastern 87 Pacific, Southeast
34 Atlantic, Eastern Central 58 Indian Ocean, Antarctic 88 Pacific, Antarctic
37 Mediterranean & Black Sea 61 Pacific, Northwest
41 Atlantic, Southwest 67 Pacific, Northeast

Figure 10.2 Major world fishing areas for statistical purposes. From FAO Yearbook of Fishery
Statistics. FAO, Rome.

When judging the importance of fishing catches and fishery products, two methods are
available. The first, "total weight landed," is the most used, and the second, "economic
value," is usually more difficult to obtain and calculate. Both methods are subject to certain
problems inherent in collection: fish and shellfish may come to the dock gutted, or heads re-
moved (as with shrimp), or shucked (as with oysters), or in the round (as with finfish); there-
fore, different weights are given for each kind of product. To calculate economic value, a
wide variety of different prices paid for fish because of size, quality, and demand must be
considered; also, supply makes the calculation of a total price difficult.
The complications do not end there because some fish are used directly as human food,
and some fish are utilized as fish meal (reduction fish) that goes into domestic animal feed;
animals that later become human food are generally classified separately. The latter are gen-
erally taken in enormous numbers and the unit price therefore is very much lower than fish
for human food.


The great fishing nations of the world (highest in production by weight of fish harvested an-
nually) share common characteristics. While all of these top nations do not possess all of
these characteristics, they generally have most of them (Fig. 10.3).
Chapter 10 Major World Fishing Nations 209

• • • • •
USSR China USA Chilt' Pt'ru

•• • • • ••
Long Coast Line
Sea Faring Tradition
D []
.:.:.:. ill]
. ...

•• •
Limited Agriculture Land--- []
Fish Important in Diet
... .

• • •
High Technological Development- [ill
:-:-:.: EJ El
.. .

• •• • • •
Government Subsidies []


Large Human PopulationlArea-

D [J :::;::'

• • • •• ••
" ' -

Deep Water Ports

Productive Waters Near
Unpolluted Coastal Water---
... [J
... []
. ...

E )
D [J

Not applicable Partiall y Applicable


Figure 10.3 General characteristics of great fishing nations. Courtesy A. Hunter.

Geographic Features

1. Long coast line, generally running north and south, covering a considerable temperature and
habitat range, supporting a wide variety of fish and shellfish species.
2. Limited land suitable for agriculture crops and land for grazing.
3. Large human population per unit of land.
4. Protected deep water ports.
5. Productive waters nearby. This feature is much more important today due to the 200-mile fish-
ing limit.

Socioeconomic Features

6. Tradition of sea faring and sea fishing.

7. Traditionally fish is an important item in the diet of the population.
S. Relatively high level of mechanical-technological development.
9. Encouragement and financial assistance (subsidies) from their governments.
10. Coastal waters not heavily polluted.

The United Nations Food and Agriculture Organization (UN /FAO) gathers and summa-
rizes statistical information on world fisheries by statistical areas, by species caught, and by
kinds of fishing gear. Reviews of several great fishing nations follow (Fig. 10.4).
210 Part Three Fisheries

lIIelrlc Tons (1IIIIIIons)


10 C
8 iii

1982 1985 1987 1990
6th 5th 4th 3rd 2nd "1st

rzlChlie 0 USA IS! Peru .Japan I8:l USSR • China

Figure 10.4 World commercial catch by leading countries 1980-1990. From Fisheries of the United
States (1991). Current Fishery Statistics No. 9100. NOAA/NMFS.

Since about 1900, Japan has been the greatest fishing nation in the world, except for the
period 1962 to 1971 when Peru led production with anchovy catches processed for fish meal.
Later, in 1988, Japanese landings fell below those of China. Japan has a large population,
over 124 million or 852 people per square mile; little of its land is suitable for farming (less
than 15 percent), and there is no grazing land. Fish consumption is high: 147 lb (67 kg) per
person annually. It is easy to see that the Japanese fish heavily on a wide range of finfish and
shellfish out of necessity.
"Japanese Fisheries Based in Overseas Areas," a report prepared by the Branch of Special
Reports (Fishery Leaflet 485) describes the expansion of the Japanese fisheries after World
War II.
During World War II most of the large Japanese fishing vessels were destroyed. Conse-
quently, few high-seas operations could be conducted until large vessels were built and the
Supreme Command Allied Powers [SCAP] (MacArthur) line imposed by Allied authorities
in 1946 was relaxed. Although the SCAP Line was changed at various times to enlarge Jap-
anese fishing areas, these changes did not permit the Japanese to fish distant offshore waters
freely. With the end of the Occupation in 1952, the Japanese began to develop high-seas op-
erations on a large scale.
As the Japanese fishing fleet expanded and Japanese vessels began to fish waters more
distant from their home shores they found some areas closed which had been fished inten-
sively by them before World War II.
These restrictions and other factors, such as the concentration of fishing in the already
overcrowded areas near the homeland, gave impetus to the building of large vessels for dis-
tant operations by the Japanese. New fishing grounds were sought and the Japanese fisher-
ies expanded to new high-seas areas of the world. As distances increased from the fishing
Chapter 10 Major World Fishing Nations 211

grounds to the homeland, bases abroad were sought and fishery operations based overseas
came into existence.
Joint overseas fishing enterprises have been sponsored primarily to invest Japanese capi-
tal in areas where rich fishery resources exist, to relieve pressure on Japanese resources by
removing vessels and fishermen from the coastal fisheries, to foster emigration, and to re-
duce the cost of operating large vessels in overseas areas by providing them with bases
nearer the fishing grounds.

The overseas fishery agreements are on a basis of:

1. joint companies to conduct fishing and processing operations;
2. contracts or concessions to supply fishery products to local markets or processing plants or for
3. technical assistance;
4. exploratory fishing;
5. refueling or transshipment bases; or
6. selling directly in a foreign port fish taken by high-seas operations.

A single agreement in a foreign country may consist of one or more of these arrangements.
Some joint fishing enterprises are conducted at the request of a foreign country to train na-
tionals in modern fishery methods and provide fishery products for the local market or for
export. Some have been established primarily to explore the possibility of tuna fishing in
waters distant from Japan.
The 200-mile exclusive economic zone (EEZ) has had a profound effect on high seas and
foreign-controlled fishing. Japan's distant water fleets that fished in other nation's waters by
permit have suffered lower catches due to quota reductions and fishing fee increases. Fuel
costs also are higher.
To overcome these problems, Japan has become heavily involved in the aquaculture of yel-
lowtail, porgy, oyster, prawn, and algae. The Japanese government pays subsidies to develop
highly productive coastal fishing grounds by means such as artificial fish reefs, nursery
ground construction, and large-scale protection of these grounds in an effort to increase pro-
duction. The Japanese emphasis on the improvement and development of her coastal fishing
grounds includes pollution abatement.
The 1981 marine fisheries catch, representing 98% of Japan's total catch, amounted to 11.1
million tons, an increase of about 2 percent from 1980. The total increase in the marine fish-
eries catch was achieved despite decreases in the catch of distant water fisheries and marine
aquaculture. Offshore and coastal fisheries easily offset the losses in the other two sectors of
marine fisheries. In 1991 Japan produced 9.3 mmt.
The decreasing distant water catch reflects the continued effect of the establishment of 200-
mile fishery zones in many countries, especially in those where catch quotas for Japanese
fishermen have been imposed. However, the rate of decline in the distant water catch, which
was as high as 20% in 1978, decreased to only 3% in 1981. The harvest of whales was 4,887
in 1981 a decline of about 6% from 1980.

Russia has not traditionally been considered an important seafaring or ocean fishing na-
tion, but many of her people are enthusiastic fish eaters. Russia is, and has been for many
years, a fish importing country. She now has about 149 million consumers within her own
212 Part Three Fisheries

At the time of the Bolshevik revolution in 1917, the Russian fishery was conducted princi-
pally in her numerous lakes, rivers, and large inland seas. Ocean fishing was scattered and
primitive, and was carried out close to shore. Only 20% of her total fish catch of approxi-
mately 500 thousand metric tons in 1922 was taken in the ocean.
After the 1917 revolution the communist regime began expanding and mechanizing the
Soviet fishing fleet. The Soviets began operating their first large trawlers in 1929, but growth
of the fisheries was slow prior to World War II, and all operations were near its Siberian
coast. After the war, the Soviets reexamined their fisheries and began a program of large
capital investment in standardized vessel construction, utilizing proven fishing and support
vessel designs. Soviet shipyards were expanded to meet increased requirements. It quickly
became evident, however, that they could not keep pace with the construction needs, and
contracts were granted outside the USSR.
By 1960 the USSR had a large and growing fleet of modern high-seas fishing vessels as
well as refrigerated transport ships and floating factories and her catch reached 3.5 mmt.
This placed her near second place in world fish production. Murmansk, only an Arctic vil-
lage before World War II, has become a large and important fishing port. Of increasing im-
portance to this tremendous growth have been fishery resources of the vast Continental Shelf
off Alaska, exploited by the Soviet Far East Fishery Administration.
Although Japan is well known for fishing around the world, the USSR operated her fisher-
ies more widely. Even with a long coast line and a continental shelf supporting diverse fish-
eries, there is insufficient production to feed her population. The Soviets had a 12 nautical
mi (22 km) territorial sea limit to other nations off her coast, yet their vessels were allowed
to fish within 3 nautical mi (5.5 km) of U.s. coasts until 1966. Their large mother ships,
some 570 ft (174 m) long and 19,000 tons, with catcher boats, caused ill feelings by operating
in areas traditionally fished by Americans. In 1966 more than 340 Soviet fishing vessels (124
large stern factory trawlers, over 150 medium side trawlers, and 63 support and processing
vessels) were fishing on Georges Bank and in the mid-Atlantic Bight. In response to this
conflict, Congress extended the u.s. area of fisheries jurisdiction to 12 nautical mi in 1966.
The Soviets were accused of using what was called "vacuum cleaner" fishing. Trawlers
would line up abreast, as many as 12 or so, and drag the ocean floor for 60 to 70 mi (111 to 129
km). The trawlers transferred their catches to factory or "mother" ships for processing. Mean-
while the small, outdated, unsubsidized American vessels, unable to catch fish at the rate of the
Soviet trawlers, took whatever they could get. Americans also had to run to port to sell their
catch and gather provisions for another trip; meanwhile, the Soviets continued to fish.
By 1965, the USSR had an estimated 106 factory trawlers and, 30 factory or mother ships
fishing from Greenland south to Georges Bank, and making larger catches each year until
low U S. catch quotas in 1974 caused the United States to establish the 200-mile limit. This
limit apparently halted the decline of the badly overfished stocks.
Soviet crews are shifted each trip; captains change ships and, crews are mixed and
changed from ship to ship. Because there are more men per vessel than in other countries'
fleets, work proceeds slowly and methodically. Good safety measures and ingenious safety
devices abound. Their vessels are away from home port for long periods of time, so many
morale-building measures exist: libraries, movies, competitive sports, and social opportuni-
ties (half the crew is women). There is much camaraderie, singing, performing, dancing, and
celebration dinners that relax the crew and relieve the monotony of the long trips.
Factory ships, developed to fish efficiently in distant waters, became too costly to operate,
are restricted by the 200-mile limit, and are poorly suited to move after new species around
the globe. This worldwide fishery regulation effectively ended a spectacular era in the his-
Chapter 10 Major World Fishing Nations 213

MiUion Met.rie Tons

10 15


Former USSR
Korea Rep



Ko .... DPRp
Counlry Tot;] 1Catch
• Cumulati\'e Catch as %
Other Asia of World Tow'

Figure 10.5 Total catch by principal producers '""bnd

in 1991 and cumulative catch as a percentage of
0 20 40 60 80 100
world total. From Yearbook of Fishery
Statistics (1991), FAO, Rome.

tory of world fisheries. Had this regulation not gone into effect, the appetites of these giant
vessels for fish may have so severely reduced the size of ocean fish stocks that they would
have become uneconomical to operate.
The Soviets have engaged in "pulse" fishing, that is, putting maximum fishing pressure
into an area and fishing the stocks nearly down to zero, then moving on. This type of fishing
is more profitable than moving large fleets continuously.
In the future, Russia will have to develop inland fisheries, specialize in deep ocean trawl-
ing, and concentrate on the possible large-scale exploitation of krill. These steps are neces-
sary to overcome the crippling effects of the 200-mile limit set in 1976 (Fig. 10.5).

China's long coastline, estimated to be between 9,600 and 12,500 mi (15,446 and 20,112
km), covers a vast range of ocean climates from tropical to cold temperate waters. She has
shallow seas along her coastline that favor a large fishing industry based on stocks of cod,
herring, sea perch, sardine, and mackerel. Several species of shellfish and sea weeds are har-
vested as well. For thousands of years, millions of peasants combined some form of agricul-
ture with either fishing or freshwater fish farming; about one-third of fish production comes
from this source.
After many years of isolationism, China adapted an "open door" policy and has estab-
lished economic zones to encourage trade with foreign investors. This movement will un-
doubtedly stimulate many sluggish industries and should create greater demand for foods
that will include seafood.
214 Part Three Fisheries

China is probably in great need of modernizing fishing vessels and adding new ones to
her fleets. From the standpoint of fisheries management, China has a dire need for reliable
data on catch and fishing effort. Reliable data on the kinds and amounts of fishes caught are
difficult to obtain, yet this is necessary to obtain the greatest yields from the sea and freshwa-
ter to feed their approximately 1.2 billion people. China landed 13.1 mmt in 1991 placing
them as the top fishing nation of the world.

Measuring along the coast, Chile is 2,650 mi (4,264 km) long. It is only 110 mi (177 km)
wide and comprises 286,000 mi 2 (740,740 km 2), bordered on the west by the Pacific Ocean
and on the east by the Andes Mountains. Islands make up about one-sixth of the country.
The bulk of the 13.6 million people in Chile live in the central region where the climate is
similar to that in California.
Fishing exports are Chile's third largest source of hard currency, after mining and agricul-
ture. Like Peru, Chile is affected by El Nino, a climate change that affects coastal upwelling
by displacing the cold, nutrient-rich coastal Humboldt current, vital to the marine food chain,
with unusually warm waters that decimate fishing. Chilean pilchard (sardine) is an impor-
tant reduction (oil and fish meal manufacture) species. Mackerels are important food fish.
Catch statistics list some 43 species of commercial by important species harvest, including
For many years salmon eggs from Japan and the Pacific Northwest were transplanted to
Chile in an attempt to establish salmon runs that would take advantage of the rich krill
stocks in the Antarctic. Suitable spawning streams for salmon are rare in Chile because of the
steep slope of the coastal areas. Clearcut success has not been achieved, although adults
have returned in small numbers from some hatchery releases. Hatcheries have produced
millions of young salmon in attempts by private corporations to ranch salmon on an eco-
nomic basis.
The greatest overall foreign interest has been in Chile's booming salmon pen culture along
its southern coast. Production is primarily targeted for the U.S. market. Chilean farmers har-
vest their salmon during the northern hemisphere winters when prices in the United States
are highest. In 1992, Chile produced 39,000 tons of farmed Atlantic and Pacific salmon, up
from 70 tons in 1980.
Within about 30 mi (48 km) of the Chilean coast, over 200 species of fish live, many of
which are important to her fishing industry. In the past, whales, seals, and sea otters were
hunted from Chilean ports. Within about the last decade rock lobsters (langostinos) have
been shipped to United States, and within the last year stone crabs produced in Chile were
shipped to south Florida to take advantage of a strong demand and Florida's closed season
on stone crab fishing.


Peru has over 500 known fish species that live in cold water, lOoC (SO°F), in southern wa-
ters and warm water, 22°C (71°F), in the north. The coastline is about 1,480 mi (2,381 km)
long without estuaries, marshes, or lagoons, and is an area having upwelling of nutrient-rich
Far and away the most important fishery is targeted at a species of anchovy, Engraulis
ringens, one of the 10 anchovy species that live in the waters off Peru.
As a result of 20 years of overfishing, the anchovy population has fallen to such a low
Chapter 10 Major World Fishing Nations 215

level that it is unlikely to fully recuperate. Finally recognizing the problem directly for the
first time, the Ministry of Fisheries in Peru banned anchovy fishing and the traditional fish
meal industry, which supplied much of the world's poultry and hog producers, in favor of
catching food fish. The ban extends along the entire coastline except for a 150 mi (241 km)
stretch just north of the Chilean border. The Fisheries Minister says that Peru should have
declared an emergency in the fishery when the catch dropped from a high of about 15 mmt
in 1971 to 3 mmt in 1973. Scientists are trying to determine what volume can be fished with-
out further endangering the species. Two species of birds, which feed largely on the an-
chovy, are important producers of guano used as fertilizer in agriculture. The decline of the
anchovy stocks has seriously affected the well-being of the bird stocks, and in tum, harmed
the fertilizer industry. Other Peruvian fisheries include members of the family scombridae,
especially the bonito, and penaeid shrimp species. In 1991, Peruvian shrimp farmers pro-
duced 3,500 mt (heads-on) shrimp.
Peru's population, over 22 million, has been doubling every 25 years and is supported by
a precarious economy of wool, metals, and fish. The loss of the anchovy fishery and industry
infrastructure has dealt this nation a serious blow. Diversification of her fisheries and up-
grading of vessels with refrigeration, together with new modem vessels, are needed.

Norway is a long, narrow country with its northern half within the Arctic Circle. Almost
every major industry and every major town is sea based. Norway, called less a country than
a coastline, has 2,175 mi (3,500 km) of coastline. If fjords and other inlets are added, the fig-
ure becomes 12,420 mi (19,983 km) which is long, but not as long as the coastline of Chile.
There are many deep fjords, and 50,000 islands and skerries (rocky isles, reefs).
The rugged terrain and climate have bred a race of individualists whose staple diet is sea-
food. The area of Norway is about 125,000 mi 2 (323,750 km 2) that supports a population of
over 4 million. Only about 4 percent of the land is tillable. A few years ago about 13 percent
of the population was engaged in agriculture, fishing, and forestry; about 4 percent of the to-
tal work force worked in fisheries. This percentage is considerably higher, of course, in the
coastal districts, and especially in the northernmost part where fishing is the only economic
Norway is historically and traditionally self-sufficient in fish. However, she does import
modest amounts of fish for processing and re-export. Fish are abundant off its coast because
the Gulf Stream's warm waters create favorable spawning and growth conditions. Although
seasonal fisheries are predominant, much fishing also is done in more distant waters. The
main species caught are herring, mackerel, capelin, and cod.
Per capita fish consumption is estimated at about 90 lb (40 kg) per person per year. In-
come studies show that many fishermen earn less than many fixed-salary land jobs pay, so
they often combine fishing with other occupations, especially farming, to earn an adequate
income. This fact, plus its hard physical life, motivates a desire to abandon fishing as a live-
lihood. Because this industry is important for Norway's economy, the government is at-
tempting to make conditions more enticing. It has also imposed strict limits on the amounts
of fish caught, because overfishing is endangering their fish stocks, especially cod, haddock,
herring, and mackerel.
Like Japan, Norway is emphasizing aquaculture to satisfy her seafood demands in the face
of declining ocean catches. In addition to their highly successful salmon and trout farming
operations, a variety of different fish and shellfish are being investigated as candidate species
216 Part Three Fisheries

for commercial aquaculture. Examples of the growth of salmon and trout farming operations
is readily apparent; Norway's Fisheries Directorate granted 83 new concessions for salmonid
farming operations in 1983. In that same year trout and salmon raised in cages, net pens,
ponds, closed off coves, sounds, and fjords were valued at U.S. $100 million. In 1990, Nor-
way produced 158,000 tons of Atlantic salmon, but in 1992 production dropped to 140,000
tons. Worldwide salmon production has seriously affected their market (Chapter 11).

United States
The United States is blessed with many favorable geographic attributes and productive
waters, generally not found in combination among the other top 10 fishing nations, that con-
tribute to high landings. The length of the coastline greatly exceeds that of the other coun-
tries, being some 90,000 mi (144,810 km) on just the mainland. Adding in the offshore states
and territories such as Hawaii, Puerto Rico, and Guam this figure swells somewhat. Not
only is the coastline long, but it has a wide variety of habitats extending from very cold
climes to subtropical waters that support over 200 commercial and recreational finfish and
shellfish species. Features that contribute significantly to productivity are large estuaries and
a broad continental shelf along the east coast; upwelling along the California coast; large riv-
ers such as the Mississippi that carry nutrients to the Gulf of Mexico that make these areas
highly fertile fishing grounds; and the Columbia River that provides a route for salmon to
reach the extensive spawning grounds in the inland areas of the Pacific Northwest. Large ar-
eas of protected waters such as Puget Sound, Long Island Sound, and the inland waters of
southeastern Alaska are desirable areas for small trollers and seiners seeking both recrea-
tional and commercial fish. Many excellent natural harbors and ports add to the long list of
natural attributes (Fig. 10.6).
For the most part, the U.S. fishing fleet has consisted of small vessels, generally 5 tons or
under. The Pacific tuna fleet has for years employed large vessels and sailed considerable
distances in search of surface-swimming tunas off the coasts of Central and northern South
America. The United States constructed two large factory trawlers, but they were taken out
of service because of high operating costs and labor problems.
The most valuable United States fisheries are also the oldest, those off New England. For
many years the Gulf of Mexico was thought to be an aquatic desert, but it was later found to
be a very productive body of water yielding vast quantities of fish and shellfish. For many
years shrimp fishermen from the Gulf and southeastern United States travelled as far south
as the waters off eastern Central America and northeastern South America; many also fished
the Campeche banks off the Yucatan Peninsula of Mexico. The 200-mile fishing limit changed
this pattern. The Pacific southwest once supported the greatest volume of fish landed any-
where in North America, mainly enormous landings of sardines, a fishery that declined dra-
To put to rest any question that the opponents of the purchase of Alaska from Russia were
wrong when they labelled the transaction as "Seward's Folly," one has only to look at the
millions of dollars taken from Alaskan waters in the salmon, bottom fish, herring, and crabs
landings, not to mention timber and other resources from the land that greatly augment these
rich aquatic resources. In 1990 alone 5.1 billion lb (2.3 billion kg) were landed in Alaska.
United States fishermen, using old vessels usually manned by elderly fishermen, competed
for many years side by side with numerous large, modern, heavily subsidized foreign fishing
vessels from countries such as Japan and Russia. In an attempt to compete with this not-so-
subtle foreign invasion on a more nearly equal footing, the United States provided modest
Chapter 10 Major World Fishing Nations 217

5.1 bllon t> (54'1.)

234.4 mlHon Ib (21.)

769.6 milian t> (9'f.)

Sout~ Aftantlc
293.1 mIflon It> (3$)

Figure 10.6 U.S. commercial landings by region 1990. From Fisheries of the United States, (1991)
Current Fishery Statistics No. 9100. NOAA/NMFS.

subsidies in the form of fishery loans for construction of new vessels and modernization of
older vessels.
Unfortunately, many of the valuable productive habitats that existed years ago have been
lost due to damming of streams and rivers for electric generating plants, heating nearshore
waters with thermal wastes, and dumping other chemical and human wastes into our coastal
waters (Chapter 13). Thus, the United States has lost some of its productive waters in the in-
terest of becoming an important industrial nation.
Many modern, high quality fish processing and packaging plants have been constructed
along the coasts of United States, with efficient distribution systems. The Alaska pollock con-
tributed greatly to U.S. production in recent years.
Despite the many advantages that the United States has as a great fishing nation, consid-
erable government support goes to agriculture instead of fishing or aquaculture. Over the
years the per capita consumption of fish has increased, but the demand is being satisfied by
importation of fish from many foreign countries, not by U.S. fishermen.


Countries with herring, sardines, anchovies, and codlike fishes take advantage of these spe-
cies to increase total landings. As shown in Figure 10.7, U.s. landings in the EEZ during
218 Part Three Fisheries


Figure 10.7 Commercial catches in the U.S.

1986 1 987 1988 1989 1990 1991 exclusive economic zone (EEZ) showing the
decrease in foreign catches and joint ventures
lID Foreign Catches. U.s. Vessel landings ISIJolnl Venture
with increase in U.s. vessel landings.

('ports by Continent

• NA $362.3 million
SA S5.6 mi~ljon
EU $509 .2 million
• />S $2. 140.0 million
. . Af $1.7 million
AU $18.3 million

Figure 10.8 U.S. exports of edible fishery products, 1991. From Fisheries of the United States (1991)
Current Fishery Statistics No. 9100 NOAA/NMFS.

Import. by Con tinent

• NA S' ,842.7 million

UII SA s.s54.3 m mi on
• EU $519. 3 million
• /JS ' million
.. K $45.1 milion
AU S 267,~ mii1[on

Figure 10.9 U.S. imports of edible fishery products, 1991. From Fisheries of the United States (1991)
Current Fishery Statistics No. 9100 NOAA/NMFS.
Chapter 10 Major World Fishing Nations 219

1986-1991 have increased while foreign catches and joint ventures landings have decreased.
For many years the United States has imported far more edible fish products than it exports
(Fig. 10.8). Exports of 3,037 million lb (1,377 million kg) and imports of 5.6 trillion lb (2.5 tril-
lion kg) were reported by the United States during 1991 (Fig. 10.9).


One might expect the per capita consumption of fish and shellfish to be very high for the top
fishing nations of the world. This is true for Japan and the USSR, first and second highest,
respectively, in weight of fish landed, and who have very high per capita consumption, 106
lb (48.1 kg) (Japan) and 95 lb (43 kg) (Russia). However, for many other nations this is not
true. India, for example, is one of the top 10 great fishing nations, yet her per capita con-
sumption is estimated at 7 lb (3.2 kg) per year. Denmark's catch is smaller than India's, yet
on the average each Dane consumes about 10 times as much, 77 lb (35 kg) fish per year.
These apparent anomalies occur because some developing countries have large land masses
and often no system of distribution of perishable fishery products is available to inland in-
habitants. Others have large segments of their populations consisting of people of low in-
come who cannot afford fish. Furthermore, landings of high-value fish in many of the
developing countries will be exported to take advantage of a better market in another coun-
try. The availability within a country of plentiful, inexpensive foods, such as beef, pork,
poultry, or plant crops will also reduce the consumption of fish. These are but a few of the
myriad interacting factors that affect the consumption of fish by various nations.
World catches of species and species groups (Fig. 10.10, 10.11) indicate the positive rela-
Million Metric Tons

SouthAlssk.a pollack
Am. pilchard ~~~fi~~~~~~~i3::}!];]
Ancboveta 1305
OIileanjBd: mack.erel 1==~~lfi.:;:;"'====~.9~
Japanese pilchard I===~~':::'::_ _ _= " ' ;
S.ipjaclr. tuna I===.~ 22.9 %
Silver C3rp I==~~
Specie< Totol Cmcb
Atlantic nelTing I==~~
&ropean pilchard I===;:;~ ~ Cumulmive Catch as iJi
Atlantic·rod 1===#:;' of World TOlal
Common carp I===f;~
Chub mackerel I==~:;'
Grasscarp I==~:;'
Yellowfin tuna ,====:=-'
Largebead bairtail
Pacific cllrped oyster
Bignead co",
Atlantic ma~ke.rel
A"JUC3l1ian beml].g 3&2%
Gulf menhaden 3&8%
Japanae scallop 39.3%
Argeotil1¢ bake 39.9%
Jap3n~e anchory 40.4%

Figure 10.10 Catch by principal species in o 20 40 60 80 100

1991 and cumulative catch as percentage of the Percentag.e
world total. From FAO Yearbook of Fishery
Statistics (1992), FAO, Rome. Species listed are those with total catch above 500.000 metric tons.
220 Part Three Fisheries

Carp'. and other Cyprinids 1=----' 5.394

Tirapias and other Cichlids
MisreL Fre.hwater Fi.hes F----, 6,320
Salmons~ Trouts, Smelts
Or.b~[" Diadromolls Fi!ibes
Flounders, Halibuu., Soles
Cod'. Hakes. Haddock> F - - - - - - , 10.467
Re<lr~he~ B" .... C<>nS"" I===:::::!.::::'~-,
Jacks, Mullets, Sauries 10.071
Herring!. Sardine!., Anchovies F=======--=~------, 21.406
Tunas, Bonitos.. Bi.l1fl.Shes 4,478
Mack:erels, Sn~k.s. Cuttlassf.
Sbaru, Rays, Chimaeras
Miscel. MarirJe FIShes F------.---, 10,390
Freshwater Crustaceans
Sea-.spiders, Crabs
Loblten, Spiny-rock lobste["$
Shrimps, Prawns
Krill, Planktonic Crustaceans
Other Marine Cru:!ilaceans
Freshwater Molluscs.
Scallops. P<Ctens
Clams, Coctfes, Arksbells
Squid~ Cuttlef" o<topu ...
Other Marine Molluscs Aquatic Animals 341
Figure 10.11 Total catch by groups of species
o 5.000 10.000 15.000 20.000 25.000
in 1991. From FAO Yearbook of Fishery
Thowand Metric Toru
Statistics (1991). FAO, Rome.

tionship between the abundance of traditional and desirable food fish stocks off the coasts of
certain countries and the standings of those countries as great fishing nations. Furthermore,
stock abundance affects the variation in per capita fish consumption from country to country
in world fisheries. This has been especially evident since the coming of the worldwide 200-
mile fishing limit.


Borgstrom, G. 1964. Japan's world success in fishing. Fishing News (Books) Ltd., London.
Borgstrom, G. and A. J. Heighway. 1961. Atlantic Ocean fisheries. Fishing News (Books) Ltd., London.
Coull, J. R. 1993. World fisheries resources. Routledge, London.
Cushing, D. H. 1974. The management of marine fisheries. University of Washington Press, Seattle.
Cushing, D. H. 1975. Fisheries resources of the sea and their management. Oxford University Press,
Cushing, D. H. 1977. Science and the fisheries. Oxford University Press, London. Edward Arnold,
Cushing, D. H. 1988. The provident sea. Cambridge University Press, Cambridge, UK.
Everhart, W. H. and W. D. Youngs. 1982. Principles of fishery science. Cornell University Press, Ithaca,
New York. (Second Edition).
Firth, F. E. (ed.). 1971. The encyclopedia of marine resources. Van Nostrand Reinhold, New York.
Chapter 10 Major World Fishing Nations 221

Food and Agriculture Organization of the United Nations. FAO Yearbook of fishery statistics. (Pub-
lished annually).
Graham, M. (ed.). 1956. Sea fisheries. Their investigation in the United Kingdom. Edward Arnold,
Gulland, J. A. (ed.). 1970. The fish resources of the ocean. FAO Fish. Tech. Pap. (97) 425 pp.
Harden-Jones, F. R. 1974. Sea fisheries research. John Wiley & Sons, Inc., New York.
Holden, M. 1994. The common fisheries policy. Orgin, evaluation and future. Fishing News Books,
Idyll, C. P. 1978. The sea against hunger. Crowell, New York.
Johnson, F. G. and R. R. Stickney. 1989. Fisheries: Harvesting life from water. Kendall/Hunt Publish-
ing Co., Dubuque, IA.
Moiseev, P. A. 1971. The living resources of the world ocean. Israel Prog. Sci. Trans. TT 71-50026.
Rothschild. B. J. 1983. Global fisheries: Perspectives for the 1980s. Springer-Verlag, New York.
Rounsefell, G. A. 1975. Ecology, utilization, and management of marine fisheries. C. V. Mosby, St
Louis, MO.
Royce, W. F. 1984. Introduction to the practice of fisheries science. Academic Press, New York.
Shapiro, S. (ed.). 1971. Our changing fisheries. U.s. Department of Commerce, NOAA/NMFS.
Warner, W. W. 1983. Distant waters: The fate of the North Atlantic fishermen. Atlantic-Little, Brown,
New York.
Chapter 11

Sea Farming (Aquaculture)

Interest in aquaculture in the Western World developed only about 30 years ago despite doc-
umentation that various types of aquaculture were practiced as early as 2000 B.C. in the Far
East. Optimistic articles predicting that sea farming or pond culture would soon provide sea-
foods cheaply and in controlled situations-a "new found" protein source-that would liter-
ally feed the world. As time passed few examples of aquaculture success were reported in
Western nations; an obvious and widespread lack of significant progress was evident. Based
on the great expectation generated by early reports, entrepreneurs in the Western World
hoped for profitable ventures and perhaps a miracle remedy for the world hunger situation.
Large corporations allotted substantial portions of their financial resources to aquaculture at
a time when in many developed countries buzz words such as "new ventures" and "diversi-
fication" were popular; unfortunately, at this same time, biotechnology of aquatic organisms
and aquaculture field trials for many species were unavailable. Accounts of failed ventures
were commonplace.
The type of aquaculture carried on successfully in the Far East was, in many cases, estab-
lished as small subsistence and part-time operations to feed members of the farmers' imme-
diate families, usually with little or no profit motive. Western entrepreneurs faced the grim
conclusion that most aquaculture ventures were big money losers. They had not realized the
high level of economic risk in aquaculture. Many were unaware of the extremely limited bi-
otechnological knowledge for desirable species, information that is required for success.
With years of experience behind us we now see new, optimistic, yet more realistic, predic-
tions. P. Larkin, in speaking of private aquaculture ventures, predicted, "My guess is that
over the next 50 years, aquaculture production will equal, if not surpass, wild production of
fish." If this prediction is to be realized, the present world aquaculture production from fresh-
and saltwater, presently about 13 mmt, must increase to about 75 mmt (assuming present
world production from capture fisheries is constant). On an annual basis, by this extrapola-
tion, world aquaculture production would have to increase about 1.2 mmt per year.
Estimates of worldwide aquaculture production are fraught with many problems: scarcity
of data and different means of data reporting from country to country, enormity of the area
estimated, diversity of practices, and similar needs (Tables 11.1, 11.2). However, there are
good reasons to accept these predictions, not the least of which is that they are made by
highly qualified fishery scientists with years of experience. In addition, aquaculture has ben-
efited since from numerous biotechnological advances, accumulated experience, and the mis-
management of capture fisheries as will be discussed briefly below.

Table 11.1 Asia Cultured Shrimp Harvests, by Country and Quantity, 1983-1991.


Country 1983 1984 1985 1986 1987 1988 1989 1990 1991

Metric Tons (live weight)

China 9,000 19,300 40,700 82,800 153,300 199,400 175,000 150,000E 140,000E
Indonesia 39,000 43,000 50,000 53,000 73,000 96,000 97,000 120,000E 140,000E
Thailand 11,600 13,000 15,800 17,900 23,600 55,600 93,500 100,000E 110,000E
India 13,000 15,000 17,000 18,000 22,000 25,000 25,000 30,000E 35,000E
Taiwan 16,800 20,700 32,700 70,000 114,500 43,700 32,300 30,200 30,000E
Vietnam 13,000 18,000 20,000 22,000 22,000 30,000E 30,000E
Philippines 12,100 28,900 29,000 31,100 35,800 44,900 47,900 54,000 30,000E
Bangladesh 4,400 7,600 14,700 14,700 14,800 16,600 18,200 23,100 25,000E
Burma 7 10 500 4,000E 4,000E 4,000E 4,000E 4,000E
Japan 2,000 2,000 2,200 2,400 900 3,000 2,800 3,000 3,500E
Malaysia 400 100 200 300 800 1,300 2,200 2,500E 2,500E
Australia 15 28 26 200 500 600 1,100E
Sri Lanka 100 400 400 700 700 800E 800E
Korea, South 50 100 100 100 200 200 300 300 300E
New Caledonia 20 40 65 100 200 400 500E 500E
Singapore 54 64 100 100 100 100 100 50E 50E
Fiji 17 20 7 8 8 7 7E 7E
Pakistan 1 8 44 22
New Zealand 9
Total 108,400 149,800 215,800 309,400 465,500 512,900 521,900 549,100E 557,800E
E: Estimate.
From Wildman, Niemeier, Nielsen, et al. (1992). World shrimp culture Vol. 1. National Marine Fisheries Service.

224 Part Three Fisheries

Table 11.2 Value of Aquaculture Production by Continents.

1894 1985 1986 1987 1988 1989 1990

28,192 40,832 45,870 69,065 106,557 146,739 178,940
North America
550,459 527,289 569,836 651,947 750,596 798,488 844,769
South America
303,918 310,776 307,092 666,690 770,231 690,552 657,772
9,297,549 10,651,060 13,356,400 16,306,630 18,990,726 18,933,415 20,722,989
1,411,578 1,535,559 1,907,982 2,252,631 2,609,089 2,882,364 3,172,516
33,869 32,470 43,145 49,454 104,470 129,711 143,562
11,942,989 13,428,281 16,666,893 20,513,130 23,934,000 24,236,147 26,455,328
Data from FAO Fisheries Circular No. 815 Revision 4. 1992.


Virtually no reliable sources of information on aquaculture were available a few years ago.
There has been a doubling in the number of books and monograms on aquaculture every 5
years since 1961. In addition to books and monographs specifically on aquaculture, there are
many nonaquaculture journals such as those in the fields of agriculture, economics, and med-
icine, that publish aquaculture articles (Fig. 11.1). Aquaculture "how to" handbooks and
manuals can be obtained from government agencies and private publishers. These literature
sources provide an invaluable background on aquaculture biotechnology that has become
available only in recent years.

Number of books
and monographs
on aquaculture
"0 30 published each
year since 1961,
doubling every
~ .... 5 years
00 20
Figure 11.1 Numbers of books and
monographs on aquaculture published each
year since 1961, doubled every five years
1961 65 '69 '73 '77 'SI through 1981. From Maclean (1988). The
Year ICLARM Quarterly January 1988.
Chapter 11 Sea Farming (Aquaculture) 225

Figure 11.2 Fertilization of ponds in many

tropical countries is accomplished by housing
farm animals at the edge of the pond as seen in
the right background. Photo courtesy FAO.
Photographer Jack Ling.

More realistic articles have become available on the practical problems and constraints in
starting up and operating aquaculture ventures. Rosenthal identifies environmental degrada-
tion, disease control, human health, and competition for resources as impediments to over-
come for successful aquaculture. He also notes that research gaps in science and technology
still exist. Competing interest groups (salmon fishermen versus salmon farmers and ranch-
ers) and legal constraints (regulatory complexities of private versus public rights) are still
While technological advances in aquaculture in the United States frequently cannot be ap-
plied due to a variety of social, legal, and economic constraints, in Third World countries
these advances have provided impetus to aquaculture development due to less stringent reg-
ulations there (Fig. 11.2).


Techniques in finfish and shellfish hatcheries are

steadily advancing with dramatic increases in
hatchability, survival, and growth of larval organ-
isms. It is generally agreed that sanitation and
hatchery rearing techniques have improved so that
high survival is possible and, indeed, is often real-
ized. The difficulties of hatching and rearing fin-
fishes of high fecundity and with small egg size
such as mullet, yellowtail, and milkfish are being
more clearly understood, and can be avoided as a
result of improved technology. Although there is
much new information, additional research is Figure 11.3 In countries where few shrimp
needed before high survivorship of some larval hatcheries exist postlarvae for stocking
finfish can be realized. One such hurdle to over- ponds are often obtained in nearshore wa-
ters and estuaries by push nets. From
come is the ubiquitous cannibalism during mass Umali (1950). U.S. Fish and Wildl. Servo Res.
rearing of larval stages (mainly carnivorous spe- Rept. No 17.
cies) and its impact on survival. Also, for some
species wild seed survive better than those reared in hatcheries (Fig. 11.3).
226 Part Three Fisheries


Species like giant clams (Indo-Pacific), queen conch (Western Atlantic Ocean), or marron
(Australian crawfish) have recently become of increased interest to aquaculturists. Other spe-
cies proposed for aquaculture in the past that have appeared unsuccessful in field or pilot tri-
als now are of renewed interest because of biotechnological advances such as new improved
feeds, control over reproduction by hormones, and improved strains (Fig. 11.4).



Figure 11.4 Despite many successful sea farming operations, a number of species studied for farming
potential have not proved to be economically feasible at present. Some examples are: (a) stone crabs,
(b) northern lobsters, (c) sea urchins, (d) groupers, etc. Photos from a variety of sources.
Chapter 11 Sea Farming (Aquaculture) 227

Today, risks to the ecology of an area resulting from the introduction of exotic species in
aquaculture ventures are so well understood that strategies have been developed to reduce
them. In 1973, the International Council for the Exploration of the Sea formulated a Revised
Code of Practice to Reduce the Risks from Adverse Effects Arising from Introduction of Ma-
rine Species. This code remains the subject of recent meetings and discussions and includes
steps to provide a greater awareness of these risks. Numerous cases of highly successful
transplantations, for example, freshwater prawns and tilapia, show that this technique can
play an important role in aquaculture, but that the degree of risk must be properly evaluated
before taking any action.


New feeds have been developed to hasten shrimp maturity. One such product on the com-
mercial market promises larger and healthier hatches of shrimp and thus provides an addi-
tion technique to such existing methods of inducing maturity as eye stalk ablation and
artificial insemination, which have been used with moderate success.
Shrimp and finfish larvae can now be fed artificially prepared feeds that replace an old,
live standby Artemia, that contain what some manufactures call "artificial plankton." Artificial
zooplankton feeds are termed "osmo-preserved copepods and cladocerans" by one manufac-
turer and are available in different sizes for feeding to different sized larvae. Microencapsu-
lated feeds in several sizes are on the market for a variety of larval fish and shellfish. These
feeds eliminate the high expenses and effort normally required to raise live feeds. In the
past, nonliving feeds that were uneaten tended to pollute hatchery tanks and kept hatchery
operators from relying on them; this objection is said to have been solved by preventing de-
terioration of the feeds in water. However, despite the progress made on formulated diets for
larval fish and shellfish total, it has not replaced live feed in mass rearing of larval stages in
hatcheries. Vitamin C, important in the diet of fish is unstable and may deteriorate quickly.
The development of a new, more stable form of vitamin C is seen as an important break-
through for the aquaculture industry (Fig. 11.5).
A variety of new feeding devices are available to fish farmers designed to reduce the cost
of the feeding operation and to be more dependable and efficient. Demand feeders on the
market for some time have fulfilled the objectives of good feeders for some species. One of
the more versatile feeding systems is computer controlled and has been very successful in
salmon pen feeding operations. All pens are programmed to receive wet or dry feeds, in
amounts required by the density of fish in each pen, the size of the fish, and the temperature
of the water (Fig. 11.6).


Since fish farming became popular in developed countries, concern was centered on feeds
that would produce the most rapid growth with the least volume of feed and at the lowest
price. The amount of fat in fish feeds has been a long-standing problem for fish farmers.
They know it increases fish growth rates. However, high dietary fats can produce excessively
fatty fish that can reduce yields, causes the product to spoil more quickly and make it more
difficult to process. Fish oil is necessary in the diet to provide omega fatty acids, a require-
ment for many fish species. Much has been written about the benefits to human health by
228 Part Three Fisheries

~i •

Figure 11.5 Feeding shrimp in large farms in

Ecuador. (Top) Especially equipped aircraft are
used to evenly distribute feed. (Bottom)
Airplanes being loaded with shrimp food.
Photos courtesy of D. Benetti.



Figure 11.6 Computer controlled feeding system for net pen farming facility. Diagram courtesy Ole
Molaug Automatisering og Produktutvikling, Bryne, Norway.

consuming omega-3. Fish and shellfish are the only source for omega-3 fatty acids. Feeding
more fats to fishes to boost their omega-3 may impart a fishy flavor to the fish and reduces
shelf life. Consumption of fish oil has recently been linked to a possible treatment for a type
of kidney disease (see Chapter 9). Comparisons of omega-3 fatty acid content in wild and
farmed fish have shown that there is general agreement between the two types of fish and
only rainbow trout show a higher level of fatty acids in farmed fish (Fig. 11.7).
A second aspect of nutrition of farmed fish is that the taste has to at least compete favora-
bly with that of wild fish of the same species. For some species the color and firmness of
flesh might vary between farmed and wild fish. Pigments can be fed to salmon to produce
Chapter 11 Sea Farming (Aquaculture) 229

EPA & DHA Mg/IOO gm fillet

.- r-------------~--------------------__,
.- _ Wild ~ Farmed ' .. 0

1200 - --

1000 - - - -- -----


000 1---------

- 1--- - -- - - -- -
Figure 11.7 Comparison of omega-3 fatty acid
(EPA & DHA) content in wild and farmed fish. 200 ~----~

EPA & DHA are long chain unsaturated fatty

acids that are believed to be responsible for the
health benefits of fish oils. From Nettleton Cbannel Red Carp Rainbow Cabo Atlantic
(1990). Aquaculture Magazine 16 (1):34-41.

a deeper red color. This method has been used on trout for many years. While foolproof
taste tests are almost impossible to design, due in part to the wide variety of taste preferences
by the taste panels, they nonetheless generally give an indication of the relative taste prefer-
ences. In many cases the average person buying fish lacks the background to distinguish be-
tween wild and farmed fish by taste or appearance.
Clearly aquaculture is playing a prominent role in making more fish available for markets
at a time when many capture fisheries are undergoing severe stock depletion resulting in
heavily curtailed production. Consumers have never before been as conscious of the need
for care in diet to insure better health and longer lives. Fish are the forefront of good nutri-
tion and can become even more desirable with additional research. Some avenues of re-
search include; incorporating some unused fats into fish feeds, improving fish feeds by the
use of some fatty acids in the family omega-6, such as linoleic acid, and researching the role
of saturated fatty acids in feeds.


Many problems and constraints characteristic of aquaculture species are serious and must be
overcome for success. Of these, parasites and diseases in farms continue to be top problems.
High crop mortalities caused by diseases have forced aquaculture facilities out of business;
other ventures were never started due to concerns by potential investors that diseases would
cause serious crop losses, and their investment. Diseases of farmed species are caused by in-
fective agents or physical and/ or chemical conditions in ponds, by nutritional deficiencies, or
230 Part Three Fisheries

Figure 11.8 Occasionally stock must be

sampled from ponds for signs of disease.
Photo courtesy of Australian News and
Information Bureau.

by a combination of these factors (Fig. 11.8). Fortunately for the aquaculturist, some fish and
shellfish parasites normally found on wild species are excluded from ponds and tanks be-
cause the environmental conditions and intermediate hosts necessary for their growth and
development are absent. Artificial feeds used in farms also reduces the chance of infection.
Unfortunately, parasites and microorganisms that do cause trouble, such as those groups
listed below, all have short direct life cycles (do not require intermediate hosts). They repro-
duce with great speed under favorable conditions, can easily invade a fish pond, and are dif-
ficult to identify and frequently difficult to eradicate.
An example of a serious infective agent occurred in warm water shrimp facilities, the tiny
viruses that cast a gloom over the future of the highly successful marine shrimp farming, in
Asia and South and Central America. The problem has brought about considerable research
attention. In 1987 Taiwanese shrimp farmers lost 65,000 tons due to a virus disease. While
viruses are still a threat to penaeid shrimp farms, viruses are now much better understood;
control and identification techniques for viral diseases have become available in the litera-
ture. Despite their minute size, some viruses can be identified by the presence of occlusion
bodies, visible under low magnification. In 1989 the U.S. Marine Shrimp Farming Program
began developing and distributing Specific Pathogen Free (SPF) Pacific white shrimp.
Bacteria of several genera such as Pseudomonas, Vibrio, and Aeromonas spp. occur naturally
in a variety of fish and shellfish and under certain conditions can cause systemic infections,
and infections of the body gills and exoskeleton. Animals fed a nutritionally deficient diet or
who are somehow injured, can become infected by these bacteria.

Noninfectious Diseases
Several diseases caused by noninfectious agents typically occur in aquaculture facilities,
apparently caused by salinity and temperature changes, such as gas-bubble disease caused by
high levels of DO in holding facilities; low DO or crowding results in high levels of stress
and tumors. Diseases caused by inadequate diet can be another important form of damage
in fish farm production because malnutrition reduces growth or contributes to death from
other causes such as increased stress or infectious parasites.
Chapter 11 Sea Farming (Aquaculture) 231

Disease Prevention and Control

Fortunately, in recent years, considerable research has been devoted to prevention and
control of diseases of aquaculture species, especially those species of greatest economic value.
A recent outbreak of "whirling disease" in Colorado trout farms is a reminder of the ever-
present threat of disease. Prevention of disease requires the full attention and ongoing active
research programs by government, academic institutions, and the aquaculture industry; and,
when they are identified, to find means to control them.


An important aspect of successful management is prevention of predation on farmed species.

Rather serious conflicts arise when numerous small fish and shrimp are taken by fish-eating
birds, such as herons, egrets, gulls, terns, mergansers, and diving ducks. Many of the fish-
eating birds that frequent aquaculture facilities are protected by federal and in most cases,
state laws under a category of migratory birds. Despite fish farms being designed with pre-
dation prevention in mind (deep ponds with steep walls to discourage wading birds, and a
variety of frightening devices, such as gas-operated exploders, fireworks, electronic noise-
makers, flashing lights, and other imaginative devices like inflatable scarecrows), bird preda-
tion can still be a big problem. Permits to kill birds at aquaculture facilities can be obtained
only after all other means of reducing bird predation have been proved ineffective.
The enormous impact that predation has on young mollusks like clams and mussels, re-
leased in nature to augment natural stocks, and the effects of cannibalism in private crusta-
cean farms, such as crabs and lobsters, is now better understood. Greater awareness of the
roles played by predation and cannibalism in shellfish aquaculture has resulted in a greater
research effort to attempt to reduce these major causes of mortality.


An important recent advance in genetic engineering for aquaculture includes progress in pro-
duction of single sex offspring using hormones (tilapia), polyploidy techniques that reduce
variation of artificial selection, and growth hormone genes that may cause fish to quickly at-
tain twice their normal weight. These new and exciting techniques can increase aquaculture
production worldwide.


One has only to glance at any of the aquaculture newsletters or magazines to appreciate the
numerous aquaculture products (water quality instruments, pumps, aerators, etc.), supplies
(feeds, postlarvae and juveniles for planting, chemicals, etc.), and services (consulting, insur-
ance, disease identification and control, etc.) available today to aid and help insure success of
aquaculture ventures. As aquaculture production increases, more and better products, sup-
plies, and services will become available.
Computer programs and spreadsheets used in industries and businesses unrelated to
aquaculture have been tailored to aquaculture over the past several years. An important use
232 Part Three Fisheries

Figure 11.9 Many foreign countries lack

trained technicians to rear fish and shellfish
through their early life stages. In this old
photograph, hatchery personnel in Thailand
are discussing means of distributing young fish
for planting in ponds. Photo courtesy of
United Nations FAO.

is in the planning stages of a new aquaculture venture. In addition, they can be programmed
to monitor water quality, amounts of feed required for maximum growth of fish and shellfish
crops, and to maintain up to the minute cash flow balance sheets for the business.
The recent increase in technicians trained in maturation, hatchery practices, disease identi-
fication and control, feeding procedures, and grow-out procedures is a giant step forward.
But a long leap remains to be taken in many countries to provide sufficient numbers of well-
trained technicians to fill the various needs of aquaculture ventures (Fig. 11.9).


Pollution resulting from aquaculture operations is expected to increases as the production

from aquaculture facilities increase. At present, the increased nutrient loading of bays and
fjords (in Norway, Scotland, and Japan) resulting from farming several species, including
salmon in net pens, is being addressed.


Conflicts that hamper the growth of aquaculture worldwide deserve mention. Areas that are
used traditionally by different user groups such as capture fishermen, recreational fishermen,
and navigation frequently result in conflicts.
British Columbia salmon fishermen, long opposed to increased net pen salmon farming of
coho and chinook salmon, complain strongly about additional competition in the market-
place for their wild-caught fish. They voice objections that the gene pool of wild salmon will
be diluted by farmed salmon and that diseases from salmon farms will spread to wild stocks
and endanger their livelihood. A case in point: The 1991 escape of an estimated 8,000 Atlan-
Chapter 11 Sea Farming (Aquaculture) 233

Figure 11.10 Large shrimp farms like this one

in Ecuador frequently are blamed for reducing
nursery areas normally used by wild shrimp
thereby reducing fishermen's catches. Photo
courtesy of D. Benetti.

tic salmon transplants from a British Columbia holding pen emphasizes their fear that this
exotic species would harm their fishing.
Georgia shrimp fishermen are catching tiger shrimp, Penaeus monodon, an exotic, native to
Southeast Asia. These shrimp escaped several years before from a South Carolina research
facility. Like the British Columbian salmon fishermen, Georgia fishermen expressed their
concerns that the exotic species will eventually compete aggressively for living space and
food in local environments. This remains to be seen.
A common complaint leveled against aquaculturists comes from shorefront property own-
ers who object that their right to quiet enjoyment of their property is violated by floating oys-
ter rafts, net pens, and similar aquaculture equipment. They considered such things
unsightly and, in addition, make small boat navigation unsafe. Furthermore, a variety of po-
tential pollutants is common in large aquaculture net pen facilities. Fish wastes (feces) where
there is a high density of fish may be very heavy in an area having little or no water ex-
change (Fig. 11.10). This can certainly warrant complaints, not only by property owners, but
by federal or state pollution agencies. Uneaten feeds, chemicals used to prevent net fouling
such as chlorine and formalin, and antibiotics are potential polluters, too. The most serious
of these are antibiotics that are used to kill sea lice, a small crustacean harmful to salmon,
even though their use requires special precautions. Sea lice also can be controlled by "para-
site pickers" or "cleaner fish" (wrasses), a cleaner and safer method. Raising the specter of
pollution by taking it out of perspective can cause conflicts to heat up.
Use of wetlands for aquaculture is incompatible with the ecology of an area. Shrimp farm-
ers may destroy nursery grounds of commercial/recreation fish species (Fig. 11.11). Example:
In Ecuador there is an existing large shrimp fishery off the coast for adult Pacific white
shrimp, Penaeus vannamei. We have learned that the estaurine mangrove areas serve as nurs-
eries. Along comes a shrimp aquaculture facility, and shrimp farmers dig up the mangroves
to make shrimp ponds. Then they catch juvenile shrimp for their ponds in what remains of
the mangrove nursery. Impacts like these on nursery grounds, together with offshore fishing
can seriously harm shrimp stocks. In the Philippines, complaints come from fishermen that
fish and shrimp ponds have caused a decline in wild fish harvest, about 420,000 tons annu-
Imports of farmed species may depress market prices of locally caught species (Fig. 11.12).
A classic case occurred in 1991 when many countries, mainly Norway, Scotland, Canada, and
Chile all had large farmed salmon production at the same time. In addition, wild caught
234 Part Three Fisheries

Figure 11.11 Floating salmon net pens have

produced vast qualities of salmon and trout in
recent years. Norway, Scotland, and Chile lead
the world in production of farmed salmon.
Photo courtesy of Skretting Tess Fish Farming
System, Stavanger, Norway.

Figure 11.12 The meat of sturgeon is high in

food value and their eggs command extremely
high prices as caviar. One species was
spawned as early as 1875 in North America.
New technology has renewed interest in
sturgeon farming in the United Stetes. In
California 45 farms are in operation. Photo
courtesy of Washington State Department of

salmon production in 1989 to 1990 also was record. This profusion caused salmon to drop in
price and resulted in a serious financial loss to all concerned. Putting aside what may have
become of the excess fish of the other countries, what is of interest here (by way of a tragic
example) is that the Norwegian farmers were holding their surplus stock in costly cold stor-
age. The world salmon market feared that some substantial portion of the frozen salmon
might be "dumped" in key markets at rock bottom prices. Precisely what became of these
frozen fish is still unclear, but close to the end of 1991, Norway's Fish Farmer's Sales Associ-
ation (a sales cooperative with a surplus of 40,000 tons of salmon in cold storage) with debts
of US$312 million, filed for bankruptcy. The U.S. International Trade Commission found the
Norwegians guilty of unfair trade practices and placed a 24% duty On fresh whole salmon
putting them out of the large, traditional U.S. market. Following more negotiation including
the European Free Trade Association, the dust is settling and Norway is clearly not at the top
of salmon production at this time. However, the economic value of salmon farming to Nor-
way is too big for them not to regroup for a comeback.
There have been many conflicts related to the growth of aquaculture during the past sev-
eral decades. Some may be more imagined than real; but, indeed, the spirit of competition is
alive and well in business, and smart entrepreneurs effectively use it to their advantage.
Chapter 11 Sea Farming (Aquaculture) 235

1,000 Metric tons

Figure 11.13 Latin American shrimp farms

increased production forced growers to seek
new markets. Asian cultured shrimp have
greatly outproduced Latin America. From
Weidner, Revord, Wells, and Manuar (1992).
82 83 84 85 86 87 88 89 90 91
National Marine Fisheries Service World
Shrimp Culture Vol. 2 Latin America. Year


Figure 11.14 The lack of basic biological data 3

on potential aquaculture species and
aquaculture trials several decades ago was a 2
severe deterrent. This is being overcome by
networking through organizations such as The
Aquaculture Information Center (U.S.
Department of Agriculture). From the U.s. 1984 1985 1986 1967 1988
Department of Agriculture.!

Avoiding some problems can be accomplished by aquaculturists by reviewing the history of

aquaculture ventures where they plan to begin a new facility. Cooperation with local people
and industries sometimes can be achieved if the new venture can demonstrate their sensitiv-
ity to public concerns, present and future. Government administrators must review all as-
pects of proposed aquaculture facilities to ascertain how such a venture might impact their
area, prior to approval or granting of permits.
In summary, aquaculture is now making considerable strides in production, in sharp con-
trast to many dismal failures in ventures experienced worldwide over the past two decades
(Fig. 11.13). Furthermore, continued increased production is predicted. With a more deliber-
ate, measured approach to aquaculture operations, a new era of aquaculture is dawning. As
in many other new business ventures, aquaculturists have made many mistakes and false
starts. Unlike many other business ventures that had technological background information
to guide new projects, aquaculturists frequently groped in the dark for success in what is
truly a complex industry.
The new beginning we are experiencing is due in a large measure to worldwide successes
of numerous species in recent years (Fig. 11.14). These successes result from a rather simple
formula using species that score high on the checklist of desirable candidate species, where
careful attention is paid to site selection, including suitable conditions for the species and
236 Part Three Fisheries

Macraalgae Culture Area

Plate a Frame Heat Exchanger +

~h~r,~,:.u '=;EmL:e=rg~e=nc=y==~
U=--==L---.J==.::..Ge.nerator r - - - - J
160 GPM

Settling Discharge 25 JIm

Pond Sand Filters

Figure 11.15 Drawing of University of

Miami's Aplysia (Sea Slug) Resource Facility
Showing water filtration facilities, macroalgae
culture area to feed aplysia, heat exchanges and
Seawater Intake Biscayne Bay
Screen a Manifold chillers to provide optimum temperature.
(Bear Cut) Drawing courtesy of T. Capo.

economic factors, and where, after projects are in operation, managers employ effective day-
to-day management of the facility. This is not to say that more mistakes will not be made,
or that no more financial losses will occur in future aquaculture ventures, but rather that they
can be substantially reduced by studying the past record of aquaculture trials: which have
succeeded and which have failed, and what were the reasons for the successes or failures.
This, of course, is easier said than done, for failures may have been due to poor management,
and that may be hard to uncover.
New, creative aquaculture ventures should not be discouraged; however, before scaling up
to a pilot project, and certainly well before a commercial operation is considered, the practi-
cality of each venture should be critically evaluated by a well-chosen team of experts who
will cover all important aquaculture aspects.
As for the future of aquaculture, the picture is bright mainly because of the growing de-
mand for seafood and limited harvests from capture fisheries. Poor management, or no man-
agement, of capture fisheries described in this volume are being blamed for severe
regulations on fisheries for many species and closing of formerly productive fishing areas.
Capture fisheries have experienced rapid escalation of fishing costs, meanwhile profitable
harvests have been less frequent than in the past (Fig. 11.15). Aquaculture is also a high
priced venture, but considering the progress made as outlined above, investment, although
Chapter 11 Sea Farming (Aquaculture) 237

risky, in more and more cases has shown the economic returns to be well worth the risk, ex-
amples of which are Atlantic salmon, Japanese yellow tail, and warm water shrimp.


Anonymous. 1990. Farm raised salmon win taste test over wild salmon. Aquaculture Magazine
Brown, E. E. 1977. World fish farming. AVI Publishing Co. Inc., Westport, CT.
Brown, E. E. 1980. Fish farming handbook. AVI Publishing Co. Inc., Westport, CT.
Courtenay, W. R. and J. R. Stauffer. 1984. Distribution, biology, and management of exotic fishes. Johns
Hopkins University Press, Baltimore, MD.
DeVoe, M. R. and R. Pomeroy (eds.). 1992. Use conflicts in aquaculture: A worldwide perspective on
issues and solutions. World Aquaculture 23(2):13-35.
Heen, K., R. 1. Monahan, and F. Utter (eds.). 1993. Salmon aquaculture. Halsted Press; Imprint of John
Wiley & Sons, Inc., New York.
Iversen, E. S. 1976. Farming the edge of the sea. Fishing News Books Ltd., Farnham, Surrey, England.
Iversen, E. S. and K. K. Hale. 1992. Aquaculture sourcebook: A guide to North American species. Van
Nostrand Reinhold, New York.
Iversen, E. S. , D. M. Allen and J. B. Higman 1993. Shrimp capture and culture fisheries of the United
States. John Wiley & Sons, Inc., New York.
Jory, D.E. and E. S. Iversen. 1986. Molluscan mariculture in the greater Caribbean: An overview.
NOAA/NMFS Mar. Fish. Rev. 47(4):1-10.
Jory, D. E., M. R. Carriker, and E. S. Iversen. 1984. Preventing predation in molluscan mariculture. J.
World Mariculture Society 15:421-432.
Landau, M. 1994. Taste in fish and humans. Aquaculture Magazine 20(3):40-47.
Landesman, 1. 1994. Negative impacts of coastal aquaculture development. World aquaculture
Larkin, Peter A 1988. The future of fisheries management: Managing the fisherman. Fisheries 13(1):3-
Maclean, J. 1. 1988. The growth of fisheries literature. Naga, the ICLARM Quarterly 11(1):3-4
Nettleton, J. A 1990. Comparing nutrients in wild and farmed fish. Aquaculture Magazine 16(1):34-41.
Pillay, T. V. R. 1990. Aquaculture: Principles and practice. Fishing New Books, A Division of Blackwell
Scientific Publications Ltd., London.
Pillay, T. V. R. 1992. Aquaculture and the environment. Halsted Press; Imprint of John Wiley & Sons,
Inc., New York.
Rosenthal, H. 1985. Constraints and perspectives in aquaculture development. GeoJournal 10(3):305-
Rosenthal, H. 1994. Aquaculture and the environment. World Aquaculture 25(2):4-11.
Ryther, J. H. 1984. Biotechnology: A potential U.S. contribution to mariculture. In Biotechnology in the
marine sciences, Proceedings of the First Annual MIT Sea Grant Lecture and Seminar, Colwell, R.
R., AJ. Sinskey, and E. R. Pariser (eds.). pp. 123-132. John Wiley & Sons, Inc., New York.
Ryman, N. and F. Utter (eds.). 1988. Population genetics and fishery management. University of Wash-
ington Press, Seattle.
Sharp, G. J. and C. Lamson. 1989. Approaches to reducing conflict between traditional fisheries and
aquaculture. World Aquaculture 20(1):79-80.
Shupe, S. J. 1982. Coastal aquaculture: Protein, profits and problems for a hungry world. Sea Grant
College Program, Oregon State University, Corvallis, OR.
238 Part Three Fisheries

Sindermann, C. J. 1986. Strategies for reducing risks from introductions of aquatic organisms: A marine
perspective. Fisheries 11(2):10-15.
Sindermann, C. J. and D. V. Lightner (eds.). 1988. Disease diagnosis and control in North American
marine aquaculture. 2nd rev. ed. Elsevier, Amsterdam.
Stickney, R. R. 1994. Principles of aquaculture. John Wiley & Sons, Inc., New York.
Stickney, R. R. 1988. Commercial fishing and net-pen salmon aquaculture: Turning conceptual antago-
nism toward a common purpose. Fisheries 13(4):9-13.
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Tiddens, A. 1990. Aquaculture in America: The role of science, government and the entrepreneur.
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Chapter 12

Management Objectives

The study of changes in population abundance, called population dynamics, is a necessary

perquisite for fisheries management. Fisheries research provides the information that allows
scientists to understand the dynamics of fish populations, and in tum permits opportunities
for managing fish stocks for maximum yields. Thus, successful fisheries management re-
quires that fisheries administrators have access to reasons for population fluctuations, envi-
ronmental quality, and socioeconomic aspects of the industry.


Population dynamics and managements programs for fish and shellfish stocks began around
the late 1920s in the United States and Canada (Fig. 12.). Western European countries pur-
sued similar studies earlier because greater fishing pressure was being placed on fisheries
stocks in limited geographic areas like the North Sea. On other hand, Japan and the Soviet
Union, both having large distant water trawler fleets and factory ships, were unconcerned
about conservation because they could roam the worlds' oceans and fish new, productive ar-
eas when fishing became poor where they were.
Fisheries research provides the necessary data required for fishery management equations.
Collectors of statistics are needed to provide long series of fisheries catches and effort
records, together with biological data that cover long time periods to even out year to year
fluctuations and provide evidence for trends within fisheries.


ACT (FCMA) OF 1976

After years of debate and discussion, this important Act became effective in 1977. It de-
clares that the United States has exclusive management over fish and shellfish, except tuna,
within 200 miles offshore, except within another country's territorial sea. Control is
granted to the United States over all creatures on the continental shelf, even beyond 200
miles (Fig. 12.2).
The Act came about as the result of need to protect our fish stocks. In 1975, more than
6,000 large foreign fishing vessels, many of which were subsidized by their governments,
were sighted fishing within 200 miles of U.S. shores in direct competition with U.S. fisher-
men. Because of their large size and profusion, these vessels easily outfished the smaller U.S.

242 Part Four Fisheries Management and Regulation


\, I I
....... --\,
,, ,, I I
, ' /--- I I
...•..././ " '" I I
/ f., ..................... "
./ ••••••••• "" I
t............... I'. I
/ ""'" .............. 1\
/ •••.•••• " " I \ ~;--I_ _ _---=-B_ _
/ ..............~",L" \A :
I ·····.1 "\ I B
. I B


Figure 12.1 Generalized history of a fishery. A, collapse which is very unusual. B, fishery can
stabilize at this point on each curve. C, by reducing fishing effort the fishery can be stabilized near the
optimum production or D, where maximum total catch is being taken, or at E, point where less than the
total catch is being taken but is being taken most economically. From Holden and Raitt (1974). FAO
Fisheries Technical Paper No. 115, revision 1.

t Am. rlcan
. s..lI't~

Figure 12.2 U.S. Exclusive Economic Zone. Photo courtesy of U.S. Geological Survey.

vessels, whose fishermen bitterly accused them of decimating fish stocks that "rightly be-
longed to U.s. fishermen."
To carry out the objectives of the FCMA, eight Regional Fishery Councils were set up.
Members who served on the Councils were drawn from science, government, and the fishing
industry. For details on the FCMA see Appendix B.
Chapter 12 Management Objectives 243


Fish and other aquatic organisms of our coastal waters comprise a living, natural resource of
considerable importance. All living resources may be classified as "renewable" because each
has the inherent potential of self-perpetuation through reproduction. The management of ex-
ploited renewable resources involves exploration to determine the size of the resource; con-
servation practices are designed to provide optimum conditions for natural reproduction and
growth, with annual use limited to harvesting the excess crop.
In a broad sense, the objective of management from the human standpoint is to maintain
the resource in abundance and utilize its products to the greatest possible advantage. From
the scientific viewpoint alone, the aim of wildlife management is to ensure production of
maximum quantities of economically important fish or desirable species. It must be realized
that there is an upper limit to the productive capacity of waters, just as there is a limit to the
yield of any specific land crop. Water productivity is set by such things as temperature, ox-
ygen content, light, and basic nutrients. The high productive capacity in certain waters
comes from an optimum combination of such environmental factors essential for organisms,
while the productive capacity of others are low because of a less favorable combination of the
same factors, or because some essential element is absent. It must be further recognized that
this capacity produces a small number of several species, or a somewhat larger number of a
single species.
Each living organism has its own set of optimum environmental requirements that are sen-
sitive to changes from the optimum in the complex of environmental factors. A species flour-
ishes when environmental conditions are close to optimum; its population decreases as
conditions depart therefrom and may be wiped out entirely if anyone factor reaches a critical
or lethal level. Thus, when one essential environmental condition reaches a limiting level
that prevents a species from flourishing, but still allows it to survive, a minor deterioration in
this condition will suddenly decimate a population. Conversely, a slight improvement in this
environmental condition will be followed by an upsurge in the size of the population. It is
the tipping of this delicate environmental balance that often leads to the violent year to year
population fluctuations that cause concern in wild population management. It also explains
the observed geographic distribution of any species because there is an area where conditions
are such that the animal thrives, but as its distribution extends in any direction from this
center, some environmental condition, or conditions, becomes limiting until areas are reached
where the animal, or animals, cannot survive.
It follows that intensive scientific research programs must be an integral part of wildlife
and fish management. For each species to be managed, we must determine its rate of repro-
duction and growth, its habits and behavior in the aquatic community, its optimum environ-
mental requirements, and the range of tolerance and effect of each environmental factor on
the success of the individual, or, collectively, the population. It is doubtful if scientists could
ever learn all there is to know about the individual effect of all environmental factors on an
animal because of the many complex interrelationships that constitute an animal's environ-
ment. However, certain management practices can be carried out after a reasonable store of
knowledge is obtained through scientific investigation of a species.
The reproductive potential of any animal species is generally more than adequate to main-
tain high population levels. Observation of aquatic animal fecundities indicates that the re-
productive potential is enormous, and because such spectacular population increases do not
occur regularly in nature, it is reasonable to assume that a very effective system of control is
exerted by the environment at some or all stages in the animal's life. The theoretical or max-
244 Part Four Fisheries Management and Regulation

imum potential of increase for each animal population is different but established for each
At all stages in the life history of an animal, from egg to adult, the environment exerts
pressures or resistances that take progressive tolls and reduce the theoretical rate of produc-
tion to the observed rate. Losses occur at all stages of development and are caused by such
things as poor egg production; lack of fertilization of eggs; destruction of fertilized eggs; or
failure of eggs to hatch; loss of young; death due to starvation, predation, disease; tempera-
ture, lack of oxygen, or the effect of toxic materials in the water; failure to locate sufficient
suitable spawning areas; natural death; and collection by man. If a population of animals is
to be maintained at a constant level then each spawning pair must produce two offspring
that reach maturity and spawn.
Population increase is really the difference between the reproductive potential of the pop-
ulation and the effects of environmental resistance. It should be apparent that, in manage-
ment practice, any technique that will reduce the weight or pressure exerted by
environmental resistance will show benefits because the population level will move upward
as the pressure is reduced and will more closely approximate the maximum population po-
tential level. Management programs are designed to reduce the environmental resistance
through environmental improvement including fertilization of waters, predator control, pol-
lution control, stream improvement, and many other similar projects.
At any specific time the number of individuals that can be removed safely from a popula-
tion is the difference between the actual population number and the essential breeding pop-
ulation number.
Therefore, the population of fish and shellfish are self-renewing resources, and thus are ca-
pable of being exploited in perpetuity. However, to preserve such resources it is necessary to
use them in such fashion that they remain capable of sustaining themselves. Because the
stocks are not only self-renewing, but the rate of renewal is dependent on the share left to
perpetuate itself (which is reduced by man's collections), it has the peculiar property that the
sustainable harvest increases with increasing fishing effort up to a certain point, when the
maximum allowable catch is obtained, and then actually decreases with increasing collecting
Ecologists describe habitat as the place where animals live, and the ecological niche as the
role they play in their surroundings or ecosystem. In addition to removing fish from an area
(thereby reducing the population), man also influences the size of populations by harmfully
altering their environment. Examples of this are construction of seawalls, draining and filling
wetlands, and pollution. These changes are harmful to populations of marine and nearshore
plants and animals, and, when coupled with removal of animals from the population, can
more quickly decimate the population than if surplus stock alone is removed.


Maximum Sustainable Yield (MSY)

It is generally accepted that the objective of conservation is to maintain the stocks of
aquatic animals at high levels so that they are not in danger of decimation if environmental
conditions become unfavorable. One concept, called maximum sustainable yield (MSY), sim-
ply means that, in theory, the yield or production from the fishery in terms of weight landed
does not cause reduction of the population, and, therefore, the yield is sustained. This can
Chapter 12 Management Objectives 245


Figure 12.3 Curve for an exploited fishery

resource showing how equilibrium yield
changes as fishing effort increases and similarly
changes between stock size and surplus
production. MSY, Maximum Sustained Yield. STOCK SIZE I FISHING EFFORT

go on in perpetuity if everything goes well. MSY is estimated from surplus production mod-
els (an increase in biomass, in excess of losses due to natural mortality. Figure 12.3 illustrates
the theory of MSY, starting at the left side of the graph, when the stock size in a virgin stock
(no fishing) maximum stock size exists with no surplus production. With exploitation, the
stock adjusts to new equilibrium levels at which point yield equals surplus production. The
MSY concept was developed for single-species management programs. The model assumes
that prey, predators, and competitors are usually unaffected by the fishery.
It is possible that the concept of MSY is more meaningful when applied to the total ex-
ploitable biomass than when applied to individual species, because the former is probably
less variable than the latter.

Optimum Sustainable Yield (OSY)

or Optimum Yield (OY)
In recent decades, the concept of OY, which embraces a broad spectrum of potentially ben-
eficial resource uses, has exerted a major and increasingly important influence on fishery re-
source management and allocation of resources. As early as 1950, many inland fishery
managers were stressing the social and economic aspects of recreational fishing on the basis
of parity with biological factors and commercial fishing as the consciously stated objectives of
fish management, in effect, OY. The OSY concept was further developed and defined in a
symposium on OSY as a concept in fisheries management held at Honolulu, Hawaii in 1974.
OY. The FCMA generally follows the thrust of the 1974 OY Symposium in defining opti-
mum yield as being the yield that:

1. will provide the greatest overall benefit to the nation, with particular reference to food produc-
tion and recreational opportunities; and
2. is presented on the basis of MSY from such fishery as modified by relevant economic, social, or
ecological factors.

There is no indication of weighing the importance or benefits under the FCMA as between
the yield objectives of "food production" and "recreationalopportunities" except as their mix
serves to "provide the greatest overall benefit to the nation." These two specified major na-
tional benefits may, therefore, logically be inferred to be coequal benefits under the law.
246 Part Four Fisheries Management and Regulation

To attain these benefits from any fishery, however, it is necessary to determine the OY of
that fishery. The OSY determination requires assessment of the relevant economic, social,
and ecological aspects. Larkin expressed the difficulties of interpreting OSY for fisheries
The decision maker is thus faced with the difficult task of putting a monetary value on con-
tentment and employment (the economist's approach) or putting a contentment value on
employment and economic performance (the politician's approach) ....
This is really what the concept of optimum sustainable yield (OSY) is all about. Biology
has nothing whatever to do with OSY except as it puts constraints on what you can do in
management. The real issues in OSY are questions of national or regional policy: What
kind of fisheries do you want to have? Do you want to make money, create employment,
provide recreation, or mix some of the above?

Assessing relevant economic factors. Economic factors are relatively easy to measure in
some commercial fisheries, but are difficult to almost impossible to assess with respect to rec-
reational fisheries. What are anglers willing to pay for their fishing? How can their oppor-
tunity to harvest fish on a personal basis be valued? A number of methods are used to
approximate such valuation: none is totally satisfactory or practicable. They include gross
expenditures, travel costs, housing costs, and costs of time devoted to fishing. New ap-
proaches, or useful refinements to the above, are urgently needed.
Assessing relevant social factors. Social factors are almost impossible to measure. How
does one measure the fun and excitement of multimillion dollar industries such as sport fish-
ing, or whale watching?
Assessing relevant ecological factors. Contrary to the traditional "full utilization" or per-
ceptions of commercial fishery interests, prey species are not necessarily "wasted" if not
caught in and processed through the commercial fisheries for direct human consumption or
for reduction. Many species of prey fish are needed to provide food for the larger carnivo-
rous (predator) species that both commercial fishermen and anglers seek.
At the 1974 symposium the new management concept, OSY,was discussed. Questions on
how it would work in practice were suggested by P. M. Roedel, a biologist with the National
Marine Fisheries Service (NMFS), who suggests 10 situations that could arise. These are
listed below with my addition of a few examples of fisheries that might apply to each situa-

1. The OSY in certain fisheries be equal to the MSY.

2. The OSY may approach zero harvest for substantial stocks that are demonstrated to fill essen-
tial niches in the food chain for more desirable species. Example fisheries: anchovy and men-
3. The OSY will for many fisheries approximate the maximum net economic yield.
4. The OSY yield may for limited periods exceed the MSY if economic or social demands so dic-
tate. Example fisheries: northern lobster.
5. The OSY from certain fisheries will require harvest rates greater than the MSY of some of their
component species, particularly in multispecies trawl fisheries.
6. The OSY for some stocks will be that which will maintain only the minimum population neces-
sary to ensure the species' continued existence.
7. The OSY from the point of view of a country having control of a stock might be to let another
nation harvest that stock at a predetermined rate in return for cash, credit, or some other sort
of rights that might or might not be fishery-oriented.
Chapter 12 Management Objectives 247

8. The OSY can be less than the conventional concept of maximum net economic yield for certain
marine stocks of primary interest to sport fishermen in developed countries.
9. The OSY will be zero harvest for species considered to be of greatest value for their aesthetic in-
terest, or for inhabitants of fragile environments that could be damaged by intrusion of man or
his gear or of environments that have high scenic values (coral reefs, underwater parks).
10. The OSY for "desirable" stocks that are already overharvested will range from zero up, depend-
ing on the level to which one desires to restore the stock and the speed with which one wants
to reach that level. Example: queen conch fisheries.

The concept of OSY management is still alive, but direction on how to apply it is still not
well defined.


A broad category of overfishing is called biological overfishing that, means in terms of the
biology of the species and the size of the stock, too many fish are being removed, that is,
more than the surplus production is being taken from the stock. This category can be broken
down into two subcategories of overfishing. One called growth overfishing, the other re-
cruitment overfishing. The former means that in many fisheries fish are caught too soon;
they are taken before they grow to the maximum biomass size. Recruitment overfishing
means that the number of spawners is severely reduced resulting in the production of insuf-
ficient offspring to replenish the population of fish of catchable sizes (recruits). When a ma-
rine fish stock in overfished simultaneously in both of these types, there is a high probability
that a decimated population will result. In fact, a disastrous collapse can result.

Biological Overfishing
Biological overfishing of a fish stock is difficult to define. Simply because the population
size of a fish stock decreases after fishing pressure begins on a virgin fish stock does not nec-
essarily mean that the stock is overfished because a surplus of members in the population
(called surplus stock) often can be removed without harm to the stock size. However, when
so many spawners are removed from a fish stock that the reproductive potential is greatly di-
minished and results in a substantial reduction in the numbers of offspring, then the stock
can be said to be recruitment overfished.
If high fishing pressure is exerted on a fish stock and a majority of the young or small fish
are taken from the stock without growing to a suitable marketable size, production in weight
from the fishery decreases and the stock is considered to be growth overfished. An extreme
case of growth overfishing can also effect recruitment overfishing by not allowing many fish
in the stock to attain maturity and spawn, with the result that the numbers of recruits to the
fishery in subsequent years may be reduced.
The concept of MSY (relationship between fishing intensity and average total catch) pro-
vides a figure for fishery managers to determine possible harm to the fish stock because this
relationship gives a yield value from the fishery beyond which overfishing will occur.
Listed below are typical warning signs of possible biological overfishing that will be dis-
cussed in detail later together with the concept of economic overfishing.

1. With increased fishing effort, fishermen's catches per unit time, or, catch per day's fishing, the
total catch from the fishery declines. In response to reduced catches fishermen may work
harder and/or go greater distances in an effort to increase their catches.
248 Part Four Fisheries Management and Regulation

700 900
- Sardina landings - Biomass
~ 600 ::
0 0
0 0
~ 500 :;
;; 300 400
c 0

300 iii
100 Figure 12.4 (Top) U.S. Pacific sardine,
Sardinops sagax, landings from the 1932 to 1933
o'a32 1$37 18012 ,,,7 1852 1'&151 1812 UMIT 1172 111177 1882 1817 1_
0 to the 1991 to 1992 season and biomass (age 2
years and older) estimates from 1954 to 1992.
As early as 1929 scientists recommended a
quota on total catches of 200,000 to 300,000 tons
2 .2
per season: however, the quota was never
implemented. Landings peaked at about
~ 1.8 650,000 tons during the 1939 to 1937 season
and dropped dramatically thereafter. From
.: 1.4
~ NMFS Our Living Oceans 1993. (Bottom)
.. Monthly nominal catch of Peruvian anchoveta,
Engraulis rigens, northern! central stock (4 to
~Co 0 .8 14""5) from January 1952 to December 1982. In
;:'" 0 .6 1960, experts cited the California sardine
OA disaster and warned that the MSY for
0.2 anchoveta was dangerously surpassed. From
o~--~~~~~~~~~~~~~~~~~~ Tsukayama and Palomares (1987). In Pauly
1953 55 57 59 61 63 65 fiT 69 71 73 75 77 79 81
and Tsukayama (eds.). ICLARM Studies and
Reviews. 15 pp.

2. The average size and age of fish in the stock decreases. Larger, older fish are removed and the
stock cannot replace them due to the heavy fishing pressure that does not permit small fish (re-
cruits) to ever reach a large size.

When a marine fish stock is overfished to a point where reproductive potential and re-
cruitment are seriously reduced, the fishery will collapse.



Excerpted from Sports Fisheries Bulletin No. 265 June, 1975:

Little attention or concern was voiced on behalf of recreational utilization of marine finfish
prior to the late 1950s. An allocation philosophy has been largely retained from this earlier
period when the predominant use of marine fish stocks was for commercial or commodity
use. There is, however, a slowly growing appreciation that maximized protein landings on
Chapter 12 Management Objectives 249

a sustained basis (maximum sustained yield) is not necessarily equated with highest value
yield from a wild fish stock, particularly one of great recreational demand.
Difficult allocation judgments that must address what is the best use, or highest social
value of a common property resource, historically have not been made by fishery regulatory
bodies. Rather, management strategies are generally developed that focus on what is best for
the resource (which is appropriate), but with a less defensible underlying strategy of main-
taining traditional user group participation in the fishery.
It is becoming increasingly apparent that aquatic resource administrators must carefully
identify the full range of benefits from the common property fishery resource. Examination
of the recent record of marine resource management decisions, reflects a growing awareness
of the complex of social, economic and ecological considerations outlined in the FCMA. This
has been particularly apparent in the territorial waters of certain coastal states where recent
analyses and management decisions based on a wider range of considerations may even be
interpreted as a trend.
Allocation of limited finfish stocks that are in great popular demanded by tourist and rec-
reational consumers is perhaps the most striking example of such shifting concern. Alloca-
tion of marine fish stocks is a highly emotional issue in coastal communities. However,
retracing the United States' record of managing common property resources, inland (nonma-
rine) fisheries, as well as terrestrial and migratory wildlife stocks, clearly reflects a history of
difficult, but now commonly accepted decisions, of setting certain resources aside for recrea-
tional, noncommodity harvesting, although traditionally exploited by market hunters for a
significant portion of our history.
Allocation decisions of a similar nature have been made for certain marine finfish stocks as
well. As early as 1917, California's San Francisco Bay-San Joaquin Delta was closed perma-
nently for market harvesting of white sturgeon. Also in California, similar actions were subse-
quently taken for striped bass, white catfish, chinook salmon, coho salmon, American shad, and
several species of croaker. Oregon "decommercialized" steelhead in the late 1960s. In Florida,
legislation has prohibited the commercial sale of sailfish, snook, bonefish, and tarpon.
The most recent, and perhaps most celebrated, case of stock allocation to the recreational
industry occurred in 1981 in Texas. For almost 100 years, Texas commercial and recreational
user groups competed for red drum and spotted seatrout. Historically, adequate data were
not available for making informed decisions on stock conservation and optimization of yield
from those stocks. Unfortunately, this remains a typical condition for many, if not most,
coastal marine fisheries today. However, in the face of declining populations of both species,
and with recently collected economic data, Texas eliminated commercial fishing for both spe-
cies on September 1, 1981. The data documented that greater benefit would accrue to Texas
citizens from a highly regulated recreational fishery for two stocks.
The Texas example is particularly important because of the care with which the social and
economic data were gathered and employed for deciding a difficult issue on merit. As re-
cently stated by a Texas fisheries administrator, the age of fishery management through sole
reliance on biological data is long gone. It has become essential to include the development
of, and reliance upon, sociological and economic data in fishery management.
The Sport Fishing Association (formerly Sport Fishing Institute) maintains that there are
other marine fish stocks of such enormous value to the recreational industry that current
commercial exploitation allowances must receive increased scrutiny. The Gulf and Atlantic
stocks of king mackerel and the Atlantic coast stocks of striped bass are two such fisheries
that come readily to mind.
250 Part Four Fisheries Management and Regulation


Yield models are graphic or mathematical representations of how yields taken by man are
affected by certain variables, usually fishing effort. Deterministic yield models, also called
logistic models or surplus stock models, assume that fishing has no effect on growth, re-
cruitment, or natural mortality of the stock. Stochastic yield models, also called dynamic
pool models, assume age of stock affects growth, recruitment, natural mortality, and yield.
In other words, the model allows variations in parameters used to estimate yield of the
In an effort to describe and understand the effect of fishing on stocks of fish various mod-
els have been proposed. Since early fishery biologists first made attempts to manage single-
species fish stocks, data had to be collected for input into recruitment, yield per recruit, sur-
plus production, and fishing mortality models. It soon became apparent that those values
had to be estimated often from small samples or uncertain measurement. Therefore, these es-
timates were sometimes inexact. The single parameter with the greatest variation, still a long
way from being understood, is the relationship between the number of adult pelagic spawn-
ers landed in a fishery and the number of young fish (offspring) that are subsequently caught
(frequently called the spawner-recruit model-see the recruitment section). Other models
that use population size, survival (mortality) rates, fishing effort, etc. are all subject to the
questions, "How accurate are my estimates? Are they suitable for the management of this
fish stock?"
Obviously, in large, complex fisheries where fishermen use different kinds of gear and ves-
sels and fish in many different areas, the accuracy of the data is more suspect than in a small
fishery where fisherman use rather standard harvesting procedures. Common sense dictates
that the fishery biologist realizes the pitfalls of data collection and makes a concerted effort
to collect the best possible data, and, when analyzing it, takes into account possible inherent
errors or irregularities that may creep in.
One of the more popular fishing yield models, due in part to its simplicity and considera-
ble applicability, is called the surplus stock model. Fishing has no effect on growth, recruit-
ment, or natural mortality. The parameters used are stock size and fishing mortality (the
index catch per unit effort CPUE is used). A disadvantage is that it requires many years of
fishing data in order to "average out" variations so that a trend can be seen. But if rates of
recruitment, growth, and natural mortality are assumed to be in a state of equilibrium, there
would be no need for estimates, and so would remove a big burden from the fisheries man-
Another model is the yield-per-recruit model (dynamic pool or stochastic model) that as-
sumes that recruitment is independent of stock size, and that the stock is in equilibrium. It
assumes that age of stock affects growth, recruitment, natural mortality, and yield; therefore,
data are required on the growth rate of a species, its natural mortality, catch, and fishing ef-
fort. Yield per recruit can be used without estimating actual numbers of fish recruited, be-
cause the yield from an average cohort during its life equals average yield of all cohorts
during any year; and the yield from a cohort is proportional to number of fish recruited to
the stock.
Virtual population analysis is a procedure that employs data on the number of fish of a
particular age in the catch and fishing effort data. The parent-progeny relationship is based
on recruits at some time after data on the catch of adults is obtained.
Chapter 12 Management Objectives 251


If the fishing industry can know in advance the size of fish stocks, their operation can be
more efficient (Fig. 12.5). Forecasting the size of runs is possible, but can be difficult; a good
example of this can be seen in the Alaskan salmon industry. The need for accurate forecasts
was especially useful in this fishery years ago when personnel, equipments and supplies
were shipped via sailing vessels from Seattle to the various canneries in southeastern Alaska,
Bristol Bay, and Kodiak. Heavy dollar losses were suffered by the industry when manpower
and supplies were either over- or underestimated, based on expected size of salmon runs,
and were frequently inaccurate. Federal agencies used information on the size of runs (fish
caught and those that escaped to spawn in the streams and rivers) and age at maturity and
looked for correlations between the number of spawners and the number of their young.
Other factors, such as conditions on the spawning grounds, time of runs, abundance of pred-
ators, and other factors, were necessary to modify the estimates of the size of returning
In 1990, an Alaska Department of Fish and Game biologist complained about the frustra-
tion of trying to forecast runs of salmon, species that can be observed in freshwater where
they hatch and live as early juveniles, when forecasts of salmon runs were found to be inac-
curate 50% of the time. In Bristol Bay, in 1989, the return of sockeye salmon to the streams
to spawn was underestimated by 13 million fish. The nagging problem of survival of the
salmon during their life at sea has faced forecasters since salmon fishing began in Alaska.
Anyone attempting to predict occurrences in nature needs solid information, a high degree
of skill, and, too often, just plain luck. Many early predictions missed the mark badly, which
told predictors that they did not have adequate information to make their estimating formu-
las, or that some unforeseen occurrence had taken place. Prediction of events is the ultimate
test of a statisticians' ability to evaluate all the factors that cause an event to happen: in this
case, knowledge of the species, and the physical and chemical factors affecting its abundance.
Successfully forecasting haddock abundance on Georges Bank was accomplished by years
of research and relatively complete fishery statistics. Also, estimates of abundance of first-
year fish were available to indicate spawning success and survival of the early larval stages.
The relatively long life of haddock (3 to 6 years), and its tendency to remain in an area, helps
greatly when predicting abundance.

iz •
~ •
Figure 12.5 Comparison of observed and
predicted trawl catches of shrimp and fish in ~ •
the southwestern Gulf of Mexico using ~

environmental parameters (independent
variables), river discharge, area of lagoon and ..

estuaries, and amount of coastal vegetation
(mangroves). From Richards and McGowan

(1989). In Biomass yields and geography of
large marine ecosystems. 287-325. Sherman
and Alexander (eds.).
'171 ,- ,.,.
,.,. ..
252 Part Four Fisheries Management and Regulation


(a) "Whites Catch (November - January)

....,..... ~o ~....... . •• .D

(b) "Reds" Catch (February to end of season)


Figure 12.6 The relationship between the total

8 catch of Australian lobsters and its predicted
value based on 4-year-old male juvenile
6 abundance and average of the puerulus larvae
abundance 3 and 4 years before
73/4 75/6 nl8 79/80 8112 83/4 85/6 87/8 (P = preliminary estimate). From Caputi,
(c) Total Catch Brown and Phillips (1988). Fins 21(2):18-22.

Predicting the abundance of short-lived animals such as warm water penaeid shrimp,
which are annual crops in southern waters, is no easy task because of the short term effects
of the environment on spawning success and the survival of their larval stages. The fore-
caster's job is made easier by long-lived fish because, when several year classes are fished to-
gether in a fishery, year to year fluctuations of recruitment success are dampened; for
example, the spring herring catches taken along the Norwegian coast include about a dozen
year classes.
Accurate population assessment of highly migratory species like tunas is extremely diffi-
cult to sample and, therefore, to obtain their abundance. Abundance of fish species caught
by both commercial and recreational fishermen is also difficult to forecast, especially because
there is incomplete or no data on the extent of the recreational catch and what proportion it
is of the total catch.
A successful system for predicting abundance of the western rock lobster (a spiny lobster,
Panulirus cygnus) off the southwestern coast of Australia has been devised. First, a mathe-
matical relationship was found between the numbers of juvenile lobsters (actually the last pe-
lagic stage, called the puerulus) that settle to the bottom in nursery areas during any year,
and the abundance of recruits into the lobster fishery 4 years later. Artificial seaweed collec-
tors placed near coastal reefs permit biologists to monitor the numbers of young lobsters set-
tling on the seaweed for protection. Using mathematic equations based on long-term counts
of pueruli and subsequent commercial catches of lobster, biologists predict the size of com-
mercial lobster catches.
Chapter 12 Management Objectives 253

Monitoring the settlement of young lobsters and prediction accuracy, enables them to ob-
tain valuable information on the effects of fishing pressure on the stocks of lobsters. The
scheme permits industry to anticipate changes in the catch roughly 4 years in advance, ample
time to take appropriate action for possible increases or decreases in the size of lobster land-
ings. The prediction system in use since 1970 is rather expensive, costing A$170,OOO annually,
but it is estimated that millions of dollars can be saved, a highly favorable cost/benefit ratio.


In the introduction we discussed the concept of fisheries as common property resources, un-
like most natural resources. This means that the raw material, fish, is available on an equal
basis to any and all takers who have the wherewithal to harvest it. Since fisheries manage-
ment began, biologists have primarily concerned themselves with the size of the resource and
the withdrawals from it (fishing and natural mortality), to the exclusion of all other factors.
It is important that fish stocks be properly protected from a biological standpoint so they can
sustain themselves; however, economists argue successfully that fisheries resources should be
utilized to achieve the greatest economic efficiency or yield from the resource, on a sustained
It has been repeated many times that commercial fishermen, unlike recreational fishermen,
do not fish for protein or fish, but for dollars. When biologists put restrictions on the size of
the catch, gear used, etc. they may not be aware of, or have ignored the negative economic
aspects of regulation on commercial fishermen.
Commercial fishermen usually have large investments in vessels, gear, insurance, licenses,
and the like, and their livelihood is threatened by numerous regulations. By necessity they
must look at the value of the harvest they can make and the total cost to them of this harvest.
In Chapter 14, we discuss techniques used by biologists and administrators that oftentimes
harm fishermen. Such restrictions are called "regulated inefficiency."
Restricting fishing effort in certain geographic areas causes the concern that if maximum
yield is not taken from fishing grounds, widespread starvation of humans in developing
countries will worsen. This is not a valid argument because the most heavily fished species
in the world are those with greatest market demand. Fishermen are not seeking to provide
protein for starving masses. Rather, the species fished provide food for populations in devel-
oped countries whose populations are already adequately fed.
Free entry into fisheries, with the resulting overcapacity of vessels and processing plants,
has had a serious affect on returns to individual fishermen, even those fishing on expensive
species. Unfortunately, in developing countries, artisanal fishermen are subject to supply and
demand relationships. When fish are abundant, prices drop because they are perishable, and
there is no way to hold them over for leaner times. Buyers take advantage of such situations
and drop the price paid dramatically. Since these fishermen cannot sell elsewhere, they sell
at whatever price they can get and continue to fish despite the reduced price. This situation
persists until the price goes so unreasonably low that they can no longer pay their bills. In
developed countries, the situation is different because the relationship between price changes
and changes in size of landings is more stable.
As a result of overcapitalization and restrictions on fisheries, a typical commercial fisher-
men in United States today is someone like this: He is an older man, with an old, and often
obsolete vessel and fishing gear. There is a better than average chance he will fish on a part-
time basis and earn a considerable proportion of his income from employment outside the
254 Part Four Fisheries Management and Regulation

toTAL COlt

Figure 12.7 Relationship between total

revenue and total cost showing the long-term
equilibrium (OAE) that would result from an
open access fishery. From Bowen and Hancock
(1989). In Marine invertebrate fisheries: Their
assessment and management. Caddy J. F. (ed.).
EFFORT 375-396. John Wiley & Sons, Inc.

fishery. He may fish essentially full-time, but to do this he may be compelled to move from
one fishery to another. These moves may be for the same species, but in a different geo-
graphic area, or for different species in the same or different geographic areas. Many fisher-
men have been pushed into these situations by regulations designed to prevent overfishing,
imposed by fisheries administrators, a consequence of treating our marine fisheries resources
as common property resources. Details on lives of commercial fishermen are given in Chap-
ter 14.
The yield-effort curve in the Figure 12.7 shows that total cost of fishing (TC) increases lin-
early as fishing effort increases. As in the previous yield-effort curve (surplus production)
the greatest yield in weight is achieved (MSY) at half the amount of effort and half the pop-
ulation size. However, the maximum economic yield (MEY) is reached at a level of fishing
effort below that of MSY in terms of weight. MEY is the difference betwewn the TC and the
total revenue (TR) earned. Profit is highest where the distance between the two curves is at
maximum. Profit, or rent, from the fishery decreases below and above this point MEY and
drops to zero where the TC line crosses the TR line. In other words, the TC equals the TR or
value of the harvest. We can say that observing the MEY is more desirable than the MSY be-
cause with less fishing effort, in addition to more profit from the fishery, the danger of bio-
logically overfishing is avoided.
Figure 12.8 shows two cases of economic overexploitation similar to the previous figure.
At point A, zero revenue is generated (TC 0 TR). This point is well below the MSY. With
lower TC there is economic overfishing and also biological overfishing at point C, because
the point is well past the MSY. Note also that in both theoretical situations, the industry-
wide profits are zero, but the weight yield to the consumer is the same. Furthermore, the
opportunity for profit along the line of total cost A is, by far, smaller than where the cost of
fishing is cheaper along line C.
A response to a change in management practice caused a conflict for the Louisiana shrimp
harvest, that is, restricting the catches of small shrimp, as is done in Texas. Researchers claim
that the supply of shrimp for peeling would decline and adversely affect the shrimp canners,
whose market is already in decline. Also, total employment in the shrimp industry would de-
cline because canneries hire a large percentage of the seasonal workers. This, in turn, would
seriously affect the economy of the small southern Louisiana towns where most of these firms
are located. Further, this change would reduce employment because the many small butter-
fly netters (who fish for small shrimp in inside waters) would be put out of business.
Chapter 12 Management Objectives 255

Total Revenue
or Catch
Equilibrium MSY or Maximum Totol Revenue
Total Cost C
of Effort CoS"\.
Figure 12.8 Two cases of economic S, Tons at
overexploitation. At Al there is no biological CI and AI
overexploitation while at CI there is. In both
examples profits to industry are zero. From
Stevenson, Pollnac, and Logan (1982).
International Center for Marine Resource
Development. University of Rhode Island,
Kingston, RI. 124 pp. EfIIS"( Eel

It is important to remember that, in promulgating regulations designed to obtain the great-

est sustained yield from fish stocks in terms of either weight landed or profit earned, the live-
lihoods of fishermen and well-being of the industry (socioeconomic aspects) must be

Seafood Prices
Seafood prices, like the tides, rise and fall. Price fluctuations for a product are not unique
to the seafood industry. But, as in agriculture, prices are subject to an important influence;
that is, the supply side of the equation is often very difficult to control because it is regulated
in part by natural phenomenon, for example, weather and ocean currents. The number of
units produced in many manufacturing operations producing durable goods can be closely
controlled, much more so than in the seafood industry. Furthermore, seafood is extremely
perishable and requires rapid and careful handling as SOOn as possible after it is caught.
Economists study relationships between price and "quantity demand," or what they call the
"demand curve" or "demand schedule." The basic concept of supply and demand operates
when supply is limited, prices go up, fishermen fish harder. The reverse is true when a spe-
cific kind of seafood is abundant and available. An important factor on the supply side is
that, in some years, large imports of inexpensive seafood from foreign waters depress prices
of domestically produced seafood seriously, affecting the livelihood of the estimated 274,000
full-time commercial American fishermen.
Value of seafood caught in a year from a certain fishery is usually based on the price paid
to fishermen, which is called "ex-vessel" or "dock-side" price. Different kinds of seafood re-
quire different processing methods and different marketing strategies. These processing and
marketing costs are passed On to the consumer in the higher prices they pay per pound of
fish at the retail level. The difference can be substantial; four to five times the price paid to
the fisherman is not UnCommOn. Economists call this "marketing margin." It is determined
by the number of middlemen and the degree of processing and marketing involved.
Of all other factors that affect the price of seafood, the most important One should be a
quality product of freshness, color, odor, and optimum size. Large, fresh-caught penaeid
shrimp rated as "pearls" bring a much higher price due to their size and color than do small
off-color shrimp. Distance to market from the landing site can add significantly to the price
of seafood because transportation is expensive. And, at certain times of year seafood prices
can be higher than at others because of seasonal preferences for seafood.
256 Part Four Fisheries Management and Regulation












Jr J



Figure 12.9 Numerous king mackerel marketing channels used for Florida landings. From Austin,
Browner, Brugger, and Davis (1977). Mackerel Workshop Report. Sea Grant Special Report No. 14.

These short-term trends in price fluctuations are superimposed on the long-term trends
traced by economists (Fig. 12.10). Examples of long-term trends are inflation or deflation,
fuel costs for fishing vessels, vessel insurance costs, and changes in consumer tastes, etc.
Considering the huge number of variables and the interaction among them, it is no wonder
that fishing is a financially risky business even when natural phenomenon are not plaguing
the industry.
Table 12.1 illustrates an example of the complexity and wide diversity of programs that
state and federal fisheries agencies have to deal with in the course of managing commercial
Chapter 12 Management Objectives 257

Table 12.1 National Marine Fisheries Service Strategic Plan for the
Conservation and Wise Use of America's Living Marine Resources

1. Rebuild overfished marine fisheries.
2. Maintain currently productive fisheries.
3. Advance fishery forecast and ecosystem models.
4. Integrate conservation of protected species and fisheries management.
5. Improve seafood safety.
6. Protect living marine resource habitat.
7. Improve the effectiveness of international fisheries relationships.
8. Reduce impediments top U.S. aquaculture.
From Strategic Plan of the National Marine Fisheries Service. Goals and objectives.
No date.

Price Supply (new)

Figure 12.10 Short run supply and demand
curves. This figure shows the effect of changes Supply (old)
in price and quantity supply and demand
determinant on supply and demand curves. P,
The shift to a new demand curve can come
about because of increases in the price
substitutes for fish. A new supply curve might P.
be defined for seasons in which fishing is P, '" Demand (new)
dangerous. From Stevenson, Pollnac, and
Logan (1982)., International Center for Marine
Resource Development. University of Rhode
Island, Kingston, RI. 124 pp. 0,0, O. QUANTITY


The concept of MSY described above was popular because it was easy to understand by sci-
entists and fishermen alike. As early as 1935, M. Graham in the United Kingdom held that
surplus stock production per unit of biomass is proportional to the difference between bio-
mass and the carrying capacity of the system. This logistic model promoted during the mid
to late 1950 by M. B. Schaefer in the United States, was widely accepted in fishery science cir-
cles. An important condition is that fishing effort is capable of being measured so that it is
truly proportional to fishing mortality rate. For these deterministic models parameters are
frequently assumed to be constant, but the reality is that surplus production in fish stocks is
highly dynamic. Another flaw inherent in the system is that stock size estimates are notori-
ously off the mark. Thus, it can be seen that the environment in which fisheries scien-
tists/managers labor is one of uncertainty. Today, the consensus is that the widespread
acceptance and application of the MSY model has caused considerable harm to some fisheries
stocks. Fishery stocks that rebounded by restrictions on the level of fishing effort include
plaice (North Sea during World War 11), Pacific halibut, and striped bass (east coast of United
258 Part Four Fisheries Management and Regulation

In the early 1970s scientists from a wide range of disciplines tried to make sense of the
lack of order at the population level by considering the system level. Chaos theory is of re-
cent origin having developed from the decades of management practices that have failed.
The title is rather frightening after reviewing all the research and numerical management
plans that have been dictated over the years to prevent just exactly what happened, mass
overexploitation of numerous important fisheries in many countries worldwide. The Califor-
nia sardine and Peruvian anchoveta are prime examples of fisheries that suffered from man's
greed, with disastrous results (Fig. 12.4).
The dictionary definition of the word chaos is "extreme confusion or disorder" and the ad-
jective chaotic is "in a state of chaos; in a completely confused or disordered condition." Au-
thors using this concept of chaotic systems say that they use it as physicists use it and that it
is an antithesis by defining it as a highly ordered system.
Scientific inadequacies include studying unfamiliar problems that are highly complex and
frequently must be addressed by trial and error. Numerous mathematical population models
have always lent a feeling of great credibility and certainty of the results. Natural fluctua-
tions in fish stock abundance frequency makes determination of historic trends of landing
difficult or impossible to recognize and categorize. Yet, these parameters have been assumed
to be constant by many fishery biologists and managers
To explain the serious overexploitation of living resources, man's greed is an important
contributor. The control of fishing effort is frequently so ineffectual that it is a just a matter
of time until overexploitation takes place. Examples are seen in land resources as well as the
oceans. The director of an NOAA fisheries management study said as late as 1986 that fish-
ery biologists could not say with a degree of confidence how many marine commercial and
recreational fishermen there are and how much fish they are harvesting from U.S. waters.
Ludwig et al. (240 1993. Science) echo the problem of human motivation in fisheries man-
agement. "Resource problems are not really environmental problems: They are human prob-
lems that we have created at many times and in many places, under a variety of political,
social, and economic systems." Will this new awareness of ignored historic lessons save val-
uable marine resources or will it be ''biopolitics as usual?"


Management-Biological Yield
Amidei, R. (ed.). 1987. Educating fisheries managers. Proceedings of a California Sea Grant Workshop.
University of California, La Jolla, CA. 43 pp.
Bell, M. C. 1986. Fisheries handbo