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DR. AXELROD'S
ATLAS
FRESHWATER
AQUARIUM FISHES
ELEVENTH EDITION
GLEN S. AXELROD, M.Sc., F.Z.S.
DR. WARREN E. BURGESS, Ph.D. | NEAL PRONEK
HERBERT R. AXELROD, Ph. D. JERRY G. WALLS
Cover photo: Aequidens sp. “Gold Saum"; photo by Dr. Harry Grier, courtesy
Florida Tropical Fish Farmers Association.
The title page photo shows Boeseman's Rainbowfish, Melanotaenia boesemani.
Photo by M.P. & C. Piednoir.
Copyrights claimed for the First Edition, 1985; Second Edition, 1986; Third Edition, 1989; Fifth Edition,
1990; Sixth Edition, 1991; Seventh Edition, 1993; Eight Edition, 1995; Ninth Edition, 1997; Tenth
Edition, 2004; Eleventh Edition, 2007; by T.F.H. Publications, Inc.
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TABLE OF CONTENTS
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Photo by M.P. & C. Piednoir, Aqua Press.
Chromobotia (formerly Botia) macracantha, the Clown Loach.
PREFACE TO THE ELEVENTH EDITION
Dr. Axelrod's Atlas of Freshwater Aquarium Fishes already is unquestionably the most com-
prehensive and renowned fish identification book ever published, relied upon by aquarium
hobbyists and professionals alike. So why would its publishers go to the effort and expense of
producing a heavily revised edition of a classic in its field that has already gone through ten
previous editions? The reason, of course, is that significant changes can make a book signifi-
cantly better, and for us at T.F.H., that's always worth a shot.
The tropical fish field changes in many ways. Fishes not seen before come onto the market—
and should be shown in the book. The names of species already familiar to fish hobbyists are
subject to change—and the book has to change with them. Better photos of species already in
the book become available—and we're glad to be able to publish them. Different groups of fish-
es become more or less important in the hobby—and the book should reflect current tastes more
accurately.
We have improved the book in all of these ways. New species have been added, satisfactory
photos have been replaced by even better ones, and photos of species that once were given
greater prominence have yielded pride of place to photos of species that for one reason or
another increase the informational value of the book.
Of course, we have continued with the apparently never-ending task of changing identifica-
tions to agree with currently accepted nomenclature. In certain cases, in fact, we've gone a little
bit beyond keeping up with currently accepted nomenclature: we've even anticipated it. That is,
we've felt sure enough about changes that we believe soon will be made that we've already
included them in this edition. (This is true of some of the names applied to the Neotropical cich-
lids.) At the same time, we've felt that the taxonomy of some other groups (the African cichlids,
primarily) is still in such a state of flux that we've refrained from making changes on as whole-
sale a basis as some taxonomists would like. Where the game of cichlid taxonomy poker is con-
cerned, we're hedging our bets and betting on the come at the same time.
Changes in nomenclature resulting from taxonomic revisions will be a continuing fact of
aquarium life, so hobbyists should get used to them if they haven't already. As is now accept-
ed practice with this Atlas, you'll be able to find many of those changes in the next edition. Until
then, you'll be able to keep up with every change on the aquarium scene through Tropical Fish
Hobbyist magazine, which will keep you abreast of new developments month by month.
Speaking of Tropical Fish Hobbyist, I'm reminded that both our magazine and T.F.H. as an
organization have been wedded to the pet field for more than 50 years. What I'm proud to look
upon as our continuity of excellence for more than half a century includes not only our pub-
lishing efforts but also our contributions in the animal health and comfort fields through our
Nylabone chews and treats and shelter products. For many years now, we have been a compa-
ny that has opened an umbrella of information and products to protect all types of pets and
make their keeping easier and more enjoyable. We are as proud of being a founding member
of such pioneering organizations as the American Pet Products Manufacturers Association as
we are of our contributions to science and conservation efforts. We hope to remain in the same
tradition for many years to come—and we know that we can do that only if we continue to win
your confidence through our dedication to providing good value.
Glen S. Axelrod, M.Sc., F.Z.S.
THE PURPOSE OF THIS BOOK
This book was created with the express purpose of making the identification of
fishes easier for hobbyists, aquarists, and scientists alike. It attempts to standardize,
as much as possible, the common names applied to fishes in the English-speaking
world, especially as they apply to those fishes that are so similar in size, shape, and
form that they cannot be distinguished readily even by an ichthyologist. These dif-
ficult fishes include—but definitely are not limited to—the popular tank-raised vari-
eties of swordtails, platies, mollies, angelfishes, and discus fishes.
As commercial breeders and aquarists develop more skills in fish breeding, espe-
cially with the ever-increasing use of hormone injections to stimulate ripe fishes to
Spawn, more and more color and fin varieties will be developed. This means that
more and more books of fish standards will be required, books that clearly identify
a fish through the use of a color photograph of the species in question.
For many years, ichthyologists have recognized the value of color illustrations in
identifying fishes, and often a scientific name is accepted if the original description
is enhanced with a suitable color illustration of the suggested species, even though
there may be significant flaws in the technical description of the fish. There is a
great distinction between color illustrations of a fish whose body was preserved for
further study after the photograph or drawing was made and one that merely hap-
pened to be photographed or drawn from an aquarium specimen. That distinction
is where this book finds its true purpose.
For more than 35 years, Dr. Herbert R. Axelrod traveled around the world visit-
ing most of the major areas that produce popular aquarium fishes. In his travels, Dr.
Axelrod had the rare opportunity to collect and photograph many of the most pop-
ular species of aquarium fishes. He then preserved their bodies and had them
examined by trained fish scientists and ichthyologists to verify their nomenclature.
In many instances, Dr. Axelrod's findings resulted in the description of new species.
While many of Dr. Axelrod's photographs lack the artistic beauty of an aquarium
scene, they are often the only authentic color photographs of a fish whose identity
is as certain as can be verified by science. Therefore, these photographs often serve
a crucial role in the proper identification of rare species. This is especially helpful,
for example, with fishes of the genus Symphysodon, the discus fishes. If Dr. Axelrod
had not photographed many of the specimens that he collected on location, we
probably still would not know what the original species and subspecies really
looked like. These photographs have allowed us to actually see “the color of life"
that taxonomists list in their papers. Several hundred of Dr. Axelrod's photographs
have been used to describe types of new species, and many of these photos are the
first, and sometimes the only known, photographs of those fishes—even today!
Thus this book is both unique and authentic, and there will never be another like
it that deals with the same groups of fishes—fishes that have made huge commer-
cial impacts on the economies of so many countries. Also, there can never be anoth-
er color photograph of a type in living colors since, by definition, a type is the sin-
gle fish selected to represent the species. Type fishes are preserved, and no living
photographs are then possible—ever!
HOW FISHES ARE IDENTIFIED
Photographs alone do not identify fishes, nor can anyone really make an absolutely positive iden-
tification of a fish without the fish's body at hand. There are, however, several other factors that lead
one to a positive identification of a particular species. Ichthyologists commonly refer to two of the
most important methods of determining speciation as meristic traits and morphometric measure-
ments, known together as meristics.
Morphometric measurements are standard measurements used to determine various dimensions
of a fish, including standard length, snout length, the length of the longest ray on the dorsal fin, or
the depth of the caudal peduncle (Moyle and Cech 2000). General instructions for making these
measurements can be found by referring to Hubbs and Lager (1964) and Strauss and Bond (1990)
(Moyle and Cech 2000). These measurements often change as fishes grow. For this reason, mor-
phometrics are usually expressed as ratios to a standard length.
Meristic traits are taxonomic characteristics that are frequently used to place initial groupings of
fishes into generic categories. These traits represent anything on a fish that can be counted, such as
fin rays and spines, vertebrae, pyloric ceca, scale rows, and lateral line pores (Moyle and Cech
2000). The most commonly encountered problem with meristic traits lies in human error, as incon-
sistent measuring techniques or hasty measurements will give rise to inaccuracies and inconsisten-
cies in the counts. However, such errors are reduced if standard methods are used. Another source
of variation is the conditions in which a larval fish develops (Moyle and Cech 2000). Any factor that
affects larval growth, such as temperature, dissolved oxygen concentration, salinity, or food avail-
ability, is likely to affect meristic characters (Barlow 1961).
A recent compilation of fish species (Eschmeyer 1998) lists 23,250 species with valid descriptions
but estimates the number to be in the range of 25,000 or more (Moyle and Cech 2000). Furthermore,
an average of 200 new species are added to the list annually. In a short period of time, the total num-
ber of valid species may reach or even exceed 30,000 worldwide. This number obviously does not
include the additional hundreds of color varieties that man has developed commercially.
In order to manage all of these fishes, a strict system of identification needed to be developed. In
1758, the famous Swedish botanist, Linnaeus, did just that. He proposed a binomial system for nam-
ing living organisms. The process of “How Fish Get Their Scientific Names" was best described by
one of the world's greatest and most respected ichthyologists, Dr. Leonard P. Schultz, in The
Handbook of Tropical Aquarium Fishes, originally published in 1955. With his permission, the fol-
lowing section has been reprinted from the work:
“This binomial system provides a generic and a specific name, usually based on Latin or Greek
words, such as Gambusia affinis (Baird and Girard). The scientific names are followed by the name
of the author who first described the species and named it according to the International Rules of
Zoological Nomenclature. When the author's name is enclosed in parentheses, it means that the
species was described in another genus and that recent authors have transferred it to the present
genus, in this case Gambusia, since Baird and Girard in their original description named this species
Heterandria affinis. It is not necessary for scientific names of animals to have specific or significant
meaning, although many do.
“Both generic and specific names may be chosen in honor of the collector, a friend, or a colleague,
as well as for a geographical locality, a ship, or almost anything. Species names must be Latinized,
however; and when named after a man they must end in i, for example, hollandi, or after a woman,
in ae, for example, aliceae. Although the International Commission of Zoological Nomenclature has
established many pages of rules governing the naming of animals, the basic theory of the system is
that of priority—the oldest name established for the genus, species, or subspecies is the valid one,
going back to the year 1758, the tenth edition of Systema naturae by Linnaeus.
“Whenever an ichthyologist thinks he has a new species, he must search the ichthyological liter-
ature all the way back to 1758 to be certain that it is not already named. This is a difficult and time-
consuming process, requiring the facilities of extensive libraries such as the Library of Congress at
Washington and those in the largest universities which have specialized in ichthyological literature.
Bibliographical indices are used, such as the three-volume work by Bashford Dean, Bibliography of
Fishes (1916-1923), along with the Pisces section of the Zoological Record, to find papers on the
group being studied. The Zoological Record, published in London since 1864, classifies natural his-
tory literature on an annual basis. Although these tools are useful, the experience of the ichthyolo-
