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- of Freshwater Aquarium Fishes ¢ 7 “Watren: E. ‘Burgess;-Ph. Wf Neal. P _ Herb\ hees rt R.A xelrod, Ph.D. iAo St ite C, x S=~S slark Avenue West il, Ont. L4d 8C1 | VAUGHAN PUBLIC LIBRARIES ad 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. T.F.H. Publications, Inc. One TFH Plaza Third and Union Avenues Neptune City, NJ 07753 Copyright © 2007 by T.F.H. Publications, Inc. All rights reserved. No part of this publication may be reproduced, stored, or trans- mitted in any form, or by any means electronic, mechanical or otherwise, without written permission from T.RH. Publications, except where permitted by law. Requests for permission or further information should be directed to the above address. Printed and Bound in China 07 08091011 135798642 ISBN13 978-0-7938-0616-4 This book has been published with the intent to provide accurate and authoritative information in regard to the subject matter within. While every reasonable precau- tion has been taken in preparation of this book, the authors and publisher express- ly disclaim responsibility for any errors, omissions, or adverse effects arising from the use or application of the information contained herein. The techniques and sug- gestions are used at the reader's discretion and are not to be considered a substitute for veterinary care. If you suspect a medical problem, consult your veterinarian. The Leader In Responsible Animal Care For Over 50 Years!™ www.tfh.com i, ee TABLE OF CONTENTS Pherdce Olesmleventi dilom® 709.0. daxasae. yee etacn old. Seouean 4 PROMO se Ol US IEO OK) cs cnc ia katt oh oe Wok oa ar ne ho ieee 2 BlOv@LishessarevUentinedi: are tusus. wiltcttd 4c ue rwan aeeaieee Ad wna alge 6 Sane sslO FEX IsUNG (aerelay cer: tte ee a seen ORs eee eee 9 Conlinental Dittand Fist: Distribulon.. oc... oc aeus asecida 6s Ade 0 ces1 0 SOU MIESUME PS TLLOnsr ew Wary ence Ie A. tiem iT oe NS eel aeh Aeve 12 FTC RS VS EOLS teens ee wir ear os ean eee EPL een en ane ee £2 i On VacOeW Sem HIS BOOK ade atoud dt inna BAaS Madd Oe Hear aa ESS eer 13 Enoography, Acknowledgments. 4 ¢<<2% wlitiws sncawd on cadenteoeeSeee n 14 IN @mtis MOriCa NiR eGiON \ eisai olin.) uci wag wheal hen see eee 15 ErtimastoneecMOn: “werair, to. Gatits iia tly Je Riemann nti ee oan aan Te) ee 109 INC OU ODICAIBROGION s @.0 p coc. end Wlca, tusinste wao cial ohana Gkugratiet: Moba anew meee ples pAT MCAEIO ENC OUN E Sits wehbe SOs let | LPO, (lei a pe ie 578 SGUUMledsir ste eQION oni jlan ss ees Cok Cree Cee ee ee 887 PeS UUSA GA UT BRGCP O L satS ara ig doses caa ivtah Touss eroaetin Beet agv ed aila ckO Y Boose We Te 983 idexsOnoclentiiciNames. (islet Meee ak ReeA e 1089 fiero ROOD Utah NGAMICS nc. sc 2c tava cca kites wae oe Nee eee 1138 Le b s coe a” 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 fishes 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 cichlids.) 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 accepted 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 difficult fishes include—but definitely are not limited to—the popular tank-raised varieties of swordtails, platies, mollies, angelfishes, and discus fishes. As commercial breeders and aquarists develop more skills in fish breeding, especially 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 popular 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 commercial impacts on the economies of so many countries. Also, there can never be another 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 identification 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 measurements, 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 number 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 naming 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 literature 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-