Mongolia

North
Korea

Afghan- China South Japan

Snakeheads
istan
Korea
an

Iran Nepal
st
ki
Pa

(Pisces, Channidae)−
India
PACIFIC
OCEAN
A Biological
Myanmar Synopsis
Vietnam
and Risk Assessment Philippines
Thailand

Malaysia
INDIAN OCEAN
Indonesia
Indonesia

U.S. Department of the Interior

U.S. Geological Survey
Circular 1251
SNAKEHEADS (Pisces, Channidae)—
A Biological Synopsis and
Risk Assessment

By Walter R. Courtenay, Jr., and James D. Williams

U.S. Geological Survey Circular 1251

U.S. DEPARTMENT OF THE INTERIOR
GALE A. NORTON, Secretary

U.S. GEOLOGICAL SURVEY

CHARLES G. GROAT, Director

Use of trade, product, or firm names in this publication is for descriptive purposes only
and does not imply endorsement by the U.S. Geological Survey. Copyrighted material
reprinted with permission.

2004

For additional information write to:

Walter R. Courtenay, Jr.

Florida Integrated Science Center
U.S. Geological Survey
7920 N.W. 71st Street
Gainesville, Florida 32653

For additional copies please contact:

U.S. Geological Survey
Branch of Information Services
Box 25286
Denver, Colorado 80225-0286
Telephone: 1-888-ASK-USGS
World Wide Web: http://www.usgs.gov

Library of Congress Cataloging-in-Publication Data

Walter R. Courtenay, Jr., and James D. Williams
Snakeheads (Pisces, Channidae)—A Biological Synopsis and Risk Assessment / by Walter R.
Courtenay, Jr., and James D. Williams
p. cm. — (U.S. Geological Survey circular ; 1251)
Includes bibliographical references.
ISBN.0-607-93720 (alk. paper)
1. Snakeheads — Pisces, Channidae— Invasive Species 2. Biological Synopsis and Risk
Assessment. Title. II. Series.
QL653.N8D64 2004
597.8’09768’89—dc22
CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Literature Review and Background Information . . . . . . . . . . . . . . . . . . . . . . . . 4
Taxonomy and Synonymy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Common Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Description and Distinguishing Characteristics . . . . . . . . . . . . . . . . . . . . . 6
Native Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Biology and Natural History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Associated Diseases and Parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
History in Fisheries and Aquaculture . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
History of Introductions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Eastern Hemisphere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Western Hemisphere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Uses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Aquarium Trade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Live-Food Fish Trade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Biological Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
U.S. Importations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Regulations as of July 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Potential Range . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Risk Assessment Process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Rating Elements of Risk Model. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Organism Risk Potential . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Species Accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Channa amphibeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Channa argus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Channa asiatica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Channa aurantimaculata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Channa bankanensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Channa baramensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Channa barca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Channa bleheri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Channa burmanica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Channa cyanospilos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Channa gachua . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Channa harcourtbutleri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Channa lucius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Channa maculata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Channa marulioides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Channa marulius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Channa melanopterus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

Contents III
 Channa melasoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Channa micropeltes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Channa nox . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Channa orientalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Channa panaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Channa pleurophthalma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Channa punctata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Channa stewartii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Channa striata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
Parachanna africana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
Parachanna insignis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
Parachanna obscura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

FIGURES
1. Depiction of the Chinese snakehead, Channa asiatica, by Walt Disney
Productions in 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2-6. Maps showing:
2. Native distribution of the family Channidae . . . . . . . . . . . . . . . . . . . . . 8
3. Introductions of snakeheads in the Eastern Hemisphere . . . . . . . . . . . . 17
4. States where snakeheads have been collected from open waters,
were cultured prior to August 2002, or are established . . . . . . . . . . . . . 20
5. States prohibiting possession of live snakeheads as of
November 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
6. Thermal range of snakeheads (Channidae) based largely on native
range of distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

TABLES
1. Currently recognized species of the family Channidae . . . . . . . . . . . . . . . . . 5
2. Parasites of northern snakehead (Channa argus) . . . . . . . . . . . . . . . . . . . . . 14
3. Species of the family Channidae currently known to be cultured for food 16
and/or aquarium fish trade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
4. Snakeheads of interest to aquarists in the U.S. . . . . . . . . . . . . . . . . . . . . . . . 25
5. U.S. importations of live snakeheads (Channidae, all species) during 29
1997-2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
6. Origin of snakehead shipments (Channidae, all species) during the 30
past 5 or more years . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
7. States prohibiting snakeheads as of July 2002 . . . . . . . . . . . . . . . . . . . . . . . 31

IV Contents
CONVERSION FACTORS, ACRONYMS, and
ABBREVIATIONS

Multiply By To obtain

centimeter (cm) 0.3937 inch

meter (m) 3.281 foot
kilogram (kg) 2.2046 pound

Degrees Celsius (°C) may be converted to degrees Fahrenheit (°F) as

follows: °F = 1.8 x (°C + 32)

FAO Food and Agriculture Organization of the United Nations

FFWCC Florida Fish and Wildlife Conservation Commission
U.S. United States
USGS U.S. Geological Survey
ppm part per million
S$ Singapore currency
$ United States currency

Unless otherwise stated in this report, measurement of fish refers to total

length, defined as the measurement made from the tip of the snout to the
posterior tip of the caudal or tail fin. Standard length refers to the measurement
made from the tip of the snout to the base of the caudal fin.

Contents V
VI Contents
SNAKEHEADS (Pisces, Channidae)—
A Biological Synopsis and Risk Assessment

By Walter R. Courtenay, Jr., and James D. Williams

ABSTRACT

Snakeheads (family Channidae) are airbreathing freshwater fishes

containing two genera, Channa with 26 species native to Asia, Malaysia,
and Indonesia; and Parachanna with 3 species native to tropical Africa.
Some snakeheads are small, reaching about 17 centimeters, but most
are much larger, the largest reported to be 1.8 meters in length. All are
considered thrust predators with most being piscivorous as adults.
A few of the smaller snakeheads and colorful juveniles of some
larger ones have been available to hobbyists through the aquarium fish
trade. Several species are highly valued as food fishes within parts of
their native ranges, especially in Asia where they are an important part
of capture fisheries and aquaculture.
Because of these uses by humans, introductions far beyond native
ranges have occurred. One Asian snakehead has been established in
Oahu, Hawaii, since before 1900. Another species was discovered
established in southeastern Florida in 2000, and a third in a pond in
Maryland in 2002. Others have been captured from natural waters of
the United States without evidence of reproduction and likely represent
released aquarium fishes. That snakeheads at or near sexual maturity
were being sold alive in ethnic food markets raised fears that they could
be introduced into novel waters. These concerns led to this study on the
biology of snakeheads. A risk assessment is included that examines
environmental and related aspects of snakehead introductions.

1
 INTRODUCTION
Snakeheads (family Channidae) are airbreathing freshwater fishes
containing two genera, Channa, native to Asia, Malaysia, and Indonesia, and
Parachanna, endemic to tropical Africa. Taxonomy of these fishes is in flux,
but leading authorities on snakehead systematics currently recognize 26
species of Channa and 3 of Parachanna. A few snakeheads are small, reaching
about 17 centimeters; most, however, are much larger, the largest reported to be
1.8 meters in length. All are considered thrust predators with most being piscivorous
as adults.
Within parts of their native ranges, some species of snakeheads are highly
valued as food fishes, particularly in India, southeastern Asia, China, and to a lesser
extent in Africa. They have long been an important part of capture fisheries and, in
recent decades, some species have been utilized in aquaculture and a few used as
predators to control density of tilapiine fishes in culture.
Because of its popularity as a food fish in southern China and adjacent
southeastern Asia, the chevron snakehead (Channa striata) has been reported as
widely introduced into islands from the western Indian Ocean eastward to Hawaii.
The northern snakehead (C. argus) has been a market leader, and is cultured in China
and Korea. This species has been exported to other nations, including Canada and the
United States where it has been sold alive in certain ethnic markets and restaurants.
Although purposefully introduced and established in Japan in the early 1900s, its
introduction and subsequent establishment in ponds, rivers, and reservoirs of
Kazakhstan, Turkmenistan, and Uzbekistan (formerly part of the Soviet Union) in the
early 1960s appear to have been accidental.
Other snakeheads utilized as food fishes include the Chinese snakehead
(Channa asiatica), blotched snakehead (C. maculata), and spotted snakehead
(C. punctata). The bullseye snakehead (C. marulius), found in the live-food and
aquarium fish trades, is now established in Broward County, Florida, and the blotched
snakehead has been established in Oahu, Hawaii, prior to 1900.
Snakeheads used in the aquarium fish trade include a few small species and
brightly colored juveniles of several large snakeheads. They are moderately popular
with hobbyists in Japan and Europe. Several species are marketed in Canada and have
been sold in the U.S., even in states where possession of live snakeheads has been
illegal for decades. Hobbyists and importers can purchase snakeheads through a
variety of sites on the Internet. Because of their highly predacious nature, however,
snakeheads have not had a large following of interested hobbyists in the U.S. Those
who purchased attractively colored juveniles of the larger species typically found that
snakeheads became incompatible with other fishes (even killing others of their own
kind), required expensive food (preferably live), and quickly outgrew their aquaria.
This apparently has led to releases of “pet” snakeheads. As a result of these habits and
their prohibition in several states, snakeheads have had a limited market in the U.S.
aquarium fish trade. The fact remains, however, that they have been available for
purchase.

2
 The earliest known record of snakehead imports into the contiguous U.S.
was published by Brind (1914). The importation consisted of about 60 juvenile fish
that we believe were blotched snakeheads. Their progeny are thought to have been
consumed by parent fish (Brind, 1914). Klee (1987) noted that a snakehead species,
the chevron snakehead, probably a misidentification of the Chinese snakehead, was in
the U.S. aquarium fish trade by 1912. Ross B. Socolof (personal commun., 2003) said
that the Chinese snakehead was the first snakehead imported for the U.S. aquarium
fish trade in the very late 1800s or early 1900s. Innes (1917) mentioned snakeheads as
aquarium fishes, but did not include individual species. Innes (1920) reported on his
having received a “breeding pair” of what he cited as Channa fasciata from a
colleague who brought the fish to him from San Francisco. He indicated that a “single
adult pair and a few young” of this snakehead had been “recently imported into
California from Southern Asia.” The photograph of this fish in Innes (1920) is clearly
that of the Chinese snakehead, and the same photograph appeared in the account of
the Chinese snakehead by Innes (1955). Armstrong (1923) purchased four progeny of
Inne’s snakehead in 1922, and included his failures and success in breeding these fish
and their care under aquarium conditions. Stoye (1935) and Axelrod and Schultz
(1955) provided descriptions, illustrations, and information on the care and breeding
of Chinese snakehead, leading us
to believe that it was available for
sale to aquarium hobbyists
through at least the 1950s.

The Chinese snakehead

is one of a few species known to
crawl short distances overland—
references to this snakehead in the
aquarium fish literature may have
played a role in creation of an
information “strip” on this spe-
cies, published by Walt Disney
Productions in 1959 (fig. 1), and
brought to our attention by Robert
Howells of the Texas Department
of Parks and Wildlife.

Figure 1—Perhaps due to its avail-

ability in the aquarium fish hobby
through the 1950s, Walt Disney Pro-
ductions published this depiction of the
Chinese snakehead, Channa asiatica,
in 1959. This is a species of snakehead
known to crawl overland for short dis-
tances. Reprinted with permission of
Disney Publishing Worldwide.
© Disney Enterprises, Inc.

3
 A comprehensive snakehead fish study, including a biological synopsis, risk
assessment, and accounts for each species, was conducted between September 2001
and September 2002 by the U.S. Geological Survey, with support provided by the
Division of Scientific Authority and Fisheries Management of the U.S. Fish and
Wildlife Service. This study was prompted for several reasons—
• The discovery of an established population of Channa marulius in Broward
County, Florida, in 2000;
• A well established snakehead (C. maculata) population in Oahu, Hawaii, since
the late 1800s;
• Recent captures of introduced snakeheads in several states; and
• Recognition that channid fishes (at or near sexual maturity) were being sold in
live-food fish markets in the U.S.
A limited number of snakeheads have been available for sale through the aquarium fish
trade for several decades, but a new pathway— introduction of these fishes in live-food
fish markets—had been largely overlooked.
Because snakeheads are highly predatory, some having the ability to travel
overland to new water bodies, the inevitable release of these fishes by hobbyists,
escapes from aquaculture, and liberation of live-food fish into U.S. waters threatens
aquatic ecosystems. This report provides a comprehensive assessment of the risks
involved with introductions of potentially invasive snakeheads into non-native waters.

LITERATURE REVIEW AND BACKGROUND

INFORMATION
TAXONOMY AND SYNONYMY
According to Nelson (1994), the group of teleostean fishes known as
snakeheads is classified as follows:

• Class Actinopterygii
• Subclass Neopterygii
• Order Perciformes
• Suborder Channoidei
• Family Channidae
Two genera are currently recognized as comprising the family Channidae.
They are Channa (Scopoli, 1777; snakeheads of Asia, Malaysia, and Indonesia) and
Parachanna (Teugels and Daget, 1984; African snakeheads). Generic synonyms of
Channa include Channa Gronow, 1763, a nomen nudum; Ophicephalus Bloch, 1793,
and its misspelled version Ophiocephalus; Bostrychoides Lacepède, 1801; and
Philypnoides Bleeker, 1849. Synonyms of Parachanna are Ophiocephalus Günther,
1861; Parophiocephalus Senna, 1924 (originally proposed as a subgenus, but
preoccupied in the fish family Anabantidae by Parophiocephalus Popta, 1905); and
Channa Scopoli, 1777. Synonyms of the 29 species of snakeheads described herein are
included in the individual species accounts contained in the section “Species Accounts.”

4
 Myers and Shapovalov (1932) reviewed the status of the genera
Ophicephalus and Channa, and they concluded that the practice of separating the two
based on presence (Ophicephalus) or absence (Channa) of pelvic fins was invalid,
based on specimens of C. gachua from India and one from Taiwan (introduced) that
lacked pelvic fins. They placed Ophicephalus as a junior synonym of Channa. Five
species of Channa lack pelvic fins.

Vierke (1991b), Musikasinthorn (2000), Musikasinthorn and Taki (2001),

and Zhang and others (2002) consider 29 species of this family as valid (table 1).
Nevertheless, 87 species and 4 subspecies have been described (Eschmeyer, 1998, in
part) and current taxonomy is in flux. Although many described species are now
considered synonyms of recognized species, there are about 20 names that cannot be
associated with valid taxa. The plethora of scientific names for snakeheads is in part
due to the sometimes dramatic color changes that occur between early and late
juvenile stages, and adult patterns, a factor then unknown and hence unrecognized by
early taxonomists using color as one of the distinguishing characteristics. Moreover,
some descriptions lack detail, illustrations, or type specimens that could assist in
solving these taxonomic mysteries. Four new species have been described since 1990,
another put into synonymy, and two removed from synonymy and recognized as valid
during that same time period. A taxonomic revision of the family is being prepared
(Prachya Musikasinthorn, personal commun., 2002) and will likely result in more
species being recognized as valid, and new species will perhaps be described.

Table 1—Currently recognized species of the family Channidae

[After Vierke (1991b), Musikasinthorn (2000), Musikasinthorn and Taki (2001), and Zhang and others (2002).
Bibliographic sources are cited in the References section of this report or in Eschmeyer (1998)]
_________________________________________________________________________________________________________

Channa amphibeus (McClelland, 1845) - Chel snakehead1 Channa marulioides (Bleeker, 1851) - emperor snakehead
Channa argus (Cantor, 1842) - northern snakehead1 Channa melanoptera (Bleeker, 1855) - blackfinned snakehead1
Channa asiatica (Linnaeus, 1758) - Chinese snakehead Channa melasoma (Bleeker, 1851) - black snakehead
Channa aurantimaculata Musikasinthorn, 2000 - Channa micropeltes (Cuvier, 1831) - giant snakehead3
orangespotted snakehead 1
Channa bankanensis (Bleeker, 1852) - Bangka snakehead1 Channa nox Zhang, Musikasinthorn, and Watanabe, 2002 –
night snakehead1
Channa baramensis (Steindachner, 1901) - Baram snakehead1 Channa orientalis Schneider, 1801 - Ceylon snakehead2
Channa barca (Hamilton, 1822) - barca snakehead Channa panaw Musikasinthorn, 1998 - panaw snakehead1
Channa bleheri Vierke, 1991 - rainbow snakehead Channa pleurophthalma (Bleeker, 1851) - ocellated snakehead1
Channa burmanica Chaudhuri, 1919 – Burmese snakehead1 Channa punctata (Bloch, 1793) - spotted snakehead3
Channa cyanospilos (Bleeker, 1853) - bluespotted snakehead1 Channa stewartii (Playfair, 1867) - golden snakehead
Channa gachua (Hamilton, 1822) - dwarf snakehead3 Channa striata (Bloch, 1797) - chevron snakehead3
Channa harcourtbutleri (Annandale, 1918) - Inle snakehead1 Parachanna africana (Steindachner, 1879) - Niger snakehead1
Channa lucius (Cuvier, 1831) - splendid snakehead Parachanna insignis (Sauvage, 1884) - Congo snakehead1
Channa maculata (Lacepède, 1802) - blotched snakehead1 Parachanna obscura (Günther, 1861) - African snakehead
Channa marulius (Hamilton, 1822) - bullseye snakehead 1,3
______________________________________
1 Proposed common name. 2 Common name tentative. 3 Species complex.

5
 COMMON NAMES
As is typical of fishes of foreign origin, there has been a history of different
English common names utilized for snakeheads. It is not unusual to find dissimilar
names used for juveniles and adults of the same species, particularly in the aquarium
fish trade. Moreover, one can find several English common names in the scientific
literature for the same species in different parts of its native range. This also is true for
common names used by indigenous people for the same species. In India, for
example, various common names for a single species are often used by people from
diverse regions, states, or cultures. For purposes of this report, we have followed
Robins and others (1991) in using common names for the two snakeheads they
treated, selected names we felt appropriate primarily from those used in the aquarium
fish trade, and have added proposed names for some species that lacked English
common names (table 1). The common names are identified in table 1 and appear in
bold type in the section “Species Accounts.”
Accounts for each of these species are detailed in the section “Species
Accounts,” which includes illustrations or photographs, source of original description,
type specimens, synonyms, common name(s), native range, introduced range, size,
habitat preference, temperature range, reproductive habits, feeding habits, characters,
commercial importance in the United States, commercial importance in native range,
and environmental concerns. Where known, the diploid chromosome number is
included. Each account also contains a map showing native range and, where known,
location or range of introductions. Literature citations for some synonyms are not
included in the “References” section but can be found in Eschmeyer (1998) or on the
Internet at http://www.calacademy.org/research/ichthyology/catalog/
fishcatsearch.html.

DESCRIPTION AND DISTINGUISHING CHARACTERISTICS

Fishes of the family Channidae are commonly referred to as snakeheads

(sometimes serpent-heads), primarily because of their somewhat elongated and
cylindrical bodies, but particularly due to the presence of large scales on the head of
most species, reminiscent of the large epidermal scales (cephalic plates) on the heads
of snakes. Another snake-like feature is the somewhat flattened head with eyes located
in a dorsolateral position on the anterior part of the head. Anterior nostrils are present
and tubular. Dorsal and anal fins are elongated, and all fins are supported only by rays.
A few species lack pelvic fins (Nelson, 1994; Berra, 2001). The caudal fin is rounded.
The mouth is terminal and large with a protruding lower jaw, which is toothed, often
containing canine-like teeth. The prevomer and palatines may or may not be toothed,
depending on species. Scales are cycloid or ctenoid. All species possess paired
suprabranchial chambers located behind and above the gills. The chambers in Channa
are bordered by two plates, one from the epibranchial of the first gill arch and the
other as an expansion of the hyomandibular. Those in Parachanna have a simple
cavity not involving processes from the first epibranchial or hyomandibular. These
chambers are not labyrinthic (Berg, 1947), but are lined with respiratory epithelium.
All species occupy freshwater and a few can tolerate extremely low salinities.

6
 Illustrations or photographs of certain species of Channa appear in Nichols
(1943), Munro (1955), Nakamura (1963), Mohsin and Ambak (1983), Masuda and
others (1984), Lim and Ng (1990), Ng and Lim (1990), Lee and Ng (1991, 1994),
Pethiyagoda (1991), Talwar and Jhingran (1992), Kottelat and others (1993), Jayaram
(1999), and Kottelat (1998, 2001a). The three species of Parachanna were illustrated
by Bonou and Teugels (1985), who provided a key for identification of Parachanna.
But, there is no single key to identify all species of Channa, at least five of which
appear to be species complexes rather than single, distinct species.

Two larger snakehead species, the bullseye snakehead (Channa marulius)

and emperor snakehead (C. marulioides), superficially resemble the native bowfin
(Amia calva) in that all three are elongated fishes, have long dorsal fins, tubular
nostrils, and an ocellus near the base of the upper part of the caudal fin. The bowfin,
however, has its pelvic fins in an abdominal rather than thoracic or anterior-abdominal
position, and the anal fin is not elongated. Moreover, the bowfin lacks a rosette of
enlarged scales on top of its head. Other than this example, there are no native fishes
in North America or within their native ranges with which snakeheads could be
confused.

Native Distribution

Species and species complexes of the genus Channa are native from
southeastern Iran and eastern Afghanistan eastward through Pakistan, India, southern
Nepal, Bangladesh, Myanmar, Thailand, Laos, Malaysia, Sumatra, Indonesia,
Vietnam, Korea, and China northward into Siberia (fig. 2).

Of the currently recognized 26 species of Channa, 8 species and

representatives of 4 species complexes occur in peninsular Malaysia, Sumatra, and/or
Indonesia. Of the same 26 species, 13 species and 1 species complex are tropical to
subtropical; members of 6 species and 2 species complexes are warm temperate to
subtropical/tropical, 2 species complexes are cold temperate to subtropical/tropical,
and 1 species is warm temperate to boreal and can live beneath ice in the northern part
of its range. The three species of Parachanna are native to Africa and are tropical
(fig. 2).

Snakeheads are non-ostariophysan primary freshwater fishes (Mirza, 1975,

1995) and have little or no tolerance for seawater. Habitat preferences vary by species
or species complex, with a majority occurring in streams and rivers. Others live in
swamps, rice paddies, ponds, and ditches. All can tolerate hypoxic conditions because
they are airbreathers from late juvenile stages. The pH range, where known, varies by
species with one, the Bangka snakehead (Channa bankanensis), preferring highly
acidic (pH 2.8-3.8) waters (Lee and Ng, 1991; Ng and Lim, 1991). At least three
species are tolerant of a wide pH range: the dwarf snakehead (C. gachua), spotted
snakehead (C. punctata), and chevron snakehead (C. striata) survived for 72 hours at
pH levels ranging from 4.25 to 9.4 (Varma, 1979).

7
 20°W 0° 20°E 40°E 60°E 80°E 100°E 120°E 140°E 160°E
60°N
Russia

Kazakhstan
Mongolia
North
40°N Korea Japan

Pak tan n-
China

is ha

n
Iran

ista
g
Af
South
Algeria Libya Korea
Egypt Saudi India PACIFIC
Arabia Taiwan
20°N
Burma
Sudan OCEAN
Vietnam
Philippines
Sri Thailand
Lanka
Malaysia
0° Kalimantan
Equator Sumatra
INDIAN
Java Indonesia
OCEAN

Madagascar
20°S
Australia

0 2,000 MILES
EXPLANATION
Native range of Channa 0 2,000 KILOMETERS
Native range of Parachanna Scale is approximate

Figure 2—Native distribution of the family Channidae.

Biology and Natural History

• Paleogeographic origins—The fossil record indicates the presence of

channid fishes during the upper Oligocene/lower Miocene in what is now western
Switzerland and eastern France (Reichenbacher and Weidmann, 1992). Nevertheless,
Ralf Britz (personal commun., 2003) noted that identification of these fishes is based
on fossilized otoliths and may not be reliable.
Lydekker (1886) reported on fossilized skull bones of snakeheads from the
Siwalik Hills, Himachal Pradesh, northern India. These fossils and additional material
of Pontian age (early Pliocene) were described in more detail by Sahni and Khare
(1977) who described Channa bhimachari, C. gregoryi, and C. romeri based on
fossilized skull-bone material found in stream sediments. Comparing these skeletal
elements to osteological preparations of recent snakeheads, they suggested that
C. bhimachari was most closely related to C. striata and C. gregoryi to C. marulius.
They were unable to determine a recent relative of C. romeri.
Boeseman (1949) described skull-bone fossils of a snakehead from
Pleistocene deposits from Trinil, central Java. Based on comparisons with osteological
preparations of recent snakeheads, he concluded that these fossilized remains were of a
species closely related to living Channa striata, and he named the species Ophicephalus
palaeostriatus. Coupled with data presented by Lydekker (1886), Boeseman (1949), and
Sahni and Khare (1977), there is convincing evidence that congeners of living
snakeheads were present in Asia by and probably well before the Pleistocene.

8
 A majority of living snakeheads are native to southeastern Asia, with most
species found from Myanmar eastward and southeastward through Malaysia, Sumatra,
Java, and Borneo (Kalimantan). Boeseman (1949) confirmed that at least one species
of what is now included in the genus Channa was present in Java in the Pleistocene.
Jocano (1975) noted that during the Pleistocene, the Malay Peninsula, Sumatra,
Borneo, and the “Sundas” to Palawan were connected by what is termed the “Sunda
Shelf.” He described conditions as “a vast dryland covering 1.8 million square kilome-
ters,” containing a “great river,” adding that many of the present river systems of Kali-
mantan, Sumatra, and surrounding localities were tributaries of that river. He also
added that this “may explain the striking similarities of fish faunas in Sumatra, Borneo,
and the Philippines,” the latter restricted to the Palawan region of the Philippines.
From an evolutionary standpoint and considering a Pliocene to Pleistocene
presence of channid fishes in Asia, it is likely that the “great river” of the Sunda Shelf
began to dry during the late Pleistocene and speciation of snakeheads in the region
was in advanced stages. This probably occurred before parts of that shelf became
isolated as peninsulas and islands, with species evolving in differing kinds of habitats
separated from their ancestral origin(s) by geographic and biological factors. Such an
evolutionary scenario explains why, for example, species known from Sumatra also
occur in the southern part of the Malay Peninsula and Kalimantan, Bangka, Billiton,
perhaps Bali, and nearby areas. Moreover, isolation of such species “populations”
between continental and insular ranges over geologic time has doubtless led to genetic
and phenotypic differences that complicate taxonomic interpretations. Similar
linkages between what are now continental areas and nearby islands (for example,
Vietnam and China with Hainan) are known from the Pleistocene (Sterling and others,
2003). India and Sri Lanka are reported to have been connected as recently as 8,000
years ago (www.tamilinfo.org).
Fossils of snakeheads have also been identified from post-Pleistocene
deposits in the Sahara Desert (Van Neer, 1989). Banerjee and others (1988) suggested
an origin of the family from the area of Yunnan Province, southern China, dating back
to the Pliocene or earlier, but based their suggestion on ecological habitats currently
occupied by snakeheads rather than from fossil evidence. A more accurate picture of
where this family evolved and its ancestor(s) is yet to be determined.

• Spawning seasons and reproductive behavior—There is a paucity of

information on reproductive biology of many species, but several conclusions can be
drawn for those that are known. Spawning seasons vary by species. Spawning in
several species occurs primarily in summer months (June through August) but, in at
least two (the Channa striata and C. punctata species complexes), breeding pairs can
be found throughout the year. Some species spawn twice to three or more times each
year. Okada (1960) reported that female northern snakehead are capable of spawning
five times per year. There are several reports that when snakeheads pair, they remain
monogamous for a spawning season, perhaps longer, but this may not apply across the
life history of any individual snakehead.
Most snakeheads build nests by clearing a generally circular area in aquatic
vegetation, often weaving the removed vegetation around the centrally cleared area.
This results in a vertical column of water surrounded by vegetation. Sometimes the

9
 surface of this column contains pieces of removed vegetation. One species complex
(Channa punctata) prepares elaborate tunnels through vegetation leading into the nest
column. In general, the male entwines his body around that of the female, with some
species appearing to “dance” in the water column as eggs are released and fertilized
(Breder and Rosen, 1966; Ng and Lim, 1990). Eggs are buoyant, due to a large oil
droplet in the yolk mass, and rise to the surface where they are vigorously guarded by
one or both parents. Some snakeheads in one species complex (C. gachua) and
C. orientalis are reported to be mouthbrooders, with the male being the mouthbrooder
of fertilized eggs and, later, fry in C. orientalis. Peter Ng (personal commun., 2002)
suggested that C. asiatica may also be a mouthbrooder. Most snakeheads, however, are
not mouthbrooders and one or both parents vigorously guard their young. One species
(C. micropeltes) is reported to have attacked and, in some instances killed, humans who
approached the mass of young (Kottelat and others, 1993). Thus, parental care, whether
by guarding or mouthbrooding, is a behavioral characteristic of snakeheads.

One might assume, based on reported spawning habits, that presence of

vegetation is mandatory for successful spawning, but this is not the case. Wee (1982)
cited Parameswaran and Murugesan (1976b) as having documented Channa gachua,
C. marulius, and C. punctata spawning in ponds lacking vegetation. Alikunhi (1953)
noted that C. striata is also known to spawn in the absence of vegetation. These
observations, however, imply that other snakeheads are also capable of reproducing in
waters lacking vegetation.

• Fecundity and early development—There is limited information on

fecundity except for snakeheads of commercial importance. Nevertheless, that
information shows a pattern that likely applies to the entire family Channidae. An
unfertilized "egg" is an oöcyte. Once an oöcyte is fertilized by fusion of oöcyte and
sperm nuclei, it becomes an egg, with an embryo resulting if fertilization is successful.
Smaller snakeheads, such as Channa gachua and C. orientalis, produce few oöcytes
(about 20 when sexual maturity is first reached and up to 200 later; Lee and Ng, 1991,
1994). Low fecundity is a general rule among mouthbrooding fishes (Breder and
Rosen, 1966). Fecundity increases greatly in larger snakeheads and appears to be
linear, increasing in volume with increasing body length. For example, Quayyum and
Qasim (1962) recorded fecundity ranging from 2,300-26,000 oöcytes for C. striata,
increasing in number with increasing body length. Large female bullseye snakeheads,
C. marulius, among the largest species, have been reported to produce as many as
40,000 oöcytes (Jhingran, 1984). A fecundity for the northern snakehead, C. argus, was
about 50,000 oöcytes (Frank, 1970). Frank's data came from Nikol'skiy (1956), who
recorded fecundity of 22,000-51,000 in northern snakeheads from the Amur basin.
Dukravets and Machulin (1978) gave fecundity rates of 28,600 to a high of 115,000 for
northern snakehead (probably from Yangtze River stock) introduced into the Syr
Dar'ya basin of Turkmenistan/Uzbekistan. They also noted that, whereas growth of
northern snakeheads is slower than that reported for this species from the Amur basin,
growth rates from both stocks became equal once sexual maturity was reached.

10
 Oöcytes, when released from the female parent, are small, ranging from about
1 mm to slightly over 2 mm in diameter, depending on species. Fertilization takes place
by the male releasing milt (sperm) on the oöcytes as they emerge from the female.
Development time to hatching varies with water temperature and, to a lesser extent, with
the species involved. For example, hatching occurred in 54 hours at 16-26 ºC and 30
hours at 28-33 ºC in Channa punctata (Kahn, 1924). In the northern snakehead,
C. argus, eggs hatch in 28 hours at 31 ºC, 45 hours at 25 ºC, and 120 hours at 18 ºC.
In general, newly hatched fry, depending on species, are about 3.0-3.5 mm in length.
• Early life history—Following yolk resorption, snakehead fry begin feed-
ing on zooplankton. Fry typically remain together until they reach early juvenile stage,
guarded by one or both adults, when they can fend for themselves (Lee and Ng, 1994).
Late juveniles of the giant snakehead, Channa micropeltes, school and feed in packs
(Lee and Ng, 1991). Although there are few reports of early life history except for spe-
cies of commercial importance, it appears that as larval snakeheads mature to early
juvenile stages, the diet changes to small crustaceans and insects, particularly insect
larvae. Presence of phytoplankton, plant material, and detritus in the digestive system
of young snakeheads, as well as adults, appears to occur from incidental ingestion.
Juveniles frequently differ in colors and color patterns from late juveniles to adults,
making young of interest to some aquarium hobbyists (Lee and Ng, 1991, 1994).
• Respiration and overland migrations—Snakeheads are highly evolved
airbreathing teleostean fishes, and several are capable of overland migration by
wriggling motions (Lee and Ng, 1991; Berra, 2001; Peter Ng, personal commun.,
2002) despite the fact their pectoral fins lack spines like those of clariid catfishes.
They possess suprabranchial chambers for aerial respiration, and the ventral aorta is
divided into two parts to permit bimodal (aquatic and aerial) respiration (Das and
Saxena, 1956; Graham, 1997). The suprabranchial chambers become functional
during the juvenile stage of growth (Graham, 1997), following which some species of
snakeheads are obligate and others are facultative airbreathers. In some large species
of snakeheads, such as Channa marulius, the young are facultative airbreathers and
adults are obligate breathers (Wee, 1982), but all species are airbreathers.
These suprabranchial chambers lie above the pharynx and gill arches, lateral
to the otic chambers of the skull. In Channa, the chambers open into the pharynx
through inhalant apertures. The chamber lining contains respiratory “islets” with
vascular papillae. The chambers can be filled with air or water. In addition, in
C. striata, there are also vascular papillae in the epithelium of the mouth and pharynx
that can be utilized for respiration; these, however, can be retracted into depressions in
the epithelium to prevent damage when feeding (Munshi and Hughes, 1992).
Some channids, perhaps all, have a circadian rhythm in frequency of oxygen
uptake. Channa marulius, for example, showed a peak in oxygen uptake at night.
Channa striata and C. gachua peaked in early night hours, and C. punctata at dusk.
These rhythms are attributed to evolution in swamp ecosystems (that is, the rhythm is
a property of the ecosystem) (Munshi and Hughes, 1992).
The number of species of snakeheads capable of overland migrations is
unknown, but several display such behavior (Khin, 1948). These migrations often are
assumed to be the result of fish relocating from drying habitats in search of those with
water, perhaps driven by instinctive behavior for better feeding conditions, or both.

11
 Overland migrations likely apply to those species whose native range is subject to
seasonal dry/wet (or monsoonal) conditions, which encompass much of western to
southeastern Asia where the majority of snakehead species exist.

The species of Channa most capable of overland migrations are those that
are somewhat flattened ventrally (Peter Ng, personal commun., 2002). These include
C. asiatica, C. gachua, C. micropeltes, C. melasoma, C. nox, C. orientalis, and
C. striata. Even large C. micropeltes are capable of “crawling” in a sinuous motion on
dry or wet land, although movement is slow (Peter Ng., personal commun., 2002).
Those snakeheads with more rounded bodies (for example, C. argus, C. lucius, and
C. maculata) have very limited ability to move on land except as young, and only
when some water is present, as under mild flooded conditions. Liem (1987) noted that
Channa, like airbreathing catfishes (Clarias and Heteropneustes), do not migrate on
land to escape drying habitats, but burrow into mud to survive droughts. These species
only migrate during or soon after heavy rains, allowing these fishes to invade new hab-
itats, which permits a wider dispersal from more crowded environments (Liem, 1987).

• Hypoxic survival—Snakeheads are either obligate or facultative

airbreathers. Therefore, survival in hypoxic waters is not problematic to these fishes.
When prevented from access to the surface, adult snakeheads of many species will
drown due to lack of oxygen (Day, 1868; Lee and Ng, 1991). Cold temperatures
reduce metabolism as well as oxygen demand, allowing such species as Channa argus
to survive under ice (Frank, 1970). Moreover, snakeheads can remain out of water for
considerable periods of time as long as they remain moist. Some snakeheads,
especially C. striata, can bury themselves in mud during times of drought (Smith,
1945). They are known to secrete mucus that helps to reduce desiccation and
facilitates cutaneous breathing (Mittal and Banerji, 1975; Lee and Ng, 1991). Fishers
in Thailand are aware of this habit and, during drought periods, will slice into the mud
until they locate the fish (Smith, 1945).

• Lifespan—No specific information appears in literature. One species

(Channa marulius) is reported to reach a total length of 1.8 m in Maharashtra State,
India (Talwar and Jhingran, 1992), a size that would suggest a relatively long lifespan.
Nevertheless, we have been unable to find an ichthyologist who knows of preserved
specimens of such a length. The typical maximum length stated for C. marulius, the
largest snakehead, is 1.0-1.2 m. Nina Bogutskaya (personal commun., 2002) stated
she had seen a specimen of C. argus that was almost 1.5 m in length, also indicating a
relatively long-lived species. Moreover, Peter Ng (personal commun., 2003) reported
that C. micropeltes is known to reach 1.5 m in length. Smaller snakeheads, such as
members of the C. gachua and C. orientalis species complexes, may not live for more
than a few years. Most larger snakeheads are reported to reach sexual maturity within
2 years, after which growth slows but fecundity increases with increasing size. The
few publications that discussed growth rates in snakeheads based on examination of
scales or otoliths were inconclusive as to the interpretation of “growth” markings.
Moreover, timeframes of these studies were of such short duration (a few years) that
they documented no evidence of maximum lifespan.

12
 • Feeding habits—Few studies
analyze the feeding habits of snakeheads.
For those species studied, however, snake-
head fry feed mostly on zooplankton follow-
ing yolk-sac resorption. Munshi and Hughes
(1992) cited Banerji (1974) that fry of
Channa punctata feed on phytoplankton.
As juveniles, they feed on insect larvae,
small crustaceans, and fry of other fishes
(Munshi and Hughes, 1992). What is univer-
sal in reports of adult feeding habits is that
all snakeheads are predators, with many spe-
cies showing a preference for other fishes,
although they may also consume crusta-
ceans, frogs, smaller reptiles, and sometimes
young birds and small mammals. Welcomme
(1985) cited C. lucius, C. micropeltes,
School of young giant snakehead, Channa micropeltes,
C. pleurophthalma and C. striata as “large feeding at the surface in Thailand. Photo courtesy of
predators eating fish of all sizes, shrimps, Jean-Francois Helias, Fishing Adventures Thailand.
prawns and crabs.” Under conditions of food
depravation, snakeheads can become cannibalistic on their young. The piscivorous
nature of snakeheads has led to the use of some species (C. striata and Parachanna
obscura in particular) to control tilapia fish populations in aquaculture.

ASSOCIATED DISEASES AND PARASITES

Investigations of diseases and parasites of snakeheads concentrate on those
species of importance in aquaculture. Hoffman and Schubert (1984) noted that most
fishes can sometimes be hosts of parasites. Snakeheads are no exception.
Jinhui (1991) listed parasitic crustaceans of Channa argus, C. asiatica,
and C. punctata from Chinese waters. A listing of known parasites of C. gachua,
C. marulius, C. punctata, and C. striata from Bangladesh was provided by Arthus and
Ahmed (2002). In that study, parasites of all but C. gachua equaled or far outnum-
bered the parasites reported by Bykhovskaya-Pavlovskaya and others (1964) for
C. argus (table 2).
Literature on parasites of snakeheads includes numerous descriptions of
new species, not detailed herein, but indicates that most studies concentrate on
cultured fishes, such as Channa argus, C. punctata, and C. striata. Chiba and others
(1989) noted that C. argus and C. maculata introduced parasites to Japan, but did not
detail the parasites involved or fish species affected. None of the parasite literature we
reviewed on snakeheads indicated that any of these represent a potential threat to
native North American fishes. Conversely, such potential has not been examined.
A disease of snakeheads that has received broad attention is epizootic
ulcerative syndrome (EUS), which causes high mortality in these fishes, particularly
Channa striata and C. punctata under intensive culture. EUS involves several

13
 Table 2—Parasites of northern snakehead (Channa argus)
[Adapted from Bykhovskaya-Pavlovskaya and others, 1964]

Parasite Group Host tissues Other fishes affected

Myxidium ophiocephali Myxosporidia gallbladder, liver ducts

Zschokkella ophiocephalli Myxosporidia kidney tubules
Neomyxobolus ophiocephalus Myxosporidia gill filaments
Mysosoma acuta Myxosporidia gill filaments crucian carp
Myxobolus cheisini Myxosporidia gill filaments
Henneguya zschokkei ? Myxosporidia gills, subcutaneous, salmonids (tubercle disease
musculature of salmonids)
Henneguya ophiocephali Myxosporidia gill arches, supra-
branchial chambers
Henneguya vovki Myxosporidia body cavity
Thelohanellus catlae Myxosporidia kidneys
Gyrodactylus ophiocephali Monogenoidea fins
Polyonchobothrium ophiocephalina Cestoidea intestine
Cysticercus gryporhynchus cheilancristrotus Cestoidea gallbladder, intestine cyprinids, perches
Azygia hwangtsiüi Trematoda intestine
Clinostomum complanatum Trematoda body cavity perches
Pingis sinensis Nematoda intestine
Paracanthocephalus curtus Acanthocephala intestine cyprinids, esocids, sleepers,
bagrid catfishes
Paracanthocephalus tenuirostris Acanthocephala intestine
Lamproglena chinensis Copepoda gills

pathogens, including motile aeromonad bacteria (for example, Aeromonas hydrophila,

A. caviae, Pseudomonas fluorescens; Prasad and others, 1998; Qureshi and others,
1999), a fungus, Aphanomyces invadans (considered a primary pathogen; Mohan and
others, 1999; Miles and others, 2001), and perhaps a rhabdovirus (Kanchanakhan and
others, 1999; Lio-Po and others, 2000). Another bacterium, Aquaspirillum sp., also
has been implicated (Lio-Po and others, 2000). EUS may have originated in India in
the 1980s, but has since been found in Pakistan, Thailand, and the Philippines, with
outbreaks reported from all of these areas during the 1990s. Snakeheads are not the
only fishes affected by this disease. It is also known to occur in airbreathing catfish
(Clarias), the bagrid catfish genus Mystus, two cyprinid genera (Cyprinus and
Puntius), mastacembelid eels (Mastacembelus), and the nandid genus Nandus in India
(Mukherjee, 1998). In Thailand, it has been found in giant gourami (Osphronemus
goramy) and climbing perch (Anabas testudineus) during an outbreak in 1996-1997
(Kanchanakhan and others, 1999).
A parasitic disease that can affect humans is gnathostomiasis, caused by a
helminth parasite, Gnathostoma spinigerum. It has been recognized as a highly
important disease with about 800 suspected cases per year in two hospitals in Bangkok,
Thailand, between 1985 and 1988 (Setasuban, 1990). Channa striata has been identified
as an intermediate host for this parasite, found mostly in muscle tissue and occurring in
100 percent of fish examined over 41 cm in length (Setasuban and others, 1991). It is
unknown if additional species of snakeheads in Thailand and other countries of
southeastern Asia may serve as an intermediate host for larvae of this parasite.

14
HISTORY IN FISHERIES AND AQUACULTURE

Most snakeheads are part of capture fisheries. Few details were found in the liter-
ature on fishing methods, but most appear to involve hook and line, traps, gillnets, or seines.
Species for which we have found no information that they are of importance
as a fishery resource include Channa amphibeus, C. bankanensis, C. burmanica,
C. cyanospilos, and C. melasoma. Some do not appear to be targets of active fisheries,
but are believed or known to be periodically available in local markets as incidental
catches. These species include C. aurantimaculata (Musikasinthorn, 2000),
C. baramensis (Ng and others, 1996), C. barca (also in the aquarium trade; Talwar and
Jhingran, 1992), C. bleheri (wild caught for the aquarium trade; Ralf Britz, personal
commun., 2002), C. gachua (Talwar and Jhingran, 1992), C. harcourtbutleri (Ng and
others, 1999), perhaps C. melanopterus, C. nox (Zhang and others 2002), C. panaw
(Musikasinthorn, 1998), C. stewartii (Talwar and Jhingran, 1992), and possibly
Parachanna africana and P. insignis. Those for which there are active commercial fish-
eries are C. argus (Berg, 1965; Baltz, 1991; Dukravets, 1992), C. asiatica (Nichols,
1943; Daiqin and others, 1999), C. lucius (for food and aquarium purposes; Ng and
Lim, 1990), C. maculata (Nichols, 1943; Atkinson, 1977; Hay and Hodgkiss, 1981),
C. marulioides (aquarium purposes; Ng and Lim, 1990), C. marulius (Sriramlu, 1979;
Rao and Durve, 1989; Talwar and Jhingran, 1992), C. micropeltes (Lee and Ng, 1991;
Dudley, 2000), C. orientalis (Rainboth, 1996), C. pleurophthalma (Lee and Ng, 1991;
Kottelat and others, 1993; Dudley, 2000), C. punctata (Quayyum and Qasim, 1962;
Bhuiyan and Rahman, 1984; Rao and Durve, 1989; also in aquarium trade, Talwar and
Jhingran, 1992), C. stewartii (minor importance in India, also in aquarium trade; Talwar
and Jhingran, 1992), C. striata (Roa and Durve, 1989; Talwar and Jhingran, 1992), and
P. obscura (aquarium and food purposes; Dankwa and others, 1999) . In addition,
C. argus, C. maculata, and C. striata are commercially fished in most areas where these
species have been introduced. Interestingly, there are cultural differences in acceptance
of using introduced C. argus as a food fish. Within its native range in China, Korea, and
southern Siberia (Berg, 1965), and within its introduced range in Kazakhstan, Uzbeki-
stan, and Turkemistan, it is considered a desirable and sought-after food fish (Baltz,
1991; Dukravets, 1992; FAO, 1994); nevertheless, it failed to become popular following
its introduction to Japan in the early 1900s (Okada, 1960).
Snakeheads known to be cultured are summarized in table 3. The most
important and widely cultured species appears to be Channa striata. This may apply,
however, only within its native range and perhaps where it has been introduced into
southern China. It is becoming evident that it has been misidentified in places where this
species has been reported as introduced (Madagascar and Hawaii in particular), and the
introduced snakehead is C. maculata (Ralf Britz, personal commun., 2002). Thus, the
many reported introductions of C. striata to Pacific Islands summarized by Eldredge
(1994) and Lever (1996) will require reexamination. Channa maculata is the second
most important snakehead cultured in China (Fang Fang, personal commun., 2002), and
its culture appears to be concentrated primarily in Guangdong Province, where it is
native. During 2001, imports of snakeheads (likely C. maculata) into the U.S. increased,
the point of export having been Ghangzhou, Guangdong Province. Until identification of
introduced “C. striata” is verified, its reputation as the most widely cultured snakehead

15
 Table 3—Species of the family Channidae currently known to be cultured for food
and/or aquarium fish trade

Channa argus1 Channa maculata

Channa asiatica Channa micropeltes2
Channa marulius Channa striata3
Channa punctata Parachanna obscura
________________
1Second most important species cultured for food. Also, the only species that has been in culture
in the contiguous United States (Arkansas).
2Appears to be the most important species cultured for the aquarium fish trade.
3Species most widely cultured for food. Being cultured in Hawaii.

remains in question. We have verified that it is in culture in Hawaii (Qin and Fast,
1996a,b,c; Qin and others, 1997; Qin, Fast, and Kai, 1997; Qin, He, and Fast, 1997; Qin
and Fast, 1998; Pam Fuller, personal commun., 2002). Moreover, it is considered to be
the most important species economically in India (Bhatt, 1970), and is being cultured
there and in Thailand, Java (Hofstede and others, 1953); Vietnam (Pantulu, 1976; Bard,
1991); the Philippines (Conlu, 1986; Guerrero, 2000); Sri Lanka (Kilambi, 1986);
Pakistan (Talwar and Jhingran, 1992); Malaysia (Ali, 1999); and Cambodia (Balzer and
others, 2002).
Channa argus is the most important snakehead cultured in China (Fang
Fang, personal commun., 2002) where it is grown in ponds, rice paddies, and reser-
voirs (Atkinson, 1977; Sifa and Senlin, 1995; Liu and others, 1998). It was being
cultured on three fish farms in Arkansas until importation, culture, sale, and posses-
sion of snakeheads was prohibited by the Arkansas Fish and Game Commission in
August 2002.
Channa micropeltes is cultured for food in Vietnam (Pantulu, 1976; Wee,
1982), Malaysia (Lee and Ng, 1991), Thailand (FAO, 1994), and Cambodia (Rain-
both, 1996), and often in floating cages (Pantulu, 1976; Rainboth, 1996). Young of
this species are sold in the aquarium fish trade where this species, at least in the U.S.,
has been the most available snakehead.
Channa marulius is cultured in ponds, ricefields, and irrigation wells that do
not support other fishes in Pakistan and India (Bardach and others, 1972). Wee (1982)
noted that it is reared in monoculture in India, where it is fed tilapia. Mirza and Bhatti
(1993) contradicted Bardach and others (1972) in stating that this species is unsuitable
for aquaculture in Pakistan due to its highly piscivorous nature. Young have been avail-
able in the aquarium fish trade and are presumed to have originated from cultured stock.
Channa punctata has been an important food fish in India, where it is fished
commercially and reared in ponds (Quayyum and Qasim, 1962; Talwar and Jhingran,
1992). Some snakehead species, including C. punctata, used in intensive aquaculture,
are subject to outbreaks of EUS, and this has been reported for this species in India
(Prasad and others, 1998; Qureshi and others, 1999).
Parachanna obscura is being cultured in the Central African Republic
(Micha, 1974), Ondo State, Nigeria (Ajana, 1983; Victor and Akpocha, 1992), Benin
(Jackson, 1988), and Ghana (Morrice, 1991).

16
HISTORY OF INTRODUCTIONS

Eastern Hemisphere

Locations where snakeheads have been introduced beyond their native

ranges in the Eastern Hemisphere are shown in figure 3.

• Channa argus—Reported as introduced into “Czecho-Slovakia” and

Russia beginning in 1949 (Holcík, 1991). No specific localities of introduction or
information on status of the releases were provided by Holcík (1991). Nina
Bogutskaya (personal commun., 2002) reported early introductions that failed into the
Volga delta and ponds in Ekaterinburg (formerly Sverdlovsk) Province in the southern
Urals. An experimental introduction was made in ponds of Moscow Province during
1949-1950 that established. In 1953, it was recommended that the species be stocked
widely, but that failed to happen. There was a report in a Russian aquarium journal in
1963 noting occurrence of this species in small lakes in the Podolsk Region, Moscow
Province, but the species is presently absent from the Moscow area. Tandon (1976)
reported that acclimatization experiments were conducted in the former Soviet Union
after 1950, and that fry were collected from ponds near Moscow and the Ukraine in
1955 and sent to Czechoslovakia for acclimatization purposes. He concluded that the
source of the original stock was the Amur basin.

20°W 0° 20°E 40°E 60°E 80°E 100°E 120°E 140°E 160°E

60°N
Russia

Kazakhstan
Mongolia
North
40°N Korea Japan
Pak tan n-

China
is ha

Iran
ista
g
Af

South
Algeria Libya Korea
Egypt Saudi India PACIFIC
Arabia Taiwan
20°N
Burma
Sudan OCEAN
Vietnam
Philippines
Sri Thailand
Lanka
Malaysia
0° Kalimantan
Equator Sumatra
INDIAN
Indonesia
OCEAN Java

Madagascar
20°S
Australia

0 2,000 MILES
EXPLANATION
Introduced range of Channa 0 2,000 KILOMETERS
Scale is approximate

Figure 3—Introductions of snakeheads in the Eastern Hemisphere.

17
 The northern snakehead was introduced into the Aral Sea basin in the early
1960s where it became established in the Amu Dar'ya, Syr Dar'ya, and Kashka-Dar'ya
rivers of Kazakhstan, Turkmenistan, and Uzbekistan (Dukravets and Machulin, 1978;
Usmanova, 1982; Guseva, 1990; Dukravets, 1992). The initial introduction was
apparently accidental, with snakeheads mixed with shipments of Asian carps. All
introductions were apparently from Yangtze basin stock (Sal’nikov, 1998), although
some authors claim the stock came from the Amur basin and were purposeful releases
to establish a food resource. Dukravets (1992) recorded additional introductions that
became established in the Sarysu River, reservoirs on the Talus River, and Chu River
of Kazakhstan during the 1980s. He also reported that 10 metric tons of northern
snakeheads was harvested from reservoirs along the Talus River in 1989.

The northern snakehead was introduced from Korea in the early 1900s and
became established in many waters of central and southern Japan (Okada, 1960;
Nakamura, 1963; Uyeno and Akai, 1984), which includes the islands of Hokkaido,
Honshu, Kyushu, and Shikoku (Hiroshi Ueda, personal commun., 2002).

• Channa asiatica—Introduced and established in Taiwan

(Musikasinthorn, 2000), probably released as a food resource.

• Channa gachua—Ismail (1989), misidentifying this snakehead as

C. orientalis (Ralf Britz, personal commun., 2003), included Kalimantan (southern
Borneo) in the native range of this species, and Kottelat (1985) noted its presence in
the Greater Sunda Islands of Indonesia (probably including Kalimantan). This may
represent an introduction, but the rationale for introducing this very small snakehead
is not apparent. Myers and Shapovalov (1932) recorded this species in Taiwan.

• Channa maculata—Introduced and established as a food resource in

Taiwan; Nara, Hyôgo, Hiroshima, Mie, and Shiga prefectures, Japan; and the
Philippines (Okada, 1960; Liang and others, 1962; Hay and Hodgkiss, 1981; Uyeno
and Akai, 1984). Ralf Britz (personal commun., 2002) confirmed that it is this species,
not C. striata, that is established in Madagascar. He has also identified this species as
present in Oahu, Hawaii, since about 1900, based on specimens examined at the U.S.
National Museum of Natural History.

• Channa melasoma—Perhaps introduced and established in Palawan,

Philippines (Kottelat, 1985). Pathway and rationale unknown.

• Channa micropeltes—We believe the presence of the giant snakehead in

Kerala State, southwestern India, and described by Day (1865a) as Ophiocephalus
diplogramma was the result of an introduction from southeastern Asia that occurred
prior to the mid-1800s.

• Channa punctata—Smith (1950) reported this species as introduced in

the vicinity of Delagoa Bay, southern Mozambique. Paul Skelton (personal commun.,
2001) stated that no snakehead has been found or reported from southern Africa since
the Smith (1950) record.

18
 • Channa striata—This species has been reported as the most widely
introduced species of snakehead. It was recorded as introduced and established in
Madagascar (Raminosoa, 1987; Reinthal and Stiassny, 1991; Stiassny and Raminosoa,
1994; Lévêque, 1998), although Ralf Britz (personal commun., 2002) stated that this
was a misidentification of C. maculata. The chevron snakehead is also recorded from
the following locations, although some of these records may prove to be C. maculata,
misidentified as C. striata: Mauritius (Parameswaran and Goorah, 1981; Welcomme,
1988, Lever, 1996); Philippines (Seale, 1908; Herre, 1924, 1934; Conlu, 1986);
Vogelkop Peninsula, Papua, Indonesia (Allen, 1991); Sundaland, Sulawesi, Lesser
Sundas, Moluccas (Welcomme, 1981; Kottellat and others, 1993; Lever, 1996).
Kottelat and others (1993) reported introductions into China but gave no specific
localities. Its presence in Papua, Indonesia, was confirmed through photographs
supplied by Gerald L. Allen (personal commun., 2002). The species was also intro-
duced into Fiji and New Caledonia where establishment is questionable, and its intro-
duction to Guam failed (Maciolek, 1984). Channa striata is regarded as a prized food
fish in many parts of southeastern Asia, and in some localities its oils are used to heal
wounds and prevent scarring. Introductions beyond its native range were primarily to
establish a live-food resource.

Western Hemisphere

Five species of snakeheads have been reported from open waters of the
United States (California, Florida, Hawaii, Maine, Maryland, Massachusetts, North
Carolina, Rhode Island, and Wisconsin), and three became established as reproducing
populations (fig. 4). One species was being cultured in Arksansas until possession of
live snakeheads was prohibited in August 2002 and a fifth species is under culture in
Hawaii.
• California •
A northern snakehead, Channa argus, was collected by California
Department of Fish and Game personnel by electrofishing in a reservoir, Silverwood
Lake, October 22, 1997. This represents the earliest known record of a live snakehead
captured from open waters of the western United States. Silverwood Lake is in the
Mohave River drainage, east-northeast of Los Angeles and north of San Bernardino in
the San Bernardino Mountains, and receives water from the California Aqueduct. The
specimen was subsequently frozen and, apparently, later discarded (Camm Swift,
personal commun., 2002). A photograph of the 71 cm specimen that weighed 3.4 kg
was taken, which allowed identification of the fish (John Sunada, personal commun.
to Camm Swift, 2002). It remains unknown if the snakehead was released into
Silverwood Lake or arrived through the California Aqueduct. The aqueduct has been
the source of other fishes in the reservoir, including inland silverside (Menidia
beryllina), striped bass (Morone saxatilis), bigscale logperch (Percina macrolepida),
and tule perch (Hysterocarpus traskii) (Swift and others, 1993; Camm Swift, personal
commun., 2002).

19
 Hawaii

EXPLANATION
STATE WHERE SNAKEHEADS WERE CULTURED−Prior to August 2002
STATE WHERE SNAKEHEADS HAVE BEEN COLLECTED−In open waters
Channa maculata−Established before 1900 in Oahu, Hawaii
Channa marulius−Established in Broward County, Florida

Figure 4—States where snakeheads have been collected from open waters, were cultured prior to
August 2002, or are established.

• Florida •
An established population of the bullseye snakehead, Channa marulius, was
discovered in residential lakes and adjoining canals in Tamarac, Broward County,
Florida, in 2000 (Florida Fish and Wildlife Conservation Commission, 2001). It is
unknown how long this species has occupied these waters, perhaps several years, but
both juveniles and adults have been collected, indicating reproductive success. This
species is a large snakehead with adults commonly reaching lengths of 120-122 cm
(Talwar and Jhingran, 1992). In Maharashtra State, India, it can reach a length of
1.8 m and a weight of 30 kg, and was observed to reach a length of 30 cm in 1 year
(Talwar and Jhingran, 1992).

The pathway for the introduction in Broward County, Florida, is unknown.

The species may have been purposefully introduced to establish a food or aquarium
fish resource or could have been released accidentally by aquarists, in which case
several must have been released almost simultaneously. Because this species is
considered as a game fish in its native range (http://www.fishingasia.com), it also
could have been released illegally to establish a new sport fishing resource. Tamarac is
located just east of Water Conservation Area II, north of Everglades National Park,
and interconnected canal systems lead into this area. It is likely that Channa marulius
will expand its range in peninsular Florida as its native range includes tropical to
temperate climates. The bullseye snakehead is considered predacious (Jhingran, 1984;
Talwar and Jhingran, 1992), especially on other fishes (Schmidt, 2001).

20
 The northern snakehead, Channa argus, is also reported from Florida
waters. Two individuals were caught in the St. Johns River below Lake Harney,
Seminole and Volusia Counties in 2000, with unconfirmed reports of an additional
three individuals caught nearby. An attempt to collect additional specimens by Florida
Fish and Wildlife Conservation Commission (FFWCC) and U.S. Geological Survey
(USGS) personnel by electroshocking in 2001 was unsuccessful, but will be repeated.
Until reproduction has been confirmed, we consider the species as present but not
established. This fish is not involved in the aquarium fish trade but has been sold in
ethnic food markets as a food fish. The most likely pathway is introduction of live-
food fish, perhaps to establish a local source.
A live northern snakehead was purchased in a live-food fish market in
Orlando, Florida, in March 2002, another indication of its availability in a state where
possession is illegal. Moreover, we found a few U.S. aquarium fish retailers that sell
snakeheads via the Internet. Three species were purchased from a reputable dealer in
Rhode Island who requested a copy of our permit to possess certain restricted fishes,
including snakeheads. Private purchases can also be made through several Internet
“chat rooms” where possession of permits is doubtlessly of no concern.
• Hawaii •
The blotched snakehead (Channa maculata), misidentified as the chevron
snakehead (C. striata), has been established on Oahu, Hawaii, since the late 1800s,
possibly introduced from southern China (Herre, 1924). For whatever reasons, it does
not appear to have been introduced into other waters of Hawaii, although Morita (1981)
reported the species from Kauai. It is now mainly confined to one or more reservoirs on
Oahu (Maciolek, 1984). Ralf Britz (personal commun., 2002) has examined two
specimens at the U.S. National Museum of Natural History, labeled as C. striata that
were collected about 1900, and confirmed that they are C. maculata. We have examined
other specimens, collected in the early 1900s on Oahu, borrowed from the American
Museum of Natural History in New York City, Bernice P. Bishop Museum in Honolulu,
the California Academy of Sciences in San Francisco, and the Field Museum of Natural
History in Chicago, and they, too, are C. maculata. Two photographs, reported to be of
C. striata in Yamamoto and Tagawa (2000) from Hawaii, are that of C. maculata. It was
those photographs that alerted us to the likelihood that C. maculata existed in Hawaii.
We believe that all past published records of C. striata in Hawaiian waters were based
on misidentifications of C. maculata.
Channa striata is now being cultured as a food fish on Oahu, first imported
in the early 1990s under permit to Arlo Fast of the University of Hawaii (Domingo
Cravalho, Jr., personal commun. 2002). Peter Ng (personal commun., 2002) reported
that he saw C. striata in a market in Honolulu in recent years. Pam Fuller (USGS,
Gainesville, Florida) purchased five chevron snakeheads in Honolulu in September
2002. This species is regarded as carnivorous with a preference for other fishes
(Mohsin and Ambak, 1983; Conlu, 1986), and was described as a “territorial ambush
feeder” (Lee and Ng, 1991). Chevron snakeheads are used to control tilapia popula-
tions in culture ponds in the Philippines (Conlu, 1986; Milstein and Prein, 1993), and
is one of the species of snakeheads capable of overland migration (Peter Ng., personal
commun., 2002).

21
 • Maryland •
A northern snakehead, Channa argus, was caught by an angler in a 1.8-ha
pond in Maryland on May 18, 2002 (Beth Rogers, personal commun., 2002). The
angler, unable to identify the fish, took three photographs of the specimen before
releasing it into the pond. Estimated total length of the specimen was 43-45 cm. On
June 30, 2002, another angler captured a larger (66-67 cm) specimen from the same
pond and dipnetted eight juveniles from the pond on the evenings of July 7-8.
Maryland Department of Natural Resources personnel subsequently captured over
100 juveniles from the pond, proving that a well established population was present.
When the pond was treated with rotenone (a pesticide used for fish management, as
well as other uses) in August 2002, more than 1,200 northern snakeheads were
recovered (Bob Lunsford and Steve Early, personal commun., 2002). In addition, at
least three specimens of the giant snakehead, C. micropeltes, have been caught in
Maryland waters in recent years (Bob Lunsford, personal commun., 2002). Presence
of this subtropical/tropical species in Maryland waters where it could not overwinter
likely resulted from releases by aquarists.

Maryland DNR fishery biologist

Bob Lunsford examines a berm of
sandbags and silt fences strategi-
cally placed to prevent pond water
overflow or overland fish migra-
tion from Crofton Pond to the
Little Patuxent River. Photo by
Walter R. Courtenay, Jr., USGS.

Biologists dipnetting for snakeheads.

Photo by Tom Darden, Governor’s
office, Annapolis, Maryland.

22
 • New England States • Northern Snakehead
Distinguishing Features
A specimen of the northern snakehead, Long dorsal fin small head large mouth big teeth
Channa argus, was collected in October 2001 length up to 40 inches weight up to 15 pounds

from Newton Pond, Sudbury, Worcester County, HAVE YOU SEEN THIS FISH ?
Massachusetts, by Massachusetts Department of
Fish and Wildlife personnel (Hartel and others,
2002). The likely source is live-food fish markets,
as this species was the most common snakehead
available in ethnic food markets and restaurants
as a live-food fish. Moreover, it is capable of The northern snakehead from China is not native to
establishment in most freshwaters of the United Maryland waters and could cause serious problems
if introduced into our ecosystem.
States. Okada (1960) reported adults as voracious
If you come across this fish,
feeders, particularly on other fishes.
PLEASE DO NOT RELEASE.
Specimens of the giant snakehead, Please KILL this fish by cutting/bleeding
as it can survive out
Channa micropeltes, have been collected from of water for several days and REPORT all catches to
Maryland Department of Natural Resources
open waters in Maine, Massachusetts, and Rhode Fisheries Service, Thank you.
Island (Courtenay and others, 1984; Fuller and
Phone: 410 260-8320
others, 1999). This tropical/subtropical species TTY: 410 260-8835
Toll Free: 1 877 620-8DNR (8367) Ext 8320
could not establish itself in those temperate waters E-mail: [email protected]
(Hartel and others, 2002). Juveniles of the species
are cardinal red with two dark stripes on either Example of a poster alerting the public about the
side of the body, and are sold by aquarium fish invasive, non-native snakehead (this poster
retailers as red or redline snakeheads. Aquarist- courtesy of the Maryland Department of Natural
oriented websites note that this species requires Resources).
much animal food and that growth is rapid. These sites often advise that once these fish
reach about 15-20 cm in length, no more than one individual should be kept in a single
aquarium because they are aggressive predators. The pathway for release into these
New England States was likely aquarists when their “pets” grew too large for their
aquaria and/or because of the costs of feeding them. Releases of this species into sub-
tropical waters in southern Florida or Hawaii could lead to establishment of this snake-
head, regarded as the most predacious channid and known to have attacked humans
(Ng and Lim, 1990; Lee and Ng, 1991; Kottelat and others, 1993).
An angler reported having caught two blotched snakeheads, Channa macu-
lata, from a bridge over the Charles River in Boston in late July 2002 (Karsten Hartel,
personal commun., 2002). We confirmed that two live snakeheads purchased in an
ethnic market in Boston in October 2001 by Karsten Hartel were C. maculata, thus
proving local availability of this species at that time.

• North Carolina •
On July 31, 2002, two anglers reported catching two northern snakeheads from
Lake Wylie, a reservoir on the Catawba River, Mecklenburg County, North Carolina
(Wayne Starnes, personal commun., 2002). North Carolina Wildlife Resources Commis-
sion biologists sampled the lake using electrofishing equipment on August 14, 2002, but
failed to find any snakeheads (Russell Wong, personal commun., 2002).

23
 • Wisconsin •
A single specimen of the giant snakehead, Channa micropeltes, was
captured by personnel of the Wisconsin Department of Natural Resources in the Rock
River near Beloit on September 4, 2003 (Karl J. Scheidegger, personal commun.,
2003). This species would not overwinter in Wisconsin and was undoubtedly released
by an aquarist.

USES
AQUARIUM TRADE
Aquarists in Japan, Europe, and, to a lesser extent, North America
have kept snakeheads as pet fish. Because these fishes are predators (some
growing quite large) and high costs are involved with providing preferred live
food, few hobbyists become dedicated snakehead enthusiasts. Judging from
questions asked in 2002 on various Internet chat rooms dealing with aquarium
fishes in general and predatory species in particular, interest in snakeheads seemed to
be concentrated among a small number of serious collectors and a slightly larger
group of amateur aquarists curious about keeping predators as pets. Most questions
appeared to originate from persons who had experience with cichlid fishes, were
curious as to whether snakeheads could coexist with other fishes (particularly
cichlids), and wanted to know how to maintain snakeheads, what to feed them, what
species could be purchased, and where they could be bought. Experienced hobbyists
typically cautioned that large aquaria are needed for several available species, that
larger snakeheads were intolerant of other fishes and typically another of their own
species, and warned of the expenses of providing live food.
Snakeheads that have been periodically available to hobbyists in the U.S. are
listed in table 4. This information was assembled in 2002 from various Internet sites in
the U.S. and Canada that represent retailers and hobbyist groups and should not be
considered a complete “shopping list.” Moreover, availability of snakehead species
from retailers has been often sporadic. For example, two retailers located in states
where importation and possession of snakeheads is legal and who advertise on the
Internet rarely have them in stock for sale. A visit
to one of these dealers in June 2002 found no
snakeheads. A salesperson said they only have
them available periodically, adding that among
the most popular are “red” snakeheads (a “trade
name” for juvenile Channa micropeltes, a species
that can reach a length of 1 m as an adult and is a
voracious predator). Snakeheads have been only
a minor part of the aquarium fish trade in the U.S.
(Marshall Myers, personal commun., 2002).

Retail aquarium store, Nashville, Tennessee.

Photo by Walter R. Courtenay, Jr., USGS.

24
 Table 4—Snakeheads of interest to aquarists in the U.S.
[Information assembled from several aquarist-oriented and retailer
websites in 2002. Common names are those often used in the
aquarium fish trade]

Channa asiatica Chinese snakehead

Rainbow snakehead, tiger
Channa bleheri
snakehead, python snakehead
Channa marulius Cobra snakehead
Channa micropeltes Red snakehead, redline snakehead
Channa punctata Spotted snakehead
Channa stewartii Golden snakehead
Parachanna africana African snakehead

Nevertheless, hobbyists, wholesalers, and retailers have been able to import snakeheads
from many exporters in India and southeastern Asia that advertise on the Internet.
Individual hobbyists occasionally advertised snakeheads for sale, whereas others
inquired about availability on Internet aquarium fish “classifieds” and chat rooms;
sometimes these ads or inquiries originated in states where possession of snakeheads is
illegal.
Ng and Lim (1990) noted that smaller, colorful snakeheads are important in
the aquarium fish trade in southeastern Asia. For example, they mentioned that
Channa gachua was selling for S$30-60 per individual, and that slightly larger species
such as C. melanoptera and C. pleurophthalma from Borneo, Sumatra, and Malaysia
can garner prices as high as S$100 per fish. These fishes are caught wild, are primarily
found in forest streams, and with deforestation occurring at a rapid pace, there is fear
of overexploitation (Ng and Lim, 1990). In the U.S., prices for C. bleheri have ranged
from $55-75 per individual for sizes of 8-15.5 cm specimens. Larger species of snake-
heads can cost well over $100 per fish, depending on size, with young individuals of
the same species fetching prices of $15 or more. Therefore, with their predatory
nature, periodic availability, and relatively high prices for purchase and maintenance,
snakeheads cannot be considered an important staple of the U.S. aquarium fish industry.
Because 14 states prohibited importation and possession of live snakeheads
prior to the Federal ban on importation and interstate transport, the potential aquarium
market for these fishes would appear to be limited. Nevertheless, there have been
violations of these prohibitions in several states (see section “Regulations as of
July 2002”).
Larger snakeheads can outgrow their aquaria and/or the interest of their
owner(s). Some have been released, as witnessed by the capture of Channa
micropeltes from the wild in waters of Maine, Maryland, Massachusetts, and Rhode
Island (see section “Literature Review and Background Information, History of
Introduction, Western Hemisphere”). Fortunately, this subtropical species cannot
overwinter in these states.

25
 LIVE-FOOD FISH TRADE

Snakeheads have long been favored food fishes in India and many parts of
Asia, particularly southeastern Asia (Lee and Ng, 1991). Some are utilized as luxury
specialty foods, available alive in aquaria for customer selection at upscale restaurants
in larger cities such as Calcutta, Bangkok, Singapore, Hong Kong, and other major
locales. They also provide easily caught food for poorer people (Wee, 1982). These
fishes are typically freshly killed, often cooked whole or prepared as filets or steaks,
fried or steamed, or included in soups. Excess catches in Thailand and Cambodia are
often dried for storage and future use (Wee, 1982; Balzer and others, 2002).
Lee and Ng (1991) noted that snakeheads can remain alive out of water for
long periods of time if kept moist. They added that some people believed that this
ability may have provided these fishes with healing properties, making them prized as
food, particularly to people with illnesses or recovering from surgery. In such situa-
tions, the fish are killed just before cooking, the thinking being that healing properties
are lost if the fish are killed sooner. They also mentioned that some people in Myan-
mar believed that one species of snakehead represented humans transformed into fish
because of their sins, and that eating one would result in the consumer becoming a
lion. Day (1875) noted that some people in India believed that snakeheads that
suddenly appeared from mud in the bottom of dried ponds after monsoonal rains
actually fell from the sky with the rains.
To illustrate the value of snakeheads in the Orient, Wee (1982) recorded
1977 market prices as $2.50/kg in Taiwan and $1.00/kg in Hong Kong. Ng and Lim
(1990) reported prices of S$10 to S$20/kg for live Channa lucius, C. micropeltes, and
C. striata, three of the larger snakeheads, in markets in Singapore in the late 1980s.

Asian food market in Nashville, Tennessee, showing tubs and aquaria containing live-food fishes, June 2002.
Photo by Walter R. Courtenay, Jr., USGS.

26
They commented that even smaller
species (such as C. gachua and
C. melasoma) are utilized as food in
some parts of Asia (such as Myanmar
and Sri Lanka), adding that
C. gachua is sometimes used as bait
to catch larger snakeheads. Peter Ng
(personal commun., 2003) com-
mented that C. gachua was sold alive
in May 2000 in fish markets and
restaurants in Xishuangbanna and
Luxi, Yunnan Province, China, for
about S$5/kg, and served in restau-
rants as whole fish soup. He also
noted that C. marulioides was sold
fresh and dry salted in Sambas and
Fish market sign at an Asian food market in Honolulu,
Sintang markets, western Kaliman- Hawaii, showing snakehead availability. Photo by
tan, in April 1998. He also saw Pam L. Fuller, USGS.
C. marulioides being sold alive in
November 1999 in Samarinda, east Kalimantan, and, in 2002, freshly sliced fillets of
C. micropeltes were sold at S$20/kg in Singapore. In cities such as Hong Kong and
Singapore, snakeheads are imported for food from Malaysia and Indonesia. This
appears to have led to declines and scarcities of such species as C. striata in Malaysia
where this fish is considered the most valuable and important snakehead in the nation
as food and for medicinal purposes (Wan Ahmad, personal commun., 2001).

As noted earlier, some snakeheads, particularly Channa striata, called

“haruan” in Malaysia, are important for medical use, particularly in Malaysia and
Indonesia. Ng and Lim (1990) and Lee and Ng (1991) noted that C. lucius and
C. micropeltes are also utilized for such purposes in both nations. Mention was made
of use in a postnatal diet and during recuperation from illnesses or surgery (Lee and
Ng, 1991). Although no specific information was given as to how the fishes were used
following surgery, a neighbor of one of the authors (WRC), a Malaysian by birth,
stated that the oils from the haruan are used to greatly reduce scarring following
surgery, adding she had seen the results and scar tissue was dramatically reduced to a
minimum. It has been demonstrated that haruan tissues contain substantial levels of
arachidonic acid, a precursor of prostaglandin, essential amino acids (especially
glycine), and polyunsaturated fatty acids required to promote prostaglandin synthesis,
important factors in wound healing (Baie and Sheikh, 2000).

Channa striata is an important food fish throughout its native range

(a species complex distributed from Pakistan eastward to southern China including
Malaysia, Indonesia, and Java). Its supposed medicinal value doubtlessly explains
why this species is often said to be the most widely introduced species of snakehead
as persons of Asian origin emigrated to other locations. Channa maculata, often
misidentified as C. striata, have been imported and released in a similar manner.

27
 BIOLOGICAL CONTROL

There is no evidence that any species of snakehead has been suggested for
use as a biological control agent in the U.S. Nevertheless, certain species of snake-
heads have been investigated or utilized as biological controls abroad. For example,
Kehar and others (1995) reported experiments in which spotted snakeheads, Channa
punctata, were used at different levels of pH and salinity in controlling mosquito
larvae. They concluded that this snakehead could be utilized as a potential biological
control of mosquito larvae in waters up to 10 ppm salinity. Mansuri and others (1979),
however, determined that this species was intolerant of salinities above 6 ppm. Never-
theless, Khora and Rao (1994) recorded the spotted snakehead from estuaries entering
the Bay of Bengal. Only young spot-
ted snakeheads feed on insect larvae
before dietary changes to larger prey
(Quayyum and Qasim, 1962).
A more common use of
snakeheads in biological control has
been as a predator in fish culture.
Conlu (1986) and Milstein and Prein
(1993) reported that chevron snake-
heads, Channa striata, are used to
control overproduction of tilapias in
culture ponds in the Philippines. Wee
Predatory teeth and jaws of a snakehead.
(1982) also noted this practice, add- Photo by Walter R. Courtenay, Jr. USGS.
ing, as did Bardach and others (1972),
that they are also used in carp culture
to control unwanted “pest fish” in culture ponds. In turn, snakeheads utilized in this
manner are also sold as products of culture. Similarly, African snakeheads,
Parachanna obscura, are used to control young of tilapias in aquaculture ponds in
Bénin (Bonou and Teugels, 1985).

U.S. IMPORTATIONS
Some species of snakeheads have been imported for several decades
for the aquarium fish trade. In the past two decades, however, snakeheads have
also been imported to the U.S. for sale in certain ethnic markets that sell live-
food fishes and some restaurants that hold live fishes in aquaria for customer
selection. In most states and the District of Columbia, such importation and
sale is legal, but there have been violations in at least six states where
possession and sale of live snakeheads is illegal. Until recently, the live-food
fish pathway for potential introduction of live fishes into U.S. waters was largely
overlooked. Because Channa maculata (misidentified as C. striata) and other eastern
Asian food fishes (common carp, Cyprinus carpio; oriental weatherfish, Misgurnus
anguillicaudatus; whitespotted clarias, Clarias fuscus; and swamp eel, Monopterus
albus) were introduced from China into Hawaii before 1900, doubtlessly encouraged
and perhaps initiated by Asian immigrants (Maciolek, 1984), this pathway should
have been of concern.

28
 Imports of snakeheads into the U.S. have been increasing in recent years
(table 5). Importation records unfortunately report quantities either in numbers or by
weight, but not both. Hence, the two columns (Number of individuals, Number of
kilograms) in table 5 consist of 51,233 fish and an additional 22,208 kg of snakeheads.
Furthermore, records do not provide a detailed breakdown of species imported or
indication of the intended reason for importation (pet trade or live-food fish markets).
Moreover, these records are probably incomplete (Marshall Myers, personal commun.,
2001) and may represent only part of the total number/weight of imported snakeheads.
This, coupled with not knowing how much of the weight represents small snakeheads
and juveniles of larger species destined for the aquarium fish trade versus market-size,
larger fish, makes projecting the total number of individuals a precarious guess at best.

Table 5—U.S. importations of live snakeheads (Channidae, all

species) during 1997-2002

Total declared
Number of Number of $ value
Year
individuals1 kilograms2 (individuals and
weight combined)

1997 372 892 5,085

1998 1,488 1,883 12,632
1999 6,044 8,512 27,718
2000 8,650 9,240 39,990
2001 18,991 1,681 21,185
20023 15,688 -- 26,077
Totals 51,233 22,208 $132,687
1 Not included in number of kilograms.
2 Not included in number of individuals.
3 Data are for January–May 2002.

Sources of imported snakeheads are varied (table 6). Again, these records
are probably incomplete, but China is clearly the major exporter of live snakeheads.
As in table 5, there is no breakdown by species.
The number of species that have been imported for the aquarium fish trade
or the live-food fish trade could not be determined. Nevertheless, Channa argus is the
most widely cultured snakehead in China (Fang Fang, personal commun., 2002), and
has been available for sale in ethnic live-food fish markets in New York (James
Stephen Lee, personal commun., 2001) and St. Louis, Missouri (Leo Nico, personal
commun., 2002). A total of 80 live individuals in transit to Seattle were confiscated in
Blaine, Washington, in 2001 (Ted Pietsch, personal commun., 2001; Mike Williams,
personal commun., 2003), and others were seized from markets in Houston, Texas
(Howells and others, 2002), Miami and Pembroke Pines (Florida Fish and Wildlife
Conservation Commission, 2001), and Orlando, Florida. Snakeheads had been illegal
in California, Florida, Texas, Washington, and 10 other states for many years prior to

29
 Table 6—Origin of snakehead shipments (Channidae, all species)
during the past 5 or more years
[1997–2002; records for 2002 extend through May 31]

Total declared
Number of Number of $ value
Country
individuals 1 kilograms 2 (individuals and
weight combined)

China 48,533 20,323 125,295

Hong Kong 2 -- 50
India 572 -- 1,498
Indonesia 300 -- 96
Nigeria 970 -- 659
Switzerland 50 -- 100
Thailand 1,084 -- 1,420
United States 25 -- 38
Vietnam 1,079 1,435 4,265

1Not included in number of kilograms.

2Not included in number of individuals.

July 2002. The first specimen of this species to have been captured from U.S. waters
was taken by electrofishing in Spiritwood Lake, a reservoir north of San Bernadino,
California, in 1997. Two individuals were caught by angling in the St. Johns River,
below Lake Harney, Seminole and Volusia Counties, Florida, in 2001 (with three more
reported as having been caught nearby); another was captured by electrofishing in
Newton Pond, Worcester County, Massachusetts, in late 2001. The discovery of an
established population of this species in a pond in Crofton, Anne Arundel County,
Maryland, proved that this species was capable of invading U.S. waters. In July 2002
C. argus was being sold in a live-food fish market in Orlando, Florida. That market
was raided by FFWCC agents who confiscated several specimens. Northern
snakeheads were reported to be in culture in Arkansas, and this may or may not have
been a source of northern snakeheads in Florida. Channa argus was likely available in
live-food fish markets in Boston, although two snakeheads purchased there in late
2001 by Karsten Hartel were later identified as C. maculata. Live-food fish markets in
Vancouver, British Columbia, also sold C. argus (Margarita Reimer, personal
commun., 2002).

In conclusion, Channa argus is known to have been the most widely

available snakehead in North American live-food fish markets, followed by
C. maculata. Channa marulius was also available in New York City live-food fish
markets (Leo Smith, personal commun., 2002). There have been no reports of
C. striata being available for sale in live-food fish markets in the contiguous U.S.,
even though it is considered the most important snakehead used for food in
southeastern Asia and is being cultured in Hawaii. Nevertheless, a freshly killed

30
C. striata was purchased from an oriental market in San Diego, California, on July 29,
2002 (Richard Rosenblatt, personal commun., 2002). There was no cloudiness in the
eyes of the specimen, indicating that it had never been frozen and may have been kept
in a live fish tank, perhaps on the premises of the market, until a very short time before
being placed on ice for sale. The specimen was deposited in the fish collection of
Scripps Institution of Oceanography (SIO 64-228). Channa striata is being sold in a
market in Honolulu, Hawaii (Pam Fuller, personal commun., 2002).
Another observation from importation data shows that imports of live snake-
heads from Ghangzhou and Shenzhen, both in Guangdong Province, China, began to
increase in the latter part of 2001 and accounted for the majority of imports through
May 2002. Channa argus is not native to southern China. Therefore, importation data
suggest that many snakeheads imported during late 2001 well into 2002 may have
been C. maculata, destined for the live-food fish trade.

REGULATIONS AS OF JULY 2002

At least 14 states specifically prohibited possession of
live snakeheads (table 7) prior to the discovery of an established
population of northern snakeheads in a pond in Crofton, Anne Arundel
County, Maryland, which was eradicated in September 2002. Since
then, the states of Arkansas, Connecticut, Illinois, North Carolina,
Rhode Island, Pennsylvania, South Carolina, Tennessee, and Virginia
have made possession of live channids illegal (fig. 5). Indiana Department of Natural
Resources approved emergency fisheries regulations on November 22, 2002, that bans
possession of snakeheads effective December 1, 2002. Kansas Department of Wildlife
and Parks prohibited possession of snakeheads in early 2003.

The U.S. Fish and Wild-

life Service published a proposed
Table 7—States prohibiting snakeheads
rule to list the family Channidae
as of July 2002
(snakehead fishes) as injurious
Alabama Idaho wildlife in the Federal Register on
Arizona Mississippi July 26, 2002 (67 FR 48855) under
California Nevada the Lacey Act (18 U.S.C. 42). The
Colorado Oregon final rule banning importation and
Florida Texas interstate transport of live snake-
Kentucky Utah
heads was published in the Federal
Georgia Washington
Register on October 4, 2002 (67 FR
62193). This ruling does not affect
possession or sale of live snake-
heads in states that do not specifically prohibit them, or importation of dead snake-
heads refrigerated or frozen for sale as food fishes into states where possession of live
snakeheads is illegal. Nevertheless, despite the Federal rule and a long-standing state
prohibition, several live Channa argus were confiscated by U.S. Fish and Wildlife
Service Inspectors in California as recently as July 2003.

31
 Hawaii

EXPLANATION
NO LEGAL VIOLATIONS REPORTED−For laws related to snakeheads.
LEGAL VIOLATIONS REPORTED−For laws related to snakeheads. This would include
illegal activity, confiscations, citations issued, or investigations initiated within the last 2 years.

Figure 5—States prohibiting possession of live snakeheads as of November 2002.

Prior to 2002, there were illegal activities involving snakeheads in states that
prohibited their sale or possession. A total of 80 live Channa argus, destined for
markets in Seattle, was discovered in 2001 on a truck from British Columbia. Speci-
mens of C. micropeltes and C. marulius were confiscated from pet shops in the Los
Angeles area, southern California, in the past 2 years. Channa argus was confiscated
from live-food fish markets in Miami, Orlando, and Pembroke Pines, Florida, and
Houston, Texas, in 2001. Illegal traffic in pet snakeheads, involving mostly C. bleheri,
was discovered in Alabama and Kentucky in the past 2 years. It has been suggested
that these snakeheads came from a distributor in Atlanta, Georgia, a state where
snakehead possession is also illegal.

POTENTIAL RANGE
Temperature is the most important environmental factor that would
determine potential range of snakeheads in the United States. Because there
are few data providing thermal tolerance ranges for snakeheads, potential
range must be inferred from distribution within native ranges (fig. 6).
The family Channidae contains 10 species that are strictly tropical
and, if introduced, would survive in only the warmest waters, such as extreme southern
Florida, perhaps parts of southern California and Hawaii, and certain thermal spring
systems and their outflows in the American west. Another four species can be considered

32
tropical to subtropical, indicating a similar potential range of
distribution as for tropical species, but with a greater likelihood
of survival during cold winters and more northward limits. One is Channa amphibeus
subtropical. Another 11 snakeheads (3 that appear to be species Channa argus
complexes) can tolerate tropical or subtropical to warm temperate Channa asiatica
conditions, indicative of species that could survive in most south- Channa aurantimaculata
ern states. One is warm temperate, and another is warm temperate Channa bankanensis
to cold temperate (Channa argus has a temperature range of Channa baramensis
0-30 ºC). In summary, there are no waters in the United States Channa barca
that, based on temperature, would preclude some member(s) of Channa bleheri
the family Channidae from becoming established. Channa burmanica
Channa cyanospilos
Channa gachua*
RISK ASSESSMENT PROCESS Channa harcourtbutleri
Channa lucius
Snakehead fishes have had a modest Channa maculata
following among U.S. aquarists for several Channa marulius*
decades. Nevertheless, they are more popular in the Channa marulioides
Japanese and European aquarium fish trade (Ralf Channa melanoptera
Britz, personal commun., 2002). It is only within Channa melasoma
the past two decades that snakeheads, limited to a Channa micropeltes
few species, have been imported and marketed in the U.S. as live- Channa nox
food fishes. Thus, pathways for introduction into U.S. waters have Channa orientalis
been through importations and releases by aquarists and Channa panaw
introductions of market-size snakeheads, the latter typically at or Channa pleurophthalma
near sexual maturity. Release or introduction of snakeheads out of Channa punctata*
both of these pathways has been documented in open waters of Channa stewartii
the U.S. All snakeheads prey on other aquatic organisms, most Channa striata*
showing a preference for fishes, and many are regarded as thrust Parachanna africana
predators, hiding and attacking prey by surprise. As with any Parachanna insignis
introduction of a nonindigenous aquatic species, there is always Parachanna obscura
the possibility that they may host parasites or diseases that could
EXPLANATION
spread to native species. Some might host human parasites, and
TROPICAL
one snakehead species has been found to be a carrier for
SUBTROPICAL
gnathostomiasis. The fact that one species has been shown as a
carrier indicates that there are others which could present a WARM TEMPERATE

similar threat to human health, yet to be investigated. COLD TEMPERATE

There are seven rating elements in the risk model (Risk * Indicates a species complex

Assessment Management Committee, 1996). Each element is

assigned an estimated level of risk, rated as high, medium, or low. Figure 6—Thermal range of
Uncertainty codes after each element rating are as follows (with snakeheads (Channidae) based
descriptions): Very Certain (as certain as we are going to get); largely on native range of
Reasonably Certain (reasonably certain); Moderately Certain distribution.
(more certain than not); Reasonably Uncertain (reasonably
uncertain); and Very Uncertain (a guess).

33
 RATING ELEMENTS OF RISK MODEL

• 1 • Estimate probability of the exotic organism being on, with, or in the pathway.
High—very certain
Four species of snakeheads have been recorded as reproducing in waters
of the United States. These are Channa argus in Crofton, Maryland (isolated
population, eradicated in September 2002), C. maculata in Oahu, Hawaii, C. marulius
in southeastern Florida, and C. striata, being cultured in confined waters in Oahu,
Hawaii, since the early 1990s. Specimens of C. micropeltes have been collected from
waters of four states, the earliest records being from Maine and Rhode Island from the
1970s. Snakeheads have had a limited market in the aquarium fish trade for several
decades and, more recently, four species of snakeheads (C. argus, C. maculata,
C. marulius, and C. striata) were being sold in live-food fish markets within the U.S.
Therefore, snakeheads have been and are in the United States pathway.

• 2 • Estimate probability of the organism surviving in transit.

High—very certain
Snakeheads are capable of breathing air, many being obligate airbreathers,
and easily transported by air or land vehicle without water as long as they are kept
moist. They have survived importation from overseas as well as interstate truck
transportation.

• 3 • Estimate probability of the organism successfully colonizing and maintaining a

population where introduced.
High—very certain
Appropriate habitats and climate are found throughout most of the United
States. This does not infer that all species of snakeheads could become established in
most of the U.S., but that there are habitats in all states, with the possible exception of
Alaska, where one or more species could establish a reproducing population.
Preferred food of snakeheads (that is, fishes, crustaceans, insects and insect larvae) is
locally abundant.
Several species of snakeheads have established in waters outside their native
ranges of distribution in the Eastern Hemisphere. These include Channa argus in
Japan, Czechoslovakia, Russia for a period of time, the Aral Sea basin (Amu Dar’ya,
Syr Dar’ya, Kaska-Dar’ya, Sarysu, Chu, and reservoirs on the Talus rivers);
C. asiatica in Taiwan; C. maculata in Taiwan, several prefectures of Japan,
Madagascar, and Hawaii; C. melasoma on Palawan, Philippines; C. orientalis in
Kalimantan and Greater Sunda Islands; and C. striata in many Pacific Islands and
most recently (early 1990s) in confined waters of Oahu, Hawaii.
Within the continental U.S., two species of snakeheads have been recorded
as established. Channa argus was established in a pond in Crofton, Anne Arundel
County, Maryland, for at least 2 years before being eradicated in September 2002.
There remains some concern that the species may have escaped into the Little
Patuxent River during that period of occupancy. In addition, specimens of this species
have been collected from the St. Johns River, Seminole and Volusia Counties, Florida,

34
a pond in Shrewsbury, Worcester County, Massachusetts, and from a reservoir serving
Los Angeles, located just north of San Bernardino, California, in 1997. There was a
reported capture of two individuals of the same species from a reservoir near
Charlotte, Mecklenburg County, North Carolina. Subsequent sampling of that
reservoir by North Carolina Wildlife Resources Commission biologists did not reveal
the presence of additional specimens. Channa marulius has been established for
several years in a series of interconnected artificial lakes and canals in Tamarac,
Broward County, Florida. This system of waterways is connected to the gridwork of
flood control canals of southeastern Florida.
Channa maculata has been established since before 1900 on Oahu, Hawaii.
Although the species was once widely distributed on Oahu, it is now largely confined to
Wahiawa Reservoir and adjoining canal systems. Yamamoto and Tagawa (2000)
reported the largest snakehead captured from waters of Oahu was “over 5 feet in length,”
and that species had to have been C. maculata. Two specimens were also reported to
have been captured by an angler
from the Charles River, Boston,
Massachusetts, in 2002. Since the
early 1990s, C. striata was
imported into Hawaii and it is now
being cultured.
Channa micropeltes,
a species largely sold through the
pet fish trade, has been collected
from open waters of Maine, Mas-
sachusetts, Maryland, Rhode
Island, and Wisconsin. This tropi-
cal/subtropical species could not
survive winters in those states.
Nevertheless, these releases, likely
made by hobbyists, is indicative of
what could happen if similar intro-
ductions of this or other tropical/ Joe Hennessey with a giant snakehead (Channa micropeltes)
caught in a Wisconsin river, September 4, 2003, undoubtedly after
subtropical snakeheads were made
being released by an aquarium enthusiast. Fortunately, this species
in states, such as Florida or Hawaii, will not survive the winter (see fig. 6). Photo by Mike Sorge,
or into thermal springs and their Wisconsin Department of Natural Resources, Bureau of Fisheries
outflows in western states. Management and Habitat Protection.

• 4 • Estimate probability of the organism to spread beyond the colonized area.

High—reasonably certain
Appropriate habitats (rivers, streams, lakes, reservoirs, ponds, canals) and
climate are suitable for establishment of snakeheads in U.S. waters. Suitable habitat for
subtropical/tropical species exists in southern Florida, Hawaii, perhaps southern Texas,
and thermal springs and their outflows in several western states. Several snakehead
species can exist in warm temperate conditions that exist in southern states. Both
Channa argus and C. maculata, especially the former, can tolerate cold climates,

35
 making the likelihood of their becoming established a probability even in some
northern states if released. Introductions into rivers, streams, or canal systems would
likely spread whereas releases into lakes or ponds could be more restrictive as to range
expansion. Nevertheless, people move fish; considering that larger species of
snakeheads are popular with anglers in several locations within their native and
introduced ranges abroad, the likelihood of anglers moving snakeheads to novel
waters from colonized areas is reasonably great.

Because most snakeheads build nests in aquatic vegetation, some might

argue that these fishes would be incapable of colonizing waters devoid of macro-
phytes. Nevertheless, at least three snakeheads, Channa gachua, C. marulius, and
C. punctata, have successfully reproduced in waters lacking vegetation. The same
may be true for C. argus that has colonized reservoirs on the Talas River of Kazakh-
stan. This suggests that there is likelihood that other species of snakeheads have the
potential to establish in waters lacking vegetation. Predictions as to where or under
what environmental conditions a nonindigenous aquatic species might or might not
become established have been proven unreliable in several instances.

• 5 • Estimate economic impact if established.

Medium—moderately certain

The predatory nature of snakeheads indicates that their introduction could

negatively impact populations of native fishes through direct predation, competition
for food resources, and alteration of food webs. Larger species of snakeheads are
considered to be “top predators” in their native ranges. Unlike U.S. highly predatory
native fishes, snakeheads are very protective of their young, thus enhancing survival
beyond early life history stages and suggesting the possibility of eventual dominance
in suitable waters. To predict what the economic impact could be to the recreational
fishing industry or to sport fishing is difficult to assess, but could prove to be
substantially detrimental over time.

The economic cost of eradication efforts would be high. Introduction of the

northern snakehead, Channa argus, to a single pond in Crofton, Anne Arundel
County, Maryland, serves as an example. The original purchase of the snakeheads that
were eventually introduced at least 2 years ago was likely no more than $40. A recent
estimate of the costs to the State of Maryland during 2002 in personnel, creating and
conducting two meetings of the Maryland Snakehead Scientific Advisory Panel,
application of herbicides and rotenone, and disposing of dead fish was about $110,000
(Steve Early, personal commun., 2003). Introduction of non-native aquatic species is
illegal in Maryland, but the perpetrator must be found and charged of such action
within a 2-year period. In this instance, the time limitation had expired before the
individual making the introduction was identified. Had that person been charged
before the limitation expired, the fine would have been $40. At present, no state
requires a liability bond before an intentional introduction is made by individuals or
an agency, and there are no laws that hold an individual (or individuals) responsible
for the costs of eradicating or controlling an unintentional introduction should the
species involved become established.

36
 The northern snakehead introduction in Maryland was a rare instance where
the fish was confined to a single pond from which it could be eradicated. The costs of
eradicating an introduced species in an isolated small lake would be greater and could
be substantial in a larger lake. Eradication from flowing waters or large lakes with
connecting drainages is physically and fiscally impossible, and the same applies to
control measures.
Some species of snakeheads are capable of short overland migrations. This
presents a potential economic threat to fish culture interests if those species enter
culture facilities from adjacent waters, such as occurred with another introduced
airbreathing predator, the walking catfish, in Florida (Courtenay and Miley, 1975).

• 6 • Estimate environmental impact if established.

High—very certain
Because snakeheads do not occur naturally in the U.S., there is no
possibility of introduced snakeheads hybridizing or interbreeding with native fishes.
Conversely, competition for food resources is probably high. Competition for habitat
is probably low except during spawning seasons. Moreover, potential to cause habitat
degradation and/or destruction is low.
All snakeheads are predators, particularly on fishes. Therefore, negative
impacts to populations of native fishes could be quite high, as well as predation on
crustaceans. Predation on other invertebrate species would be moderate to low, based

A blue tilapia (Oreochromis aureus), an introduced species in Thailand, was sheared in half by a
giant snakehead (Channa micropeltes). Photo courtesy of Jean-Francois Helias, Fishing Adventures
Thailand.

37
 on literature references supplied in individual species accounts. Larger snakeheads,
however, are known to also feed on birds (particularly young waterfowl), amphibians,
small reptiles (snakes, lizards), and small mammals.

Potential to transfer pathogens (parasites, diseases) is largely unknown.

Nevertheless, all snakehead species are hosts to at least several species of parasites (see
table 2). At least two snakehead species utilized in intense aquaculture, Channa
punctata and C. striata, are susceptible to epizootic ulcerative syndrome (EUS), a
disease believed to be caused by several species of bacteria, a fungus, and perhaps a
retrovirus. EUS is not specific to snakeheads and has affected other fishes, such as
clariid catfishes, bagrid catfishes, two cyprinid genera, mastacembalid eels, a nandid fish
in India, and giant gourami and climbing perch in Thailand. There have been no studies
undertaken to examine transfer of parasites or diseases to native North American fishes.

Adverse impacts on native wildlife and wildlife resources would likely be

few, other than through predation. Ecosystem balance, however, could be substantially
modified should snakeheads become established in waters with low diversity of native
fishes and low abundance or absence of native predatory species.

Adverse impacts on threatened and endangered species would likely be

high. Of all the taxa listed as endangered or threatened in U.S. aquatic habitats, 16
amphibians, 115 fishes, and 5 of the 21 crustaceans (surface dwelling crayfish and
shrimp), would be the most likely to be affected. Based on habitat requirements and
life history, amphibians and surface dwelling crustaceans would generally be less
likely to be affected by introduced snakeheads than would fishes. The possibility of a
nonindigenous predator in the aquatic community with any listed amphibian or
crustacean would constitute a threat.

Likelihood and magnitude of the effect on designated critical habitats of

threatened or endangered species would be significant on the living component of the
aquatic ecosystem. Depending on the habitat, snakeheads have the potential to
detrimentally alter aquatic communities. The most likely scenario would be an
alteration of the fish and crustacean community structure through predation. For listed
fishes there could be competition for food in addition to direct predation. Like
amphibians, fishes and crustaceans listed as threatened or endangered species,
candidate taxa of these three groups or aquatic organisms would likewise be at risk.

Introduction of a small number of snakeheads (for example, less than five)

into isolated spring habitats could result in extinction of endemic spring-adapted
fishes or crustaceans. Introductions of fishes considered to be far less aggressive than
snakeheads (that is, guppies, Poecilia reticulata) in such habitats have had major nega-
tive impacts (Courtenay and others, 1985). Snakeheads would not have to establish a
reproducing population to reduce or eliminate a fish or crustacean species confined to
a small section of a stream or isolated spring habitat. A small number of snakeheads
introduced, but not established, in a stream or lake would likely have less of an
impact. Nevertheless, any snakehead that becomes established in a water body would
represent a significant threat and could potentially put any listed amphibian, fish, or
crustacean at risk of local extinction.

38
 There is a likelihood that damage to ancillary wildlife resources through
control measures could be substantial. Netting and/or electrofishing would be too
selective on size classes to remove a population of snakeheads, even in an isolated
situation. Despite preliminary fears that rotenone would be ineffective against
airbreathing snakeheads, the Crofton, Anne Arundel County, Maryland, eradication
program on Channa argus in September 2002 proved to be effective. Young northern
snakeheads captured from the pond were exposed experimentally to several different
ichthyocides, and rotenone did kill the fish. Nevertheless and as expected, when
rotenone was applied to the three adjacent ponds in Crofton, it also killed all other
fishes. An estimated 500 kg of native fishes died and were disposed of (Bob Lunsford
and Steve Early, personal commun., 2002). Control methods in a nonisolated pond or
lake, or in flowing water (streams, rivers) situations would be ineffective in
eliminating snakeheads whether or not they were established.

• 7 • Estimate impact from social and/or political influences.

Low—moderately certain
Snakeheads have been in the U.S. aquarium fish trade and hobby for several
decades. Due to their predatory nature, compounded by the high costs of housing and
feeding larger snakehead species, they have had a limited following by hobbyists.
Therefore, snakeheads have never represented more than a very minor component of
the U.S. aquarium fish trade. Consequently, economic impact to the aquarium fish
trade through prohibition of importation or interstate transport of live snakeheads
would be minor.

Importation of snakeheads for the live-food fish market in the mainland U.S.
is a more recent trend, to our knowledge dating back to the most recent decade or two.
Although snakeheads have been available in live-food fish markets in Hawaii for a far
longer period of time (likely several decades), only one market (in Honolulu) was
selling live snakeheads for food purposes as of 2002 (Mike Yamamoto, personal
commun., 2002). Markets that sell live freshwater food fishes also sell species other
than snakeheads, including catfishes, tilapias, carp, eels, hybrid striped bass, and
sometimes swamp eels. These are typically Asian ethnic food markets, and they
frequently carry a large variety of frozen, imported marine and freshwater food fishes.
Therefore, as in the aquarium fish trade, snakeheads are only a minor component of
live-food fish sales.

Economic impact to the live-food fish trade would be minor following a ban
on importation and interstate transportation of live snakeheads, as these fishes can be
imported frozen or dead on ice for sale. Until Arkansas passed an emergency rule
banning importation, possession, and sale of live snakeheads in late July 2002, only
three fish farmers in that state were reported to be culturing snakeheads (Channa argus)
for the live-food fish market. There were no other culture facilities in the mainland
U.S. known to be raising snakeheads. There is, however, one aquaculture facility on
Oahu, Hawaii, that has been rearing C. striata since the latter part of the 1990s.

39
 Fish produced by that facility can be shipped for sale either dead on ice or frozen to
any state or U.S. territory, or sold within Hawaii. As a result, Federal prohibition of
importation and interstate transport of live snakeheads would not present a significant
negative impact to most U.S. aquaculture interests. Although political entities could
be negatively impacted by the costs of eradication or attempts to control introduced
snakeheads, no political entity is known to support importation, culture, sale, or any
other use of live snakeheads.

ORGANISM RISK POTENTIAL

Organism within Colonization

Probability of
pathway Entry potential potential Spread potential
establishment HIGH → HIGH → HIGH → HIGH → HIGH

Consequence of Economic Environmental Perceived

establishment LOW → HIGH → HIGH → HIGH

Probability of Consequences of
Organism risk
etablishment establishment
potential HIGH → HIGH → HIGH

LOW = acceptable risk = organisms of little concern (does not justify mitigation)
MEDIUM = unacceptable risk = organisms of moderate concern (mitigation justified)
HIGH = unacceptable risk = organisms of major concern (mitigation justified)

40
SPECIES ACCOUNTS

41
42
 Channa amphibeus (McClelland, 1845)
Chel Snakehead

After Shaw and Shebbeare, 1938

Original description: Ophiocephalus amphibeus McClelland, 1845:274-282. Description of four species of fishes
at the foot of the Boutan Mountains. Journal of Natural History, Calcutta, 5(18):274-282. Type locality: vicinity of
tributaries of the Teesta (=Tista) River below the Boutan (Bhutan) mountains. Neotype: ZSI F 11435.
Synonyms: Ophiocephalus amphibius [sic], Day, 1877:365.
Ophiocephalus amphibius [sic], Shaw and Shebbeare, 1938:119-121 (vicinity of Chel River, northern
Bengal).
Channa amphibia [sic], Jayaram, 1981.
Channa amphibius [sic], Talwar and Jhingran, 1992.
Channa amphibious [sic], Eschmeyer, 1998:92.
Common names: In northern Bengal: bora cheng (Mechi), borna (Rabha) (Shaw and Shebbeare, 1938).
We suggest Chel snakehead.

Native range: Endemic to Chel River basin, Reproductive habits: No specific information,
Brahmaputra River drainage, northeastern India and but probably builds a cylindrical nest in vegetation and
Bhutan (Musikasinthorn, 2000). produces pelagic eggs.
Introduced range: No introductions known.
Size: To about 25 cm. Feeding habits: No specific information, but
likely a carnivorous predator as an adult.
Habitat preference: No specific information,
but distribution indicates preference for rivers, streams, Characters: Gular part of head without patch of
ponds, perhaps swamps in the Chel River basin, Brah- scales. Dorsal fin rays 50; anal fin rays 35; pectoral fin
maputra drainage, of northeastern India and Bhutan rays 15, pelvics 6; lateral line scales 81; cheek scales 9;
(Musikasinthorn, 2000). Shaw and Shebbeare (1938) scale rows above lateral line 5, below lateral line 13;
reported that during rainy periods, young are found “in predorsal scales 17; two large scales on underside of
flooded paddy-fields enclosed by forest; large fish can both sides of lower jaw. Head length 27.6 percent stan-
be found in pools of dried streams in forests.” dard length; anal fin length less than 50 percent of
pectoral fin length. Mouth large, maxilla extending far
Temperature range: No specific information. beyond posterior margin of eye. Many small conical
Habitat preference and range indicate a warm temper- teeth in premaxilla; three large conical teeth in
ate to subtropical species. prevomer; about four medium-sized canines on each

43
side of palatine; row of about five canine teeth on each Commercial importance in native range:
dentary, smaller than those on palatine; many small Unknown.
conical teeth on outer part of dentary. Characters based Environmental concerns: Unknown, but the
on neotype (Musikasinthorn, 2000). species is probably a predator on other fishes and
invertebrates.
Commercial importance in the United States:
Unlisted on aquarist-oriented websites. Negligible like-
lihood of being imported for sale in aquarium fish trade
or live-food fish markets.

80°E 100°E
Nepal
Bhutan China

India
EXPLANATION Myanmar Laos
DISTRIBUTION OF 20°N
Channa amphibeus Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate
Malaysia

Equator Indonesia
0°

Channa amphibeus

44
 Channa argus (Cantor, 1842)
Northern Snakehead

After Berg, 1933

Original description: Ophicephalus argus Cantor, 1842:484. General features of Chusan, with remarks on the flora
and fauna of that island. Ann. Mag. Nat. Hist. (n.s.), 9(60):481-493. Type locality: Zoushan Dao (Chusan Island),
China. Syntypes: BMNH 1843.7.21.3; BMNH 1860.3.10.
Synonyms: Ophicephalus pekinensis Basilewsky, 1855:225, tab. 9, fig. 3.
Ophicephalus argus warpachowskii Berg, 1909:200.
Ophiocephalus argus kimurai Shih, 1936:81, fig. 5.
Common names: northern snakehead; Amur snakehead; eastern snakehead; ocellated snakehead; zmeegolov
(Russian, Ussuri Basin and Lake Khanka region; Berg, 1965); her-yu (China; Evermann and Shaw, 1927); ga mul
chi (Korea); kamuruchi, raigyo (Japan; Bailey and Haller, 1977; Uyeno and Arai, 1984; Hosoya, 2002).

Native range: Middle and lower Heilong Japan (Nakamura, 1963; Uyeno and Akai, 1984),
(Amur) River basin; Songhua (Sungari) River, considered as a “careless importation” (Okada, 1960).
Manchuria; Tunguska River at Khabarovsk, Russia; Chiba and others (1989) recorded the dates of these
Ussuri River basin; Lake Khanka (Herzenstein and introductions as 1923-1924, noting that the species was
Warpachowski, 1887; Berg, 1965; Popova, 2002); reproducing in experimental or natural ponds and
Korea, except northeastern region (Okada, 1960; Berg, adding “Predation on native species” as a remark. In
1965); rivers of China southward and southwestward to Japan, present and established in Hokkaido, Honshu,
upper tributaries of the Chang Jiang (Yangtze) River Kyushu, and Shikoku (Hiroshi Ueda, personal
basin in northeastern Yunnan Province (Evermann and commun., 2001).
Shaw, 1927; Kimura, 1934; Nichols, 1943; Mori, 1952; Holcík (1991) reported this species as introduced
Berg, 1965; Xinluo and Yinrui, 1990; Ruihua, 1994). into “Czecho-Slovakia” and Russia, beginning in 1949.
Reported from Guangdong Province, China (Pearl Holcík did not provide specific localities of introduction
River Fisheries Research Institute, 1991), likely an or information on status of the releases. Bogutskaya and
introduction there. Widely distributed in Chinese Naseka (2002) listed Channa argus as having been
reservoirs (Sifa and Senlin, 1995). transplanted within Russia. Nina Bogutskaya (personal
Introduced range: Occurrence of this species in commun., 2002) reported failed early introductions of
the upper reaches of the Bei (Beijiang) River, Guang- the northern snakehead into the Volga Delta and ponds
dong Province, China, is apparently the result of an in Ekaterinburg (formerly Sverdlovsk) Province in the
introduction (Pearl River Fisheries Research Institute, southern Urals. An experimental introduction was
1991). Introduced from Korea in the early 1900s and made in ponds of Moscow Province during 1949-1950
established in many waters of central and southern that established. In 1953, it was recommended that this

45
 for the snakeheads. In one pond that contained less
vegetation, rudd (Scardinius erythrophthalmus) were
used to supplement the food base. Snakeheads were
periodically sampled from 1961-1964 to determine
growth rates. Winters between 1961-1963 were unusu-
ally severe. All of the ponds became covered with ice
during winter. The winter of 1963 was particularly cold
over a long period of time (air temperatures often
reaching -30 ºC and ice cover present for over 3 months).
Due to decaying vegetation in two of the ponds produc-
ing hydrogen sulfide, a lack of oxygen, and thick ice on
the surface, snakeheads and all other fishes perished in
these ponds. Snakeheads did survive in the third pond,
Juvenile northern snakehead, about 4 cm long, captured which was also being used experimentally to test if
from pond in Crofton, Anne Arundel County, Maryland, July 9, brown trout (Salmo trutta) could be used to control
2002. Photo by Algerina Perna; reproduced with permission
of the Baltimore Sun.
overpopulation of perch (Perca fluviatilis). During the
winter 1963, the supply of roach and white bream was
eliminated in the third pond by snakeheads and brown
trout; the food base was supplemented with perch fry
species should be stocked widely but that failed to
and snakeheads survived, but failed to grow. It was
happen. There was one report in a Russian aquarium
concluded that the northern snakehead could be accli-
journal in 1963 noting occurrence of this snakehead in
matized to Czechoslavakia as long as sufficient food
small lakes in the Podolsk Region, Moscow Province,
existed. Interestingly, when the pond was cleared of
but the species is presently absent from the Moscow
vegetation for fish removal at the end of the experimen-
area. Tandon (1976) reported that acclimatization
tal period, the northern snakeheads buried themselves
experiments were conducted in the former Soviet
in mud on the pond bottom making their capture difficult.
Union after 1950, and that fry were collected from
ponds near Moscow and the Ukraine in 1955 and sent Borisova (1972), Amanov (1974), Dukravets and
Machulin (1978), Usmanova (1982), Guseva (1990),
to Czechoslovakia for acclimatization purposes. He
and Dukravets (1992) documented introduction of
concluded that the source of the original stock was the
Channa argus into the Aral Sea basin (about 45º N) in
Amur basin, not the Yangtze of China. In further
the early 1960s and its presence in the Amu Dar’ya,
support of the Amur basin having been the source of
Syr Dar’ya, and Kashka-Dar’ya rivers of Kazakhstan,
the stock, Zhadin and Gerd (1963) noted that “rearing
Uzbekistan, and Turkmenistan. The initial importation
of many Amur fishes possessed of frost resistance and
in 1961 of the northern snakehead was accidental when
great plasticity was recently initiated in water bodies in
it was included with target phytophagous cyprinids
the European U.S.S.R.” and included the northern
(grass carp, Ctenopharyngodon idella, and silver carp,
snakehead among species imported.
Hypophthalmichthys molitrix) that were destined for
Frank (1970) was more specific on introduction use in ponds adjacent to the Syr Dar’ya River. Intro-
of northern snakehead to Czechoslovakia. He stated duction into the ponds occurred at “the ’Akkurgan’
that the first shipment (three individuals) was made in fish combine” (Amanov, 1974). Snakeheads escaped
December 1956 by V. Nikol'skiy of the Department of from ponds in 1964 and soon became established in
Ichthyology, Moscow University. These fish were the Syr Dar’ya (Amanov, 1974). Dukravets and
introduced into a pond near Nižbor during Spring 1957 Machulin (1978) reported that this snakehead migrated
that later flooded into a nearby creek, resulting in the downstream into the Aral Sea about 1965. Similar
loss of the snakeheads. Nikol'skiy authorized a second escapes from ponds along the Kashka-Dar’ya in south-
shipment of 51 fish less than a year in age in December ern Uzbekistan in 1964 also led to establishment. It was
1960. Several of these were released into three "natural also released with carp (Cyprinus carpio) in 1964 into
ponds" near the Elbe River, with the remainder intro- Chimkurgan Reservoir in the Kashka-Dar’ya basin
duced to the same ponds during summer 1961. These from a fish farm (Amanov, 1974; Usmanova, 1982;
ponds contained small cyprinids, roach (Rutilus rutilus) Baltz, 1991). Baltz (1991), however, cited 1963 as the
and white bream (Blicca bjoerkna), that served as prey year of introduction into the Aral Sea basin. At present,

46
the Aral Sea has almost disappeared and there are now adult Channa argus to the biologist who reported the
hundreds of kilometers between the terminal reaches of capture in 1997 and he responded that this appeared to
these rivers and the former coastline of the Sea (Nina be the same species (John Sunada, personal commun.,
Bogutskaya, personal commun., 2002). Moreover, the 2002). It is unlikely that another species of snakehead
salinity of the Aral Sea (10-12 ppm in about 1990; could survive in this high-altitude reservoir.
Baltz, 1991) is well above the tolerance limits of Two specimens were captured from the St. Johns
snakeheads. River below Lake Harney, Seminole and Volusia Coun-
All introductions into western Asia and eastern ties, Florida. The first was collected February 11, 2000,
Europe were thought by some to have originated from and the second on March 4, 2000. There are uncon-
China, perhaps from Amur basin stock of Channa firmed reports of three additional specimens having been
argus. Sal’nikov (1998), in a comprehensive study of caught nearby (Franklin N. Snelson, Jr., personal
the inland waters of Turkmenistan, stated that the Aral commun., 2001). An attempt was made by USGS
Sea introduction source was the Yangtze basin of personnel to collect additional specimens at this site
China. He reported the initial introductions of northern by electrofishing in 2001, but none were collected. Addi-
snakehead into Uzbekistan, with later migration tional attempts to collect this species will be undertaken.
through the Aral Sea into the Amu Dar’ya and other On May 14, 2002 a Maryland angler caught a
waters of Turkmenistan. Dukravets and Machulin 43-45 cm long snakehead in a 1.8-ha retention pond
(1978) and Usmanova (1982) noted that the characters behind a shopping mall in Crofton, Anne Arundel
of the introduced stock mostly agreed with those of County, Maryland. The angler took photographs of the
northern snakehead from the Amur basin. Dukravets unusual looking fish before releasing it back into the
(1992) reported additional introductions of this fish pond. A month later, the angler took one of the photos
beginning in 1976 from the lower part of the Sarysu to the Maryland Department of Natural Resources
River, north of the middle Syr Dar’ya, followed by (MDDNR) headquarters in Annapolis for identifica-
further releases into reservoirs of the Talas and Chu tion, and one of the biologists thought it might be a
Rivers, north and northeast of Tashkent, the capitol of snakehead. That same day, the photograph was emailed
Kazakhstan, during the middle 1980s. The species was to the USGS in Gainesville, Florida, where we readily
thriving in irrigation reservoirs and isolated ponds on identified the fish as a northern snakehead and immedi-
the Talas River, and more than 10 metric tons were ately notified MDDNR. In fact, we had just completed
commercially harvested from the reservoirs in 1989 and submitted a document on the risk assessment and
(Dukravets, 1992). The latitude of this area of Kazakh- evaluation potential for establishment (among other
stan parallels those of the Vermont/Quebec border, biological factors) to the U.S. Fish and Wildlife Service
Minneapolis, Minnesota, and Salem, Oregon, in North on the entire family of snakeheads about a week prior
America, indicating a possible northern range of distri- to when the Crofton pond fish was initially caught.
bution should this snakehead become established in
A second snakehead (about 66 cm long) was
U.S. waters.
caught by a second angler in the same retention pond
As noted earlier, presence of this species in on June 30, 2002. On the evenings of July 7-8, he
Guangdong Province, southern China (Pearl River dipnetted eight juvenile snakeheads from the pond,
Fisheries Research Institute, 1991), is likely the result which we subsequently confirmed as northern snake-
of an introduction. heads. This was the first proof that the species was
This species is recorded from open waters of established as a reproducing population. Later that
the United States. A large snakehead was captured by week (July 11), MDDNR biologists captured an addi-
electroshocking by California Fish and Game Depart- tional 99 juveniles ranging 5-6 cm in length, using
ment personnel in Silverwood Lake, located in the backpack electrofishing units close to shore (Steve
San Bernardino Mountains, north of San Bernardino, Early, personal commun., 2002). During the following
California, on October 22, 1997. The lake receives week, more juveniles about the same size were
water from the California Aqueduct and serves as a captured by electrofishing. The pond and two adjacent
reservoir for Los Angeles. A photograph of the speci- smaller ponds were treated with herbicides by
men was taken and the specimen was frozen but later MDDNR personnel on August 18, 2002. Two dead
discarded without being preserved. The photograph has juveniles, about 10 cm in length, were collected from
not yet been located. We sent a digital photograph of an the largest pond on August 20, 2002.

47
 All three ponds were treated with rotenone on (Uchida, 1933). It appears to occupy waters, usually
September 4. No snakeheads were found in the two with vegetation, close to shore, and also feeds in schools
smaller ponds. Dead fish ceased rising to the surface of (Nina Bogutskaya, personal commun., 2002).
the largest pond on September 7. The lengths of recov- Temperature range: 0 to >30 ºC (Okada, 1960).
ered northern snakeheads included two ranging from
69-70 cm, two 59 cm individuals, two ranging from Reproductive habits: Soin (1960) provided a
41-43 cm, more than 800 from 10-10.5 cm, and 566 detailed, illustrated report on spawning and development
young juveniles about 3.5 cm. The total number of of the northern snakehead from within its native range.
snakeheads recovered was greater than the number of His study was based on observations made at a fish farm
native fishes killed (Bob Lunsford, personal commun., adjacent to the Songhua Jiang (Sungari River), north-
2002), although the biomass of natives was greater eastern China, in July 1958. He found fertilized eggs,
(Steve Early, personal commun., 2002). about 1 day old, within an open nest (=cleared of
vegetation) about 1 m in diameter and 60-80 cm deep.
A specimen of northern snakehead was captured
The eggs were guarded by two adults. Eggs were
by Massachusetts Department of Fish and Wildlife
described as buoyant due to presence of a large lipid
personnel in Newton Pond, Shrewsbury, Worcester
droplet that was more than three-quarters the diameter of
County, Massachusetts, on October 4, 2001 (Karsten
the egg. Each egg had a diameter of 1.8-1.85 mm and the
Hartel and Todd Richards, personal commun., 2001;
yolk was bright yellow. At 1 day of age, the length of an
Hartel and others, 2002). The likely source for both the
embryo removed from a fertilized egg was 3.2 mm.
Florida specimens and the one from Massachusetts is
Hatching occurred 12 hours later at temperatures of
live-food fish markets.
23-25 °C, about 2 days after spawning, and each larva
In early August 2002, Gene Polk from Charlotte,
was about 4.5 mm long. Respiration was through the
North Carolina, reported that he and an angling partner caudal vein, the subintestinal vein that covers much of
caught two northern snakeheads from Lake Wylie, a the yolk sac, and the enlarged ducts of Cuvier. Pectoral
reservoir on the Catawba River, near the U.S. Route 74
fins appeared about 1 day after hatching and larvae were
bridge. These fish were identified as Channa argus about 5.7 mm long. Yolk was noticeably resorbed, ducts
(Wayne Starnes, personal commun., 2002). of Cuvier were reduced in size, and respiration was
The USGS received an unconfirmed report of an primarily through the subintestinal and hepatic veins that
angler who caught a snakehead near Rockport, south- covered about two-thirds of the yolk sac surface. An oral
eastern Texas, that he believed was a northern snake- aperture was present and an operculum covered the
head based on illustrations in Howells and others developing gills. A length of 7.1 mm was reached at
(2002). He reportedly released the specimen without 2 days following hatching and most of the yolk was
photographing it and has not followed up on his prom- resorbed. Nevertheless, the large lipid droplet was still
ise to provide a collection location or size of the fish present and caused what remained of the yolk sac to
(Robert Howells, personal commun. 2002). protrude laterally, producing a pair of cystiform struc-
Size: To 85 cm (Okada, 1960), although Nina tures. Melanophores had become well developed in the
Bogutskaya (personal commun., 2002) reported seeing skin, giving the larvae a very dark color. By 3 days old,
captured specimens in Russia approaching 1.5 m total very little yolk remained, external blood vessels became
length. In the Amur River where the species is native markedly reduced, respiration by gills had begun, and
and the Syr Dar’ya of Kazahkstan where it is intro- the cystiform outpocketings remained visible on top of
duced, males tend to be larger than females, with a what remained of the yolk sac. Pectoral fins, used in
higher dorsal fin, wider interorbital distance, and locomotion, had enlarged, and the larvae, at 7.3 mm in
longer snout, postorbital distance, and upper jaw. length, had begun to feed. Larvae remained near the nest
Moreover, morphometrics vary with age (Dukravets surface, grouped together, guarded by adults. About
and Machulin, 1978). 2 weeks after hatching, larvae were approaching 11 mm
Habitat preference: Stagnant shallow ponds or in length, the yolk sac and cystiform processes had
swamps with mud substrate and aquatic vegetation; slow disappeared, fin rays were visible in the pectorals, color-
muddy streams (Okada, 1960). Also occurs in canals, ation was black, and the epibranchial cavities that will
reservoirs, lakes, and rivers (Dukravets and Machulin, later be used for aerial respiration had begun to develop.
1978; Dukravets, 1992; Nina Bogutskaya, personal A length of 2 cm was reached by the fourth week after
commun., 2002). This species is an obligate airbreather hatching, pelvic fins were developing, epibranchial

48
breathing cavities had become functional, and body then begin feeding on small crustaceans and fish larvae.
color had changed from black to brown. Larvae of this Parental care continues through the post-larval stage.
size had lost their aggregative behavior and moved to Feeding habits: Post-larvae feed on plankton,
slightly deeper water. Scales did not develop until the juveniles on small crustaceans and fish larvae, adults on
early juveniles had reached a length of 4-4.5 cm, when fishes, frogs, crustaceans, and aquatic insects. This
the mottled pattern of dark blotches characteristic of the species is reported to be a voracious feeder (Okada,
species had appeared. 1960). Dukravets and Machulin (1978) noted that in the
Many of the reports on this species vary consider- Syr Dar’ya basin, northern snakeheads fed on 17 species
ably from what was observed following its introduction of fishes, including young and fish up to 33 percent of
into Maryland. The northern snakehead has been said to the predator’s body length. Larger prey included loach
reach sexual maturity in about 3 years at a length of 30- (Cobitis), bream (Abramis), carp (Cyprinus carpio), and
35 cm in the Amur and Syr Dar’ya, although some can perch (Perca fluviatilis). Other food items included cray-
spawn during the second year (Dukravets and Machulin, fish, dragonfly larvae, beetles, and frogs, as well as plant
1978). Okada (1960) reported spawning at 2 years at a material that was probably ingested incidentally with
length of 30 cm in Japan, and Nikol’skiy (1956) indi- prey. Guseva (1990) summarized dietary changes in
cated the same for northern snakeheads in the Amur the Amu Dar’ya basin related to age based on Guseva
basin, differing from the age report given by Dukravets and Zholdasova (1986). Northern snakeheads feed on
and Machulin (1978). This species builds a mostly circu- “crustacean zooplankton, Cladocera,” copepods, and
lar nest of pieces of aquatic plants, about 1 m in diame- small chironimid larvae for the first month of life. Once
ter, in shallow aquatic vegetation. The water surface a length of 4 cm is reached, they begin to feed on fishes
above the nest is cleaned by the parents, and spawning and when juveniles become 13-15 cm, fishes dominate
occurs at dawn or in early morning. The female rises 64-70 percent of the diet. Juveniles up to 30 cm feed
near the surface and releases eggs, which are then fertil- almost exclusively on fishes (90 percent of diet), mostly
ized by the male. Eggs are pelagic, nonadhesive, spheri- small goldfish (Carassius) and roach (Rutilis). More-
cal, yellow, and about 2 mm in diameter. Eggs hatch in over, in the Amu Dar’ya basin, northern snakehead feed
28 hours at 31 ºC, 45 hours at 25 ºC, and 120 hours at only from late March to the end of October. Feeding
18 ºC. Number of eggs released ranges between 1,300- begins when water temperature reaches 10 ºC with
15,000, with an average of 7,300. Wee (1982) cited 45 percent of its annual food consumption completed
Frank (1970) who reported fecundity for this species of by May. Roach dominated in the diet, but commercial
about 50,000 oöcytes. The report by Frank (1970) was species such as carp, zander (Sander), bream, grass carp
based on examination of individuals in Czechoslovakia, (Ctenopharyngodon idella), and various catfishes were
imported from Moscow, Russia, and apparently of Amur also consumed. Another 46 percent of annual food
basin stock. Additionally, fecundity in the Syr Dar’ya consumption occurred during June and July at tempera-
basin is higher than reported for the species in the Amur tures of 20-27 ºC with carp, zander, grass carp, and
basin by Nikol’skiy (1956), about 22,000-51,000 in the bream dominating (50 percent of diet; Guseva, 1990).
Amur basin and a low of 28,600 to a high of 115,000 in Feeding declined by autumn as water temperatures
reservoirs and lakes of the Syr Dar’ya basin (Dukravets decreased to 12-18 ºC and ceased when temperatures
and Machulin, 1978). Berg (1965) reported that the dropped below 10 ºC.
northern snakehead spawned five times per year in the This species is reported to feed in schools, with
Amur basin. Dukravets and Machulin (1978), however, most activity at dusk into early night and again before
noted that it spawns two to three times in the Syr Dar’ya dawn, typically in vegetation close to shore. It is also
basin, and in one group of lakes only once per year, typi- reported that actively feeding adults make grunting
cally from May to June at a water temperature of 18- noises "like pigs" (Nina Bogutskaya, personal commun.,
20 ºC. Newly hatched larvae are about 4 mm in length 2002). Similarly, Soin (1960) noted clicking sounds
and black. Larvae remain in the nest, guarded by one or produced by the northern snakehead in ponds adjacent to
both parents, until the yolk is resorbed and body length the Songhua Jiang (Sungari River), northeastern China,
is about 8 mm. Larvae leave the nest as a group after as the fish rose to the surface to breathe air.
yolk resorption and begin feeding on plankton. A post- Characters: Gular part of head without patch of
larval stage follows until a length of about 18 mm is scales. Head somewhat depressed anteriorly; interor-
reached at which time aerial respiration begins. Young bital area flat; eye above middle of upper jaw. Mouth

49
large, reaching far beyond eye. Villiform teeth present live snakeheads was legal. This was verified in July
in bands with some large canine-like teeth on lower jaw 2002 when it was also learned that two additional
and palatines. Lateral line scales 60-67; 8 scale rows Arkansas aquaculturists were then culturing northern
above lateral line to dorsal fin origin; 12-13 scale rows snakehead. The report included information that the
below lateral line to anal fin origin. Dorsal fin elongated, original snakehead aquaculturist had been approached
with 49-50 rays; anal fin with 31-32 rays. Origin of by a live-food fish importer in New York and asked if
pelvic fin beneath 4th dorsal fin ray. Pectorals extending he could culture snakeheads for sale in U.S. markets.
beyond base of pelvic fins. Apparently, the second two fish farmers decided that
The body has a very distinctive color pattern. they too might profit from snakehead culture. Thus, as
Background color is golden tan to pale brown with a of July 2002, there were three domestic sources of
series of dark blotches on the sides and saddle-like cultured northern snakehead, although the importation,
blotches across the back interrupted by the dorsal fin. culture, and possession of snakeheads in Arkansas
The species is capable of darkening its background were prohibited on July 29, 2002.
colors to the point of almost obscuring the blotches Commercial importance in native or intro-
(personal observation). The upper blotches on the sides duced range: This species is cultured in ponds, rice
are typically separate anteriorly, but more posterior paddies, and reservoirs in China (Atkinson, 1977; Sifa
blotches may coalesce with ventral blotches. There is a and Senlin, 1995) and considered a valuable commer-
dark stripe from just behind the eye to the upper edge cial fish in Korea (Berg, 1965). FAO (1994) listed a
of the operculum with another dark stripe below from production of 500 metric tons in Korea in 1992. It is the
behind the orbit extending to the lower quadrant of the most important snakehead cultured in China, with most
operculum. Coloration of juveniles is virtually the culture activities centered in the Chang Jiang (Yangtze)
same as in adults, a characteristic atypical for many basin (Fang Fang, personal commun., 2002). It never
snakehead species. became a popular food fish following introduction into
Coloration of the northern snakehead is quite Japanese waters (Okada, 1960). In tributaries of the
similar to that of the blotched snakehead, Channa Aral Sea, Kazakhstan, large populations of this species
maculata. A key character for separating these two is are fished commercially with an annual catch reported
the bar-like markings on the caudal peduncle. In of 1-5 metric tons (Baltz, 1991). Dukravets (1992),
C. maculata, the most posterior dark bar (usually however, reported an annual catch of 10 metric tons
complete) is preceded and followed by pale bar-like from reservoirs on the Talas River, Kazakhstan.
areas, whereas in C. argus, such pale markings are Environmental concerns: This species is
absent and the final dark marking is irregular, often described as a voracious predator on other fishes, and
blotch-like. also feeds on freshwater crustaceans. Moreover, its
Commercial importance in the United States: native range (24-53º N) and temperature tolerance
Generally not listed on aquarist-oriented websites. (0-30 ºC) indicates a species that, if introduced, could
Nevertheless, this species has been imported for sale in establish feral populations throughout most of the
live-food fish markets and has been the most widely contiguous United States and possibly some waters in
available snakehead in the U.S. It likely has comprised adjoining Canadian provinces. Because it was found to
the largest volume and greatest weight of live snake- be established in Maryland in 2002, perhaps also in
heads imported into the U.S. until 2001. The authors Florida, and was the most widely available snakehead
obtained specimens that were procured alive from fish sold as a live-food fish in the U.S., the likelihood of its
markets in New York, Houston, Pembroke Pines, and becoming more widely established is real.
Orlando, as well as photographs of live northern snake- Fourteen states banned possession of live snake-
head in markets in New York and St. Louis that were heads prior to August 2002. Among those states is Texas
taken by Leo Nico of the USGS. Photographs from the where possession of live snakeheads has been illegal for
St. Louis market were taken in mid-July 2002, after almost four decades (Howells and others, 2002). During
northern snakeheads were found established in Mary- Summer 2001, game wardens from the Texas Parks and
land; the storeowner had the fish marked as “mudfish.” Wildlife Department discovered large numbers of
Also, see below under Environmental concerns. Channa argus being sold in live-food fish markets in
A fish farmer in Arkansas was discovered cultur- Houston. Raids resulted in seizure of the fish and ship-
ing northern snakeheads in 2001 when possession of ping documentation that traced the fish to a distributor

50
who had obtained the fish from a source in New York could be purchased, raising the probability of introduc-
(Robert H. Howells, personal commun., 2001). The tions that could become established, as occurred in
distributor was an aquaculture facility that had imported Crofton, Maryland, with Channa argus.
northern snakeheads for sale to retailers in Texas, and
Snakeheads are also prohibited in Washington.
agents confiscated the fish (Howells and others, 2002). At 11:35 a.m. on April 30, 2001, a driver for a Canadian
Florida also prohibits possession of live snake- fish wholesaler declared a shipment of live ling cod in
heads. Following the discovery of the bullseye snake- Blaine, Washington. The shipment was bound for a sea-
head, Channa marulius, that was established in food distributor in Seattle. Wildlife Inspector Michael
Tamarac, Broward County, Florida, in spring 2001, Williams, U.S. Fish and Wildlife Service, inspected the
agents of the FFWCC raided two live-food fish markets, shipment to find three open boxes containing fish he
one in Miami, Miami-Dade County, and another in thought were “unusual looking.” When he asked the
Pembroke Pines, Broward County. They confiscated driver what they were, his reply was that they were
several live C. argus (Florida Fish and Wildlife Conser- snakeheads that had been pond raised in China and
vation Commission, 2001). Channa argus was also shipped without water to Canada, adding this was the
confiscated from a live-food fish market in Orlando, first time his employer had made such a shipment. Upon
Orange County, by FFWCC agents in July 2002. examining one box, Williams noticed the fish moved
Despite Florida's prohibition, one of the authors and, on further investigation, found that most were alive
(JDW) purchased a live northern snakehead from a fish and some “capable of vigorous movement.” Williams in-
market in Orlando, Orange County, in March 2002. formed the driver that possession of live snakeheads was
The tank in which the fish was found was labeled in violation of Washington State regulations. The driver
“Chinese catfish,” although the proprietor explained to was asked to kill the fish and began striking them with a
the buyer that the fish was not a catfish but “a very board. Williams notified the Washington Department of
special fish.” The Center for Aquatic Resources Studies Fish and Wildlife and, after returning to the truck, found
of the USGS Florida Integrated Science Center has a that the fish were still alive despite the driver’s attempt to
permit to possess certain restricted fishes including kill them. He seized the 80 fish at noon and placed them
snakeheads. This purchase was made to prove that ille- in a freezer. Washington Department of Fish and Wild-
gal fish can be purchased in Florida. FFWCC agents life authorities arrived about 12:30 and removed the fish
confiscated four live northern snakeheads from this from the freezer. Most were still alive. State authorities
market on July 16, 2002. took possession of the fish to proceed with penalties
The FFWCC licenses pet dealers, importers of against the companies involved (Mike Williams, per-
restricted and tropical fishes, and retailers and whole- sonal commun., 2003). The shipping invoice listed the
salers of non-native aquatic species. This last item fish as “Fresh Snakehead Fish–Product of China.” The
includes markets that sell live-food fishes (Kyle Hill, fish were subsequently identified as northern snakeheads
personal commun., 2002). The Division of Aquaculture (Ted Pietsch, personal commun., 2001).
of the Florida Department of Agriculture and Con- Discovery of an established population of north-
sumer Affairs certifies and inspects fish culture facili- ern snakeheads in Maryland, catches by anglers of two
ties, and deals with permits for restricted species to fish verified and three unconfirmed specimens from the
farmers (Paul Zajicek, personal commun., 2002) but St. Johns River in east-central Florida, and others from
does not license retailers or wholesalers that sell live- a California reservoir, a pond in central Massachusetts,
food fishes. Restricted fishes can only be sold to people and two from a reservoir in North Carolina, coupled
who possess a valid Aquaculture Certificate from with proof of their availability in live-food fish markets
Department of Agriculture and Consumer Affairs. at or near a size when they are reproductively mature,
Moreover, a retailer of live-food fishes who does not were reasons for substantial environmental concern.
apply for a license becomes an illegal “unknown” to
regulatory agencies. The markets in Miami, Pembroke Despite the fact that snakeheads are now prohib-
Pines, and Orlando that were selling northern snake- ited from importation, several live specimens of northern
heads did not have licenses to possess or sell live fresh- snakehead were confiscated in California in July 2003.
water fishes (Kyle Hill and Barry Cook, personal Comments: The diploid chromosome number
commun., 2002). Clearly, there have been sources other of Channa argus is 48 (Lee and Lee, 1986; Wu and
than aquarium fish dealers from which live snakeheads others, 1986).

51
 60°E 80°E 100°E 120°E 140°E 160°E

60°N
Russia

Kazakhstan
Aral
Sea
Mongolia
Uz
be
kis Kyrgyzstan
tan North
40°N Turkm
enista Tajikistan Korea
n

Afghan- China
Iran istan Japan
South
Pakistan Korea

Saudi PACIFIC
India
Arabia OCEAN
20°N

INDIAN
OCEAN Philippines
Thailand

EXPLANATION 0 1,000 MILES

DISTRIBUTION OF Channa argus
Native range 0 1,000 KILOMETERS
Introduced range Scale is approximate

Distribution of Channa argus in the Eastern Hemisphere

52
 Channa asiatica (Linnaeus, 1758)
Chinese Snakehead

UF 127103, 135 mm standard length. Specimen purchased in an Asian market in Kansas,

October 2002. Photo by Buck Albert, USGS, Gainesville, Florida.

Original description: Gymnotus asiaticus Linnaeus, 1758:246. Systema Naturae, ed. 10. Holmiae, 1:i-ii + 1-824.
Type locality: Asia. Holotype: ZIU 171.
Synonyms: (?)Ophicephalus miliaris Cuvier, 1831:439.
Channa ocellata Peters, 1864:392.
Channa fasciata Steindachner, 1866:480.
Channa sinensis Sauvage, 1880:58.
Channa orientalis Karoli, 1882:147.
Channa formosana Wu, 1929:73; Myers and Shapovalov, 1932:36.
Cuvier (1831) described Ophicephalus miliaris based on a description by Jean-Jacques Dussumier of a
snakehead from Canton (Guangzhou), China. Dussumier’s descriptions were made from live or fresh specimens and
were considered quite accurate by Cuvier (Bauchot and others, 1990). Figure 10 in Bauchot and others (1990), an
illustration of a snakehead lacking pelvic fins and based on Dussumier’s description, appears to be Channa asiatica.
Therefore, we have treated O. miliaris as a possible synonym of C. asiatica. Nevertheless, Peter Ng (personal
commun., 2003) noted that a closely related species, C. nox, that has very similar markings to C. asiatica, is
available for sale in live-food fish markets in Ghangzhou, raising the possibility that O. miliaris might be a synonym
of that species. Resolution of this situation is complicated because there are apparently no types of O. miliaris.
Common names: Chinese snakehead; chi hsing yü (Yangtze River basin, China); hua-t’sai-yü (Tungting Lake on
Chang Jiang [Yangtze] River); kôtal (Japan; Uyeno and Akai, 1984; Hosoya, 2002).

Native range: China, middle and lower Chang length for this species. They also commented that
Jiang (Yangtze) basin, and Xun River basin in Guangxi growth is rapid during the first 2 years of life and slows
and Guangdong provinces (Kimura, 1934; Pearl River thereafter. They recommended regulating a minimal
Fisheries Research Institute, 1991). Also reported from harvestable age of this species at 2.6 years for resource
Hainan Island, China (Kimura, 1934) where it is likely conservation purposes.
native rather than introduced. Habitat preference: No information available.
Introduced range: Taiwan (Musikasinthorn, Probably a riverine species.
2000); Japan, Ishigaki Shima Island in the Ryukyu Temperature range: No information available.
Islands (Uyeno and Akai, 1984; Hosoya, 2002). Klee Nevertheless, distribution within native range (32-22º N)
(1963) reported that this species was “occasionally indicates a warm temperate to subtropical species.
found in Florida waters,” but we have found no Reproductive habits: No information found
evidence that it is established. concerning reproduction in natural habitats. Hosoya
Size: Up to 34 cm (Daiqin and others, 1999). By (2002) stated this species does not build a nest but pro-
studying annual ring development on scales, they noted vides parental care in Japan where the species was
a linear relationship between scale length and body introduced. Breder and Rosen (1966) summarized this

53
species, in aquaria, as producing floating eggs, the eggs Commercial importance in the United States:
about the “size as that of the head of a pin.” Reproduc- Ross B. Socolof (personal commun., 2003) reported
tively active females become paler with a pink cast, Channa asiatica as the first snakehead to have been
with dark brown markings becoming darker and silver imported to the contiguous United States for the aquar-
markings becoming brighter. Reproductively active ium fish trade. Subsequent accounts of this species in
males also become darker. The prespawning female the aquarium fish literature (Innes, 1920, 1955;
rises to the surface of an aquarium, gulps air, and rolls Armstrong, 1923; Stoye, 1935; Axelrod and Schultz,
from side to side. The male then rises, circles the 1955) are indicative of its availability through much of
female, and they embrace, rising to the surface with the the early to mid-1900s. Typically not listed on aquarist-
male squeezing the female. They are reported to have oriented websites.
rolled once, then sank, releasing themselves, repeating Commercial importance in native range:
the sequence after a few minutes. In aquaria, spawning Nichols (1943) commented that it was never seen for
can occur every 6 to 10 days. Reproductive activities sale in China. Nevertheless, it is sold in the aquarium
appear to occur at night. Hatching in aquaria occurs in fish trade outside its native range in Singapore (Ng and
about 24 hours at 28 ºC. There are some indications Lim, 1990). Daiqin and others (1999) recommended
that the male may be a mouthbrooder. Both male and restricting the legal harvestable age to 2.6 years, a clear
female are reported to aggressively protect against any- indication that the species is being fished commercially
thing introduced into an aquarium when eggs or fry are in China. Ping Zhuang (personal commun., 2002) noted
present. that this species is now common in aquaculture in China.
Environmental concerns: Probably a thrust pred-
Feeding habits: No information found, but ator on other fishes and invertebrates like many other
likely a thrust predator. snakeheads. According to Ross B. Socolof (personal
Characters: Gular region of head without patch commun., 2003), this was the first snakehead imported
of scales. Pelvic fins absent. Dorsal fin with 44 rays; into the U.S. for the aquarium fish trade, the earliest
anal fin 26 rays; pectoral fin 14 rays. Lateral line scales imports occurring in the late 1800s or early 1900s.
57; predorsal scales 12; rows of scales between dorsal Comments: The diploid number of chromo-
origin and lateral line 5; scale rows between lateral line somes of Channa asiatica is 46 (Wu and others, 1986).
and midline of belly 16. Color
100°E 120°E
pattern distinct among snake-
North
heads with dark chevrons on 40°N Korea
sides and large ocellus centered
on caudal peduncle. The center South
EXPLANATION Korea
of the ocellus is not round but DISTRIBUTION OF
China
Channa asiatica
more quadrangular in shape
Native range East
(Bureau of Aquatic Products
Introduced range
Industry, 1988; Lee and Ng, China

1991), unlike that illustrated in 0 1,000 MILES

Sea
Pearl River Fisheries Research
0 1,000 KILOMETERS
Institute (1991). The most
Scale is approximate Vietnam
closely related snakehead is Myanmar Taiwan
Laos
20°N
Channa nox, which has 47-51 Hainan Island
dorsal fin rays, 31-33 anal fin South China
Thailand
rays, and is black on the upper Sea
half of the body. Like Bay of Cambodia
C. asiatica, C. nox also lacks Bengal Philippines
pelvic fins and has a large black
ocellus on the caudal peduncle. Thailand

Channa asiatica

54
 Channa aurantimaculata Musikasinthorn, 2000
Orangespotted Snakehead

Upper image: holotype, KUMF 3135, 190.8 mm standard length. Lower image: ventral view of head showing
enlarged scales on lower jaw, paratype, KUMF 3136, 163.7 mm standard length. Reprinted with permission
from Prachya Musikasinthorn, author, and Tomoki Sunobe, Secretary of the Ichthyological Society of Japan,
from: Musikasinthorn, Prachya. 2000. Channa aurantimaculata, a new channid fish from Assam (Brahmaputra
River basin), India, with designation of a neotype for C. amphibeus (McClelland, 1845). Ichthyol. Res.
47(1):27-32.

Original description: Channa aurantimaculata Musikasinthorn, 2000:27-32. Channa aurantimaculata, a new

channid fish from Assam (Brahmaputra River Basin, India), with designation of a neotype for C. amphibeus
(McClelland, 1845). Ichthyological Research, 47(1):27-37, figs. 1-5. Type locality: Dibrugarh town, Digrugarh,
Assam, India. Holotype: KUMF 3135. Paratypes: KUMF 3136; NSMT-P 55735; ZSI uncataloged, collected with
KUMF 3136.
Synonyms: No synonyms.
Common names: orangespotted snakehead; naga-cheng (Assam, India).

Native range: Endemic to middle Brahmaputra Reproductive habits: No specific information,

River basin, northern Assam, India (Musikasinthorn, but probably a nest builder with pelagic eggs like the
2000). majority of channid fishes.
Introduced range: No introductions known. Feeding habits: No information, but more than
likely a carnivorous predator as adults.
Size: To about 40 cm.
Characters: Patch of scales on gular part of head
Habitat preference: Forest streams, ponds, and absent. Head elongated, mouth large, interorbital region
swamps adjacent to the Brahmaputra River in subtropi- almost flat. Dorsal fin rays 45-47; anal rays 28-30;
cal rainforest conditions (Musikasinthorn, 2000). lateral line scales 51-54; cheek scales 8-12; predorsal
Temperature range: Unknown, except preferred scales 13-15; total vertebrae 50-52; two large scales
habitat and known range is subtropical. on each side of undersurface of lower jaw; pelvic fin

55
length less than 50 percent of pectoral fin length; Commercial importance in the United States:
cephalic sensory pores single without satellite openings. None known. Recently described species, not listed on
Small conical teeth in premaxilla with an additional aquarist-oriented websites and unknown for sale in
series of somewhat larger conical teeth anteromedially; live-food fish markets.
small teeth and 3 large conical teeth on prevomer; a Commercial importance in native range: No
row of conical teeth with 5 or 7 large canines on each specific information, but reported in markets in Assam,
side of palatine; dentary with medium-sized conical India (Musikasinthorn, 2000).
teeth on each side, also with 5 to 6 large canine-like Environmental concerns: Unknown, but
teeth (Musikasinthorn, 2000). probably a predator on other fishes and invertebrates.

80°E 100°E
Nepal
Bhutan China

India
EXPLANATION Myanmar Laos
DISTRIBUTION OF 20°N
Channa aurantimaculata Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate
Malaysia

Equator Indonesia
0°

Channa aurantimaculata

56
 Channa bankanensis (Bleeker, 1852)
Bangka Snakehead

Reprinted with permission from P.K.L. Ng from: Lee, P.G., and P.K.L. Ng. 1991. The snakehead fishes of the
Indo-Malayan region. Nature Malaysiana 16(4):112-129.

After Bleeker, 1878

Original description: Ophiocephalus bankanensis Bleeker, 1852:726. Nieuwe bijdrage tot de kenne der
ichthyolische fauna van het eiland Banka. Natuurkd. Tijdrschr. Neder. Indië 3:715-738. Type locality: Bangka
Island, Malaysia. Locality of holotype unknown.
Synonyms: No known synonyms (Ng and Lim, 1990).
Common names: Bangka (or Banka) snakehead; runtuk (Kalimantan).

Native range: Sumatra: southeastern rivers (Hari swamps (Lee and Ng, 1991; Ng and Lim, 1991). Also
and Musi basins) of mainland; Bangka Island; rivers of found in mouths and middle reaches of rivers (Lee and
central, southern, and western Kalimantan (southern Ng, 1994).
Borneo; Roberts, 1989; Ng and Lim, 1991; Kottelat Temperature range: Preferred range 26-30 ºC
and others, 1993); peat swamps of Selangor, peninsular (Lee and Ng, 1994).
Malaysia (Lee and Ng, 1991). Reproductive habits: No specific information,
Introduced Range: No introductions known. but likely a nest builder with pelagic eggs.
Feeding habits: No specific information, but
Size: No specific information; Musikasinthorn probably a carnivorous predator.
(2000) examined specimens up to 14 cm standard
Characters: Patch of scales present on gular
length.
part of head. Large canine-like teeth on prevomer and
Habitat preference: Prefers submerged vegeta- palatines. Lateral line scales 55-68; scale rows between
tion in tannin and humic acid enriched backwaters (pH lateral line and dorsal origin 4½; scale rows below lateral
2.8-3.8) and moderately fast-flowing streams in peat line and anal fin origin 7-9; preopercular scales 5-10.

57
Dorsal fin rays 31-45, anal fin rays 20-31. Superficially Commercial importance in native range:
resembles Channa lucius, head taller and more blunt in Probably not of commercial importance due to habitat
C. bankanensis (see Ng and Lim, 1990, p. 142, fig. 5C, preferences. Lee and Ng (1991) stated that this species
D), and body more compressed in C. lucius (Lee and is not popular with local anglers due to its small size.
Ng, 1991). Somewhat rounded, dark blotch on opercu- Environmental concerns: Unknown, but proba-
lum of adult, not elongated as in C. lucius (Ng and bly a predator on other fishes and invertebrates. Habitat
Lim, 1990). preferences indicate the species might only become prob-
Commercial importance in the United States: lematic in highly acidic waters. Stoye (1935) mentioned
Not listed on aquarist-oriented websites. availability of this species as an aquarium fish.

100°E 120°E 140°E

China

Myanmar PACIFIC
20°N
OCEAN
Laos
Thailand Philippines
Vietnam

EXPLANATION Cambodia
DISTRIBUTION OF
Channa bankanensis Malaysia
Malaysia
Native range
INDIAN Indonesia Equator
0°
OCEAN
0 500 MILES
Indonesia Indonesia
0 500 KILOMETERS
Scale is approximate

Channa bankanensis

58
 Channa baramensis (Steindachner, 1901)
Baram Snakehead

After Steindachner, 1901

Live coloration (14.7 cm)

Reprinted with permission from H.H. Ng and P.K.L. Ng from: Ng, H.H., and others, 1996. Revalidation of
Channa baramensis (Steindachner, 1901), a species of snakehead from northern Borneo (Teleostei:
Channidae). Sarawak Mus. J. 48(69 n.s.):219-226.

Original description: Ophicephalus baramensis Steindachner, 1901:435, pl. 17. Kükenthal’s Ergebnnisse einer
zoologischen Forschungriese in den Molukken und Borneo. Abh. Senckenb. Naturforsch. Ges. 25:409-464,
pls. 17-18. Type locality: Baram River, northern Sarawak (northern Borneo). Syntypes: SMF 860; SMF 8473.
Synonyms: Ophicephalus melasoma (Bleeker, 1851) fide Weber and de Beaufort (1922); Myers and Shapovalov
(1932); Roberts (1989); Ng and Lim (1990); and Rainboth (1996). Ng and others (1996) revalidated the species and
Musikasinthorn (2000) also recognized the species as valid.
Common names: Baram snakehead; barama snakehead.

59
 Native range: Northern Sarawak, Brunei, and profile of C. melasoma distinctly sharper. Postorbital
western Sabah (northern Borneo). Also occurs in the depth greater in C. baramensis than in C. melasoma
Sadong basin, southern Sarawak, and the Segama (33.3-34.1 percent of head length versus 27.9-31.8
basin, eastern Sabah (Martin-Smith and Hui, 1998). percent of head length in C. melasoma) (Ng and others,
Introduced range: None. 1996). There are also distinct differences in coloration.
Size: To about 22 cm. In closely related C. melasoma and C. cyanospilos,
Habitat preference: Known from blackwater black pigment (melanin) appears as evenly spread over
swamps (Ng and others, 1996) and small to moderate- each scale, whereas in C. baramensis, melanin is
sized streams, clear or turbid, in secondary growth or concentrated in the central part of most scales in adults
forest areas (Inger and Kong, 1962). and most specimens smaller than 120 mm standard
Temperature range: No specific information, length. Moreover, there is a distinct barred pattern on
but native range is tropical (about 3-6º N). the caudal fin in adult C. baramensis that is absent in
Reproductive habits: Habits can be inferred C. melasoma and C. cyanospilos of similar lengths.
from those of its closest relative, Channa melasoma. This character, however, cannot be used to identify
Doubtlessly a nest builder like other channids with specimens smaller than about 120 mm standard length
probably only one parent guarding eggs and young. (Ng and others, 1996).
Likely a nocturnal species. Commercial importance in the United States:
Feeding habits: Like Channa melasoma, proba- Unknown to have been imported for any purpose.
bly a nocturnal thrust predator that feeds on other
Commercial importance in native range:
fishes, small reptiles, crabs, insects and insect larvae.
Reported as caught by anglers, indicating use as a food
Characters: No patch of scales on gular region.
fish (Ng and others, 1996). Probably only of minor
Dorsal fin rays 38-40; anal rays 23-26. Predorsal scales
commercial importance.
8-9; lateral line scales 51-52. This species appears to be
most closely related to Channa melasoma, and some Environmental concerns: This species is proba-
characters overlap in both species (Ng and others, bly a nocturnal thrust predator. Its limited range in the
1996). Nevertheless, adults (120 mm or more standard tropics would restrict its ability to establish in all but
length) can be separated as follows: Lateral head the warmest waters of the U.S.

100°E 120°E 140°E

China

Myanmar PACIFIC
20°N
OCEAN
Laos
Thailand Philippines
Vietnam

EXPLANATION Cambodia
DISTRIBUTION OF
Channa baramensis Malaysia
Malaysia
Native range
INDIAN Indonesia Equator
0°
OCEAN
0 500 MILES
Indonesia Indonesia
0 500 KILOMETERS
Scale is approximate

Channa baramensis

60
 Channa barca (Hamilton, 1822)
Barca Snakehead

After Hamilton, 1822

Original description: Ophiocephalus barca Hamilton, 1822:67, pl. 35, fig. 20. An account of the fishes found in
the River Ganges and its branches. Edinburgh and London. i-vii + 1-405, pls. 1-39. Type locality: Brahmaputra
River, near Goalpara, Assam, India. Types unknown.
Synonyms: Ophicephalus nigricans Cuvier, 183:431.
Common name: barca snakehead.

Native range: Endemic to Ganges and on palatines. Scales on top of head large; 9 scale rows
Brahmaputra River basin, India and Bangladesh between preopercular angle and posterior border of
(Musikasinthorn, 2000). Bhuiyan (1964) cited its orbit; predorsal scales 15; 60 to 65 scales in longitudi-
presence in eastern and some areas of western Pakistan nal series. Dorsal fin rays 47-52; anal fin rays 34-36;
but this may be a misidentification. pectoral rays 16; pelvic fin rays 6. Life colors violet on
Introduced range: No introductions known. back fading to dull white with purple cast on sides;
Size: To 90 cm (Talwar and Jhingran, 1992). back and sides with large black blotches, as are dorsal,
Habitat preference: Large rivers (Talwar and anal, and caudal fins; fin edges red; pectoral fins red
Jhingran, 1992). with numerous black spots.
Temperature range: No specific information. Commercial importance in the United States:
Nevertheless, its native range is located between about Typically not listed on aquarist-oriented websites.
25-27º N, suggesting it is a warm temperate species. Likelihood of being imported for sale in aquarium fish
Reproductive habits: No detailed information, trade or live-food fish markets has been low to proba-
but like other snakehead species, it is assumed to clear bly nonexistent.
a nest in nearshore vegetation, lay pelagic eggs which, Commercial importance in native range:
following fertilization, rise to the surface where they While reported as common in the Brahmaputra River,
are guarded vigorously by one or both parents until Assam, India, it is said to be of minor importance as a
hatching. fishery resource. Nevertheless, it is considered an
Characters: Body elongated, mostly rounded. excellent food fish (Talwar and Jhingran, 1992).
Mouth large; lower jaw with a few canines behind a Environmental concerns: Like other snake-
single row of villiform teeth that widen to 5 or 6 rows heads, adults are carnivorous predators, most preferring
at jaw symphysis; 2 or 3 large teeth on vomer and some other fishes as food.

61
 80°E 100°E
Nepal
Bhutan China

India
EXPLANATION Myanmar Laos
DISTRIBUTION OF 20°N
Channa barca Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate
Malaysia

Equator Indonesia
0°

Channa barca

62
 Channa bleheri Vierke, 1991
Rainbow Snakehead

Reprinted with permission from Jörg Vierke from: Vierke, Jörg. 1991a. Ein farbenfroher neuer
schlangenkopffish aus Assam Channa bleheri spec. nov. Das Aquarium 259:21.

Original description: Channa bleheri Vierke, 1991a:20-24. Ein farbenfroher neuer Schlangenkopffisch aus Assam
Channa bleheri spec. nov. Das Aquarium 259:20-24. Type locality: upper part of Dibru River, near Guijan,
Brahmaputra River basin, northeastern Assam, India. Holotype: ZFMK 16555. Paratype: ZFMK 16556.
Synonyms: No synonyms.
Common name: rainbow snakehead.

Native range: Endemic to the Brahmaputra female, with the two wrapping around each other. This
River basin, Assam, India (Musikasinthorn, 2000). behavior increases at spawning, near the surface, and
Introduced Range: No introductions known. the spawning act can last up to 30 seconds.
Size: To about 20 cm. The eggs released are transparent and float to the
Habitat preference: Forest streams, ponds and surface. They are small, round, and 0.9-1.1 mm in
swamps in tropical rainforest conditions (Musikasin- diameter. An oil globule, about 0.6-0.7 mm in size, is
thorn, 2000). present in each egg. The egg mass, with eggs close
Temperature range: Unknown, except preferred together, appears to be made of foam. Both the male
habitat and known range is subtropical. and female initially tend the egg mass and display an
interesting behavior. They take the eggs into the oral
Reproductive habits: No specific information cavity and expel them through the gills, presumably to
concerning wild populations. Vierke (1991b), however, remove materials that may settle on egg surfaces. The
described reproductive behavior in aquaria in detail. He eggs adhere to each other at the surface. Following
noted that females are smaller than males of the same
hatching, both parents guard the larvae.
age, and that males grow faster than females. A male
selects the nesting site, but it is the female that appears The young remain around the parents, often with
to initiate courtship behavior. Two days prior to spawn- body contact between them. Young can often be found on
ing, there is frequent body contact between male and the heads of parents, appearing to be feeding on mucus.

63
They appear to nip the parents, and removal of young This species appears to be most closely related to
from parents at this stage seems to slow growth of the Channa burmanica (Peter Ng, personal commun., in
separated individuals. Vierke, 1991b). They differ in several characters.
Channa burmanica has 51 lateral line scales, 28 anal
Feeding habits: No information concerning wild
rays, and 8 predorsal scales, whereas these counts in
populations, but likely a carnivorous predator as an C. bleheri are 45-46, 24, and 6-7, respectively. The
adult. Vierke (1991b) noted that in aquaria, rainbow rainbow snakehead also has a longer caudal peduncle
snakeheads will feed on worms similar to bloodworms. than C. burmanica with 9 scales from the posterior end
When fed guppies (Poecilia reticulata), they will eat of the anal fin to the caudal fin base in C. bleheri and
guppies they can easily catch, but typically tire of chas- 4+2 in C. burmanica (Vierke, 1991b).
ing this prey, eventually tolerating their presence. Commercial importance in the United States:
Characters: No area of scales in gular region. This species is sometimes listed on aquarist-oriented
No pelvic fins. Dorsal rays 36-37; anal rays 24; predor- websites and has been available for sale through aquar-
sal scales 6-7; lateral line scales 45-46. One or 2 large ium fish retailers. Because of its attractive coloration, it
scales on undersurface of lower jaw. appears to have been increasing in popularity as an
aquarium species. An aquarium fish dealer in Kentucky
Live adults have large irregular red or orange was found to be selling this species illegally, having
spots (white in preserved specimens) that sometimes imported them from a supplier in Atlanta, Georgia,
coalesce (Musikasinthorn, 2000). This is the most where snakeheads are also illegal (Major David Casey,
colorful of all snakehead species. Newly hatched larvae personal commun., 2002).
are colorless. When they reach a length of 1 cm, the Commercial importance in native range: This
dorsum and sides of the body becomes “canary” species is caught commercially for the aquarium fish
yellow, with a dark bar on the head, angled from the tip trade (Ralf Britz, personal commun., 2002) and is not
of the lower jaw, through the eye, to the upper margin known to be cultured for this purpose.
of the operculum; the ventral side is colored smoke Environmental concerns: If released into U.S.
gray to black. As young continue to grow, their color waters, it could become established in subtropical Flor-
becomes more pale and an ocellus appears on the ida, Hawaii, perhaps southeastern Texas, and thermal
posterior part of the dorsal fin. Later they begin to springs and their outflows in western states. Probably
change to adult coloration (Vierke, 1991a). predacious on other fishes.

80°E 100°E
Nepal
Bhutan China

India
EXPLANATION Myanmar Laos
DISTRIBUTION OF 20°N
Channa bleheri Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate
Malaysia

Equator Indonesia
0°

Channa bleheri

64
 Channa burmanica Chaudhuri, 1919
Burmese Snakehead

After Chaudhuri, 1919

Left: dorsal view; Right: ventral view

Original description: Channa burmanica Chaudhuri, 1919:284-286, fig. 4. Report on a small collection of fish
from Putao (Hkamti Long) on the northern frontier of Burma. Records of the Indian Museum 16:271-287. Type
locality: Putao Plains of northern Myanmar, river Sen-Ben-Ti. Holotype: ZSI F9755.
Synonyms: Synonyms unknown.
Common name: Burmese snakehead.

Native range: Endemic to headwaters (Kiu Reproductive habits: No specific information.

River, perhaps Lang basin) of the Ayeyarwaddy The Burmese snakehead may show reproductive habits
(=Irrawaddy) River in northern Myanmar, between the similar to its closest relative, Channa bleheri.
Kumon and Shan-ngaw mountain ranges. Feeding habits: No specific information.
Introduced range: No introductions known. Perhaps similar to that of Channa bleheri.
Characters: No area of scales in the gular
Size: Maximum size unknown. Chaudhuri’s
region. No pelvic fins. Dorsal rays 38; anal rays 28.
(1919) largest specimen (one of four) was 106 cm
Lateral line scales 51 (50 pored scales), with lateral line
total length. The species doubtlessly reaches a greater
dipping ventrally after the 12th scale (scale in disjunc-
length, but cannot be considered as one of the moderate-
tion without pore). Predorsal scales 8.
to-large snakehead species.
This species appears to be most closely related to
Habitat preference: No specific information in Channa bleheri (Peter Ng, personal commun., in
Chaudhuri’s (1919) publication. Vierke, 1991b). See account for C. bleheri for species
Temperature range: No specific information. differences.
The type locality (Putao Plains) is about 27º N, indicat- Commercial importance in the United States:
ing a subtropical to warm temperate species. None known.

65
 Commercial importance in native range: in much of peninsular Florida, Hawaii, perhaps south-
Unknown. ern Texas, and thermal springs and their outflows in the
American west if introduced.
Environmental concerns: Doubtlessly a
predator and likely feeding on other fishes. Like
Channa bleheri, this species has potential to establish

80°E 100°E
Nepal
Bhutan China

India
EXPLANATION Myanmar Laos
DISTRIBUTION OF 20°N
Channa burmanica Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate
Malaysia

Equator Indonesia
0°

Channa burmanica

66
 Channa cyanospilos (Bleeker, 1853)
Bluespotted Snakehead

Reprinted with permission from P.K.L. Ng from: Lee, P.G., and P.K.L. Ng. 1991. The snakehead fishes of
the Indo-Malayan region. Nature Malaysiana 16(4):112-129.

After Bleeker, 1878

Original description: Ophicephalus cyanospilos Bleeker, 1853:256. Bleeker, P. Disgnostiche beschrijvingen van
nieuwe of weining bekende vischsoorten van Sumatra. Tiental V-X. Tijdrschr. Neder. Indië 4:243-302. Type locality:
Telok Betong (presently Bandar Lampung), southern Sumatra, Indonesia. Holotype locality unknown.
Synonyms: (?)Ophiocephalus striatus Weber and de Beaufort, 1922.
Channa sp. Ng and Lim, 1990.
(?)Channa striata Ng and Lim, 1990.
Common names: None known. The authors propose bluespotted snakehead as this character remains obvious in
preserved specimens and is unknown in other channids from Indonesia or Malaysia (Ng and Lim, 1991).

Native range: Sumatra and probably peninsular Temperature range: No specific information.
Malaysia and Kalimantan (Kapuas basin, western The known native range of this species is between 3º N
Borneo; Ng and Lim, 1991). Also found during 1995- and 6º S, indicating a tropical, equatorial taxon.
1996 in Riau and Jambi, central Sumatra (Peter Ng, Reproductive habits: No information located.
personal commun., 2003). Probably a nest builder that guards its eggs and young
Introduced range: No introductions known. like other snakeheads.
Size: To at least 20 cm (Ng and Lim, 1991). Feeding habits: No information found. Likely a
Habitat preference: No specific information, thrust predator as other snakeheads.
but known from a tributary in the Sungei Alas (Alas Characters: No patch of scales in gular region
River) basin, northern Sumatra. of head. Dorsal fin rays 38-43; anal fin rays 24-26.

67
Lateral line scales 51-55; predorsal scales (posterior to Pale blue spots or blotches in the throat region of
cephalic shields) 8. Small canines present on lower jaw. C. cyanospilos are similar to those in C. melasoma, but
Pale blue spots on the lower half of the body from gill in the latter species the spots and blotches form a
cover to caudal peduncle, remaining visible in marbled pattern; C. striata lacks any blue spots in the
preserved specimens. throat region but has brown streaks and spots (Ng and
Weber and de Beaufort (1922) listed this species Lim, 1991).
as a possible synonym of Channa striata, a practice Commercial importance in the United States:
followed for many decades. Ng and Lim (1990) cited Not listed on aquarist-oriented websites and probably
the species in this same manner. Ng and Lim (1991), has been unavailable in the aquarium fish trade. Not
however, recognized the species as valid as did Kottelat known to have been imported or available in live-food
and others (1993) and Musikasinthorn (2000). Ng and fish markets.
Lim (1991) allied C. cyanospilos with C. melasoma,
rather than C. striata, based on morphological features, Commercial importance in native range: No
particularly with regard to the shapes of the throat information found. Probably occasionally caught by
region and ventral surfaces of the gill cover. angling.
Channa cyanospilos can be separated from Environmental concerns: Likely a thrust preda-
C. melasoma by lower jaw length (5 percent standard tor with a diet that includes fishes. Nevertheless, this
length in C. cyanospilos, 12-13 percent in species is tropical and probably would survive only in
C. melasoma). Both species also have 8 predorsal extreme southern Florida, Hawaii, and thermal springs
scales behind the cephalic shields (7 in C. striata). and their outflows if introduced.

100°E 120°E 140°E

China

Myanmar PACIFIC
20°N
OCEAN
Laos
Thailand Philippines
Vietnam

EXPLANATION Cambodia
DISTRIBUTION OF
Channa cyanospilos Malaysia
Native range Malaysia
Possible native INDIAN Indonesia Equator
range 0°
OCEAN
0 500 MILES
Indonesia Indonesia
0 500 KILOMETERS
Scale is approximate

Channa cyanospilos

68
 Channa gachua (Hamilton, 1822)
Dwarf Snakehead

After Munro, 1955

Upper image: ZRC 41100, 107.7 mm standard length, from Perlis, Malaysia. Lower image: ZRC 41656,
46.5 mm standard length, from Kerala, India. Reprinted with permission from P.K.L. Ng from: Ng, H.H.,
P.K.L. Ng, and Ralf Britz. 1999. Channa harcourtbutleri (Annandale, 1918): a valid species of snakehead
(Perciformes: Channidae) from Myanmar. J. South Asian Nat. Hist. 4(1):57-63.

Original description: Ophicephalus gachua Hamilton, 1822:68, 367. An account of the fishes found in the river
Ganges and its branches. Edinburgh and London, i-xii + 1-405. Type locality: ponds and ditches of Bengal, India.
Type specimens unknown.
Synonyms: Ophiocephalus aurantiacus Hamilton, 1822:69, 368, pl. 23, fig. 22.
Ophicephalus marginatus Cuvier, 1829:230, fide Roberts, 1993:38, Menon, 1999, and Ng and others,
1999:57.
Ophicephalus limbatus Cuvier, 1831, no p., pl. 201, fide Roberts, 1993:40, Menon, 1999:275, and
Ng and others, 1999:57.

69
 Ophicephalus marginatus Cuvier, 1831, no p., pl. 201, fide Roberts, 1993:38, Menon, 1999:275, and
Ng and others, 1999:57.
Ophicephalus coramota Cuvier, 1831:414, fide Menon, 1999:275 and Ng and others,1999:57.
Ophicephalus fuscus Cuvier, 1831:414, fide Menon, 1999:275 and Ng and others, 1999:57.
Ophicephalus montanus McClelland, 1842:583, fide Menon, 1999:275, and Ng and others, 1999:57.
Philypnoides surakartensis Bleeker, 1849:19, fide Menon, 1999:275, and Ng and others, 1999:57.
Ophiocephalus apus Canestrini, 1861:77, pl. 4, fig. 7.
Ophiocephalus kelaartii Günther, 1861:472, fide Talwar and Jhingran, 1991;1019, Menon, 1999:275,
and Ng and others, 1999:57.
Ophiocephalus guachua [sic] var. malaccensis Peters, 1868.
Ophicephalus gachna [sic] Jordan and Seale, 1907.
Ophiocephalus gachua kelaarti Günther: Munro, 1955:100.
Species is in need of revision; status of many synonyms uncertain (Ng and Lim, 1990; Ng and others, 1999).
Roberts (1989), Lim and others (1990), Talwar and Jhingran (1992), and Rainboth (1996) treated this species as a
synonym of Channa orientalis, but Ng and others (1999) disagreed with this conclusion. Day (1889), Coad (1981),
Lim and others (1990), Kottelat (1998), Musikasinthorn (2000), and others considered C. gachua to be valid. For
purposes of this report, we follow the most recent authorities in recognizing the species as valid, but also realizing
that it doubtlessly represents a species complex.
Common names: dwarf snakehead; frog snakehead; brown snakehead; dolli or dauli (Pakistan); dheri dhok
(Hindu); para korava (Tamil); doarrah (Punjab, India); chen-gah (Assam, India); cheng (West Bengal, India);
dheridhok, chainga (Bijar Province, India); chenga (Orissa Province, India); malamatta-gudisa, erramatta, tatimatta-
gudisa (Andrah Pradesh Province, India); koravu, vattudi (Kerala Province, India); mohkorava, mottu (Karnataka
Province, India; Talwar and Jhingran, 1992), parandal kanaya (Sri Lanka; Pethiyagoda, 1991); bakak (Malay; Lim
and Ng, 1990).

Native range: Bampur-Haliri basin and Mashkel (=Taiwan) that lacked both pelvic fins. Because pelvic
River, southeastern Iran (Coad, 1979); Kabol (Kabul) fins were absent, they tentatively identified the speci-
drainage of Afghanistan (Coad, 1981); eastern and men as C. orientalis but noted other specimens of
western Pakistan (Qureshi, 1965); India, Sri Lanka, C. gachua from India that also lacked pelvics based on
Bangladesh, Myanmar, Thailand, Laos, Cambodia, reports by earlier scientists. They suggested that this
Malaysia, Indonesia (Borneo), Java, southern China fish may have been introduced to Taiwan. This snake-
(Mukerji, 1933; Mendis and Fernando, 1962; Fernando head can be easily transported for great distances with
and Indrassna, 1969; Roberts, 1989; Pearl River Fisher- significant altitude and temperature changes (Day,
ies Research Institute, 1991; Pethiyagoda, 1991; 1877). Peter Ng (personal commun., 2003) found this
Talwar and Jhingran, 1992; Kottelat, 1998, 2001a; snakehead in Bali in 2000, perhaps within its native
Musikasinthorn, 2000). Kullander and others (1999), range or an introduction.
citing Das and Nath (1971), recorded this species from Size: This species is often referred to by authors
the Punch Valley, in a tributary of the Jhelum River, in as the smallest snakehead, reaching a length of about
the Kashmir Valley of northeastern Pakistan/western 17 cm (Kottelat, 1998). Lee and Ng (1991) stated that it
Kashmir. Peter Ng (personal commun., 2003) reported rarely exceeded 20 cm. The smallest snakehead,
its occurrence in the Toba area of northern Sumatra, however, is Channa orientalis, a species that rarely
and Riau and Jambi in central Sumatra in 1996, as well exceeds 10 cm in length (Pethiyagoda, 1991).
as in Yunnan Province, China, in 2000. Habitat preference: Appears to prefer clear
Introduced range: There may have been intro- pools, shallow streams, and swamps, particularly in
ductions that went unrecorded, but Kottelat’s (1985) forested areas. Nevertheless, Pethiyagoda (1991) noted
reference to Channa orientalis from the Greater Sunda that it is common “in streams and ponds,” but is toler-
Islands doubtlessly refers to some species of the ant of “very stagnant, poorly oxygenated, turbid water.”
C. gachua complex. Myers and Shapovalov (1932) Deraniyagala (1929) stated that it “flourishes in ponds
cited a specimen of C. gachua from Formosa rendered so stagnant as to prove toxic to most fishes.”

70
Said to be largely a nocturnal fish. Habitat destruction, per spawning, with the male orally brooding develop-
due to deforestation near Singapore, is considered a ing eggs and fry (Lee and Ng, 1991, 1994), further
threat to this species (http://www.sci-ctr.edu.sg/ssc/ evidence that Channa gachua is a species complex.
imglib/vertebra/channa.html). Kottelat (1998, 2001a,b) This is one of three species of snakeheads known to
reported this species from hill streams in Laos, Thailand, spawn in ponds lacking vascular aquatic plants
Cambodia, and Vietnam. Lim and others (1990) recorded (Parameswaran and Murugesan, 1976b).
finding it in forest streams in peninsular Malaysia. This Bhuiyan and Rahman (1982) measured fecundity
snakehead is reported to tolerate a wide range of pH from 30 female Channa gachua collected near
levels with 100 percent survival over 72 hours from pH Rajshahi, Bangladesh, which ranged from 487 (94 mm
3.10 to 9.6 (Varma, 1979). specimen) to 4,482 (164 mm specimen). Mean fecun-
Lee and Ng (1994) reported the species “from dity of the 30 specimens was 2,307 oöcytes for a speci-
rivers, lakes, ponds, well-shaded, small forest streams men having a mean length of 132 mm. The relationship
less than 20 cm deep, hillstreams not continuous to between fecundity and length is largely linear, as is the
mountain ranges and in the upper zone of rivers.” They relationship between fecundity and length of ovaries.
indicated that it is found from sea level to an altitude of Mishra (1991) described mature (stage V)
1,520 m in India and up to 1,430 m in Malaysia, in oöcytes as ranging from 2.1 to 2.6 mm in diameter with
waters with flow rates from stationary to rapids with a the highest percentage of stage V oöcytes in July from
pH range of 3.1-9.6. Lee and Ng (1991) stated that the specimens collected near Berhampur, Orissa, India.
species is also found in mountain brooks up to 3,600 m The highest gonadosomatic index was 6.8 and occurred
above sea level. Mukerji (1931) reported capture of two in June. Estimated fecundity ranged from 2,539 to
specimens from the Billigirirangan Hills of southern 7,194 in 15 mature specimens ranging from 13.4 to
India at an altitude of 751 m. The species has been 17.2 cm in length. Again, the relationship between
reported surviving in brackish water, but this is doubted fecundity and length, as well as fecundity and body
by Lee and Ng (1991). Srivastava and others (1980) weight, was largely linear.
noted decreasing body weight of this species in salini- Feeding habits: Lee and Ng (1994) summarized
ties as low as 5 ppm. They are capable of overland food preferences as including “mouse, rat, frog, tadpole,
migrations (Lee and Ng, 1991), and Deraniyagala fish, Ephemerophtera and other insects, mosquito
(1929) commented that it is “exceedingly active on larvae, prawn (Macrobrachium sp.), crab (Irmengardia
land, pregressing by a series of leaps.” johnsoni), and other crustaceans.” They cited the
Temperature range: Lee and Ng (1994) indi- species as a nocturnal predator living at or near the
cated that this species can tolerate temperatures in hot substrate and quite capable of migrating overland.
springs in Sri Lanka to as low as 13 ºC. The species Characters: No patch of scales in the gular
complex, however, is reported from as far north as region. Pelvic fins present, although Talwar and Jhing-
Afghanistan (with cold winters) to Borneo and Java ran (1992) stated that pelvics may be present or absent.
(equatorial tropical). Pethiyagoda (1991) cited Lateral line scales 39-47; 3½ scales between lateral line
Deraniyagala (1932) as having recorded this species and base of anterior dorsal rays. Dorsal rays 32-37;
from hot springs at 36.5 ºC. anal rays 20-23. Lower jaw with 10-20 canines poste-
Reproductive habits: This species has been rior to a single row of villiform teeth, the latter expand-
cited as a mouthbrooder (Lee and Ng, 1991, 1994; ing to about 7 rows at the jaw symphysis. Dorsal, anal,
Kottelat, 2001a) and confirmed by Ralf Britz (personal and caudal fins with white (translucent in preserved
commun., 2003). An interesting factor here is that specimens) margin; ocellated spot often present near
other authors (for example, Munro, 1955) made no posterior end of dorsal fin. The ocellated spot, however,
mention of oral brooding of fertilized eggs and young may appear only in juveniles and females as occurs in
for the dwarf snakehead. Based on Kahn (1924) and Channa orientalis. Lim and others (1990) noted that
Deraniyagala (1929), Breder and Rosen (1966) stated while the fish is alive the dorsal, anal, and caudal fins
that spawning in India occurs with the female swim- are margined with red or yellow, and that the pectoral
ming upside-down under the male, with eggs being fins have semiconcentric rings and a dark area at
released and fertilized in groups of 200-300 every the base.
minute or two. Females in Indonesia and Malaysia, Commercial importance in the United States:
however, are reported to produce from 20 to 200 eggs This species is occasionally mentioned on aquarist-

71
oriented websites and has been available for sale from snakehead is utilized as food by “only the poorest
certain aquarium fish dealers. Its small size makes it classes” in Sri Lanka, adding that it is used as live bait
more appealing as an aquarium species. Too small to be to catch larger species, such as Channa striata and
sold in live-food fish markets. C. marulius.
Commercial importance in native range: Environmental concerns: Like other channids,
Talwar and Jhingran (1992) cited the species as of this species is a thrust predator, capable of breathing
minor importance in India. According to comments on atmospheric oxygen. This species is noted by many
aquarist-oriented websites and in Ng and Lim (1990), authors as migrating overland.
members of this species complex appear to be of Comments: The diploid number of chromo-
importance in the aquarium fish trade with individuals somes for Channa gachua is 78 from India (Banerjee
captured from the wild for sale or export. Ng and Lim and others, 1988) and 112 for specimens from Thailand
(1990) cited individuals being sold for S$30-60 in (Donsakul and Magtoon, 1991), a strong indication that
Singapore, although Lim and Ng (1990) listed the this represents a species complex.
species as endangered there. Pethiyagoda (1991) noted
that it is largely unused as food or in the aquarium trade
in Sri Lanka. Deraniyagala (1929) commented that this

60°E 80°E 100°E 120°E

North
40°N
Korea

Afghan-
Iran istan China South
Nepal Korea
Bhutan
Pakistan

Saudi India
Arabia Myanmar
20°N

Laos
Bangladesh Thailand
Vietnam

Cambodia
Thailand
Malaysia

Equator
0°

INDIAN OCEAN Indonesia

EXPLANATION 0 1,000 MILES

DISTRIBUTION OF
Channa gachua 0 1,000 KILOMETERS
Native range Scale is approximate

Channa gachua

72
 Channa harcourtbutleri (Annandale, 1918)
Inle Snakehead

Upper image: ZRC 42556, 111.3 mm standard length, Myanmar: Inle Lake. Lower image: ventral view of
head illustrating pigmentation, ZRC 42556, 111.3 mm standard length. Reprinted with permission from P.K.L.
Ng from: Ng, H.H., P.K.L. Ng, and Ralf Britz. 1999. Channa harcourtbutleri (Annandale, 1918): a valid species
of snakehead (Perciformes: Channidae) from Myanmar. J. South Asian Nat. Hist. 4(1):57-63.

Original description: Ophiocephalus harcourtbutleri Annandale, 1918:54, pl. 2, fig. 7; pl. 4, figs. 16-17. Fishes
and fisheries of the Inlé Lake. Rec. Indian Mus. (Calcutta), 14:33-64, pls. 1-7. Type locality: southern Shan State,
Myanmar (Burma). Holotype: ZSI F9439/1.
Synonyms: (?) Ophioceplahus gachua (non-Hamilton, 1822) Boulenger, 1899:199.
Ophiocephalus harcourt-butleri Annandale, 1918:54.
(?) Ophicephalus gachua (non-Hamilton, 1822) Hora and Mukerji, 1934:135.
Ophiocephalus harcourtbutleri Tint Hlaing, 1971:517.
Channa orientalis Kottelat, 1989:20; Talwar and Jhingran, 1991:1019.
Channa harcourtbutleri Ng and others, 1999.
Common name: Inle snakehead.

Native range: Yawnghwe and nearby areas of Temperature range: No specific information in
Myanmar, particularly Inlé Lake in southern Shan State literature. Nevertheless, Inlé Lake lies at about 20.7º N,
(Ng and others, 1999). indicating that Channa harcourtbutleri is a tropical/
subtropical species. Inlé Lake is at an altitude of about
Introduced range: Not known to have been 1,000 m (Kullander and others, 2000).
introduced. Reproductive habits: No specific information in
Size: To about 16 cm. literature. Ralf Britz (personal commun., 2003) indi-
cated that it is unknown if this species is a nest builder
Habitat preference: Annandale (1918) noted or mouthbrooder.
that the species occurred in Inlé Lake on “muddy Food preferences: No specific information in
bottom in sluggish streams … hiding as a rule among literature. Likely a thrust predator as are other snake-
weeds.” heads (Ng and others, 1999).

73
 Characters: No patch of scales in gular region The two species also differ in coloration (Ng and
of head. Lateral line curves downward between scales others, 1999) with C. harcourtbutleri never having an
15-16. Lateral line scales 44-45; transverse scales 4; ocellus at the posterior end of the dorsal fin (occurring
predorsal scales 4. Dorsal fin rays 34-38 (mode 36); in subadult C. gachua).
anal fin rays 23-26; pectoral fin rays 14-15 (mode Commercial importance in the United States:
14) (Ng and others, 1999). This species is most Not known to have been imported for any purpose.
closely related to Channa gachua from which it Commercial importance in native range: Sold
can be distinguished as follows: Head profile of in live-food fish markets in the vicinity of Inlé Lake,
C. harcourtbutleri flatter and less convex than in Myanmar (Ng and others, 1999).
C. gachua resulting in a smaller postorbital head depth Environmental concerns: Although this species
(30.9-35 percent head length versus 39.8-44 for is tropical/subtropical and could potentially establish if
C. gachua), its length 32-34 percent standard length released in southern Florida, Hawaii, and warm thermal
versus 26.8-31.9 percent, and width 16.9-19 percent springs and their outflows, its food habits are largely
standard length versus 18.6-21.7 percent. Moreover, unknown (Ralf Britz, personal commun., 2003). Like
the snout of C. harcourtbutleri is more convex when other snakeheads, it may be a predator (Ng and others,
viewed dorsally (Ng and others, 1999, figs. 3b,4a). 1999).

80°E 100°E
Nepal
Bhutan China

My
India an
ma
EXPLANATION r Laos
DISTRIBUTION OF 20°N
Channa harcourtbutleri Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate
Malaysia

Equator Indonesia
0°

Channa harcourtbutleri

74
 Channa lucius (Cuvier, 1831)
Splendid Snakehead

Upper image: adult. Lower image: juvenile.

Reprinted with permission from P.K.L. Ng from: Lee, P.G., and P.K.L. Ng. 1991. The snakehead
fishes of the Indo-Malayan region. Nature Malaysiana 16(4):112-129.

After Bleeker, 1878

Original description: Ophicephalus lucius Cuvier in Cuvier and Valenciennes, 1831:416. Histoire naturelle des
poissons 7:i-xxix + 1-537, pls. 170-208. Type locality: Java. Holotype: RMNH, whereabouts unkown.
Synonyms: Ophiocephalus polylepis Bleeker, 1852:578.
Ophiocephalus bistriatus Weber and de Beaufort, 1922.
Ophiocephalus bivittatus Károli, 1822. Name preoccupied by Ophiocephalus bivittatus Bleeker, 1845,
changed to Channa bistriata by Weber and de Beaufort, 1922.
Ophicephalus siamensis Günther, 1861 (Musikasinthorn and Taki, 2001).
Channa bistriata Weber and de Beaufort, 1922 (young of C. lucius [Alfred, 1964]).
Common names: splendid snakehead; forest snakehead; ikan bujok or ikan ubi (Malaysia), runtuk (Kalimantan),
trey kanh chorn chey (Cambodia); bujok (Mayay; Lim and Ng, 1990).

75
 Native range: Rivers of southeastern Sumatra Lateral line scales 58-65; 5½ scales between lateral line
and the Kapuas basin of western Kalimantan (southern and base of anterior dorsal rays. Dorsal rays 38-41; anal
Borneo; Roberts, 1989); Mekong basin of Laos rays 27-29. Sides of body with series of distinct, dark
(Kottelat, 2001a). Kottelat (1985) and Ismail (1989) “porthole” blotches, oblique bars on belly, and slightly
included China, Vietnam, Laos, Thailand, Malaysia, elongated, dark blotch on operculum (Ng and Lim, 1990;
Kalimantan, Java, and Sumatra in the native range. Lee and Ng, 1991). Juveniles with three dark stripes from
Peter Ng (personal commun., 2003) collected this head to caudal fin base.
species during November 1999 and April 2000 in Ismail (1989) stated that this species has one or
central Sumatra, southern Sarawak, and the Mahakam two rows of primarily canine teeth on the prevomer and
and Kayan basins of eastern Kalimantan. palatines.
Introduced range: No introductions known. Commercial importance in the United States:
Size: To 40 cm. Rarely mentioned on aquarist-oriented websites and
Habitat preference: Kottelat (2001a) stated probably of little or no importance in the domestic
preference for streams in forested areas where the aquarium fish trade. No information on its past avail-
species lives among vegetation. Lim and Ng (1990) ability in live-food fish markets.
indicated a preference for forest streams and peat Commercial importance in native range: Ng
swamps. Lee and Ng (1994) said the species occurs in and Lim (1990) listed this species as the third most
lakes, ponds, shaded forest streams, peat swamps, and highly prized food fish in southeastern Asia, where
mid-depths of rivers, showing a preference for faster channids bring S$10-20/kg when fresh. Only Channa
moving waters. pH range is 5.5-6.0 and the species is micropeltes and C. striata are more popular in live-
said to be crepuscular or nocturnal (Lee and Ng, 1994). food fish markets. It is also sold in the Singapore
Rainboth (1996) cited preferred habitat as “slowly aquarium fish trade (Ng and Lim, 1990). It is sold
moving streams and rivers as well as lakes, ponds, and fresh, frequently alive, in Cambodia (Rainboth, 1996).
reservoirs from Thailand to Indonesia. Usually found Environmental concerns: This species is known
in areas with much aquatic vegetation as well as to be a nocturnal thrust predator, with a preference for
submerged, woody plants…” other fishes. Its natural range includes tropical and
Temperature range: Lee and Ng (1994) subtropical climate zones, indicating that if introduced,
recorded the fish in waters between 24-29 ºC, indicating this species could establish only in similar climates or
a tropical/subtropical species. thermal springs and their outflows.
Reproductive habits: A nest builder like other Comments: The diploid number of chromosomes
channids, with both parents guarding developing eggs of Channa lucius is 48 (Donsakul and Magtoon, 1991).
and larvae (Lee and Ng,
1994). No information 100°E 120°E 140°E
located on fecundity. China
Food preferences:
Myanmar PACIFIC
Lee and Ng (1994) cited the 20°N
OCEAN
species as “a midwater or Laos
surface predator relying on Thailand Philippines
camouflage to ambush its prey, Vietnam
mainly fish.” Rainboth (1996)
EXPLANATION Cambodia
stated that this species preys
“on fishes, prawns, and crabs DISTRIBUTION OF
Channa lucius Malaysia
and slightly less on shrimps.” Native range Malaysia
Characters: Patch of
INDIAN Indonesia Equator
scales present on gular region 0°
OCEAN
of head. Large canine teeth
present on prevomer and 0 500 MILES
Indonesia Indonesia
palatines. Upper profile of
head somewhat concave. 0 500 KILOMETERS
Scale is approximate

Channa lucius

76
 Channa maculata (Lacepède, 1802)
Blotched Snakehead

Reprinted with permission of Tokai University Press from Masuda, H., K. Amaoka, C. Araga, T. Uyeno, and
T. Yoshino (eds.). 1984. The Fishes of the Japanese Archipelago. © Tokai University Press, Tokyo, Japan.

Original description: Bostrychus maculatus Lacepède, 1801:140, 143. Histoire naturelle des poissons 3:i-lxvi +
1-558, pls. 1-34. Type locality: none designated. No type specimens.
Synonyms: Ophicephalus guentheri Sauvage and Dabry de Thiersant, 1874:4.
Ophicephalus lucius Koller, 1927:41, pl. 1, fig. 7.
Ophicephalus tadianus Jordan and Evermann, 1902:1289.
Ophiocephalus marmoratus Brind, 1914:11.
Common names: blotched snakehead. No other English names known; hei-yü (Tungting Lake, China; Nichols,
1943); tai wan li (Mandarin); lê-hî (Taiwan; Shen and Tzeng, 1993); Taiwan-dojô (Japan and Taiwan; Uyeno and
Arai, 1984; Hosoya, 2002); fibata (Madagascar; Paul V. Loiselle, personal commun., 2003).

Native range: southern China, south of the from 3 to 6 inches in length, were caught by a sailor of
Chang Jiang (Yangtze) basin and Hainan (Nichols, a merchant vessel and brought to New York direct”
1943; Okada, 1960; Hay and Hodgkiss, 1981; Uyeno (Brind, 1914). There are no records of C. maculata
and Arai, 1984); northern Vietnam (Kotellat, 2001a). having been introduced to either locality. The illustra-
Introduced range: Taiwan (established); intro- tion that accompanied his description of this fish,
duced into Japan beginning in 1916 from Taiwan, although somewhat poor, appears to be of C. maculata
established in Nara, Hyôgo, Hiroshima, Mie, and Shiga and most certainly is not of any snakehead native to
prefectures, Japan, as of 1960 (Okada, 1960; Liang and Singapore or Malaysia. Moreover, Brind (1914) com-
others, 1962; Hay and Hodgkiss, 1981; Uyeno and mented that this fish “can stand cold water.” Therefore,
Arai, 1984; Shen and Tzeng, 1993); Philippines we have included O. marmoratus as a synonym of
(Uyeno and Arai, 1984), and Madagascar (misidenti- C. maculata and conclude that either the localities from
fied as Channa striata; Raminosoa, 1987; Welcomme, which these specimens were obtained were in error or
1988; Reinthal and Stiassny, 1991; Stiassny and Rami- that they were acquired from Chinese traders in those
nosoa, 1994; Lever, 1996; Lévèque, 1998). Brind two cities. A somewhat similar situation involving a
(1914) described a snakehead, Ophiocephalus marm- bagrid catfish, Hemibagrus elongatus, the holotype of
oratus, from specimens supposedly acquired in Sin- which was supposedly from Singapore, was reported
gapore and Malacca, Malaysia. “About 60 specimens, (Kottelat and others, 1998) where the actual type locality

77
is believed to be China. Those authors also noted that Size: To 33 cm (Okada, 1960) but reaches a
although the type of H. elongatus might have come length of more than 1 m when fully mature (William S.
from Singapore, there had been active trade in Sing- Devick, personal commun. to Paul L. Shafland, 2002).
apore of fishes imported from China for purposes of Habitat preference: Streams, lakes, ponds and
“food, aquaculture or accidentally” that would explain ditches in southern China; prefers shallow waters with
its Chinese origin. There are no type specimens of vegetation (Okada, 1960; Hay and Hodgkiss, 1981).
O. marmoratus. Temperature range: Native range is subtropical
to warm temperate. Nevertheless, Okada (1960)
This species may be one of the most widely
reported that in Japan, this species tolerated “seven
introduced snakeheads in the Indian and Pacific Ocean
days in 7 ºC air temperature out of water.” Atkinson
basins. As of August 2002, it was becoming apparent
(1977), however, cited it as a “tropical” species. Never-
that many reports of introductions and established
theless, this snakehead has become established far
populations of Channa striata in such places as Hawaii
north of its native range and assumed climate toler-
and Madagascar were based on misidentifications of
ances in Japan, following its introduction there.
C. maculata (Ralf Britz, personal commun., 2002).
Reproductive habits: Builds a circular, open
We have confirmed (October 2002) that the snakehead
nest in vegetation. Eggs float to surface and are
species recorded from Hawaii since the late 1800s is
guarded by parents. Spawns in Japan in early summer
indeed C. maculata and not C. striata. See comments (Okada, 1960).
under Introduced range in the account for C. striata.
Feeding habits: Reported to feed on crusta-
Channa maculata was introduced into Madagas- ceans, large insects, frogs, and fishes (Okada, 1960;
car about 1978 by former President Didier Ratsiraka Hay and Hodgkiss, 1981); described as a “fierce preda-
who had seen snakeheads at an aquaculture facility tory fish” that “hides among rocks or aquatic plants
during an earlier visit to North Vietnam. The species he until its prey approaches, then it quickly attacks, kills,
saw in North Korea was likely C. argus, but the species and swallows its victim” (Hay and Hodgkiss, 1981).
that was shipped to President Ratsiraka from China was Paul V. Loiselle (personal commun., 2003)
C. maculata. The shipment was divided equally, one reported that a fisherman in Madagascar had observed
group stocked into ponds at the presidential summer young blotched snakeheads slithering onto land, allow-
residence near Antananarivo, adjacent to the head- ing their bodies to be covered by ants, then returning
waters of the Betsiboka River. The remaining fish were to the water where the ants floated at the surface and
stocked into ponds at Ratsiraka’s home near Vatoman- were devoured by the juvenile snakeheads. He also
dry on the east coast. Subsequent floods from commented that amphibian populations in Madagascar
monsoons washed snakeheads out of ponds in both are probably being negatively affected by the intro-
localities and into adjacent natural waters. By 1986, duced blotched snakehead. Historically, frog tadpoles
C. maculata was well established in floodplain lakes of in the central highlands were preyed upon to some
the Betsiboka basin. Nearly the same occurred on the extent by native eels (Anguillidae), but now those same
east coast of Madagascar. There is an extensive canal waters contain considerable numbers of Channa
system (Pangalanes Canal) that includes many inland maculata.
lakes along the Indian Ocean coast of Madagascar that Characters: No patch of scales on gular region.
extends for hundreds of kilometers north to south. Head profile slightly depressed. Dorsal rays 40-46;
Snakeheads entered this canal system from ponds near anal rays 26-30. Lateral line scales 41-60 with lateral
Vatomandry and spread rapidly, being recorded about line continuous; transverse scales below lateral line 11;
200 km north of Vatomandry near Toamasina, the 9 scale rows between posterior rim of orbit to upper
northern terminus of the Pangalanes Canal, several edge of operculum. Color pattern similar to that of
years later. It also dispersed southward to the Mangoro Channa argus; dark stripe from tip of snout through
River. The species apparently was also moved to other orbit extending to almost above anterior base of pectoral
areas of Madagascar by Sino-Malagasy merchants fin; second dark stripe from posteroventral corner of
(Paul V. Loiselle, personal commun., 2003). It was orbit to posteroventral edge of operculum; sides of
initially misidentified as C. striata beginning with body with two rows of large, dark blotches extending
Raminosoa (1987). Reports of C. striata from Mauri- posteriorly to anterior caudal peduncle; blotches in the
tius are also possible misidentifications of C. maculata. form of two bar-like markings on caudal peduncle;

78
dark markings toward mid-dorsal part of back extending personal commun., 2002), apparently imported in the
up onto proximal part of dorsal fin. A key character for early 1990s (Domingo Cravalho, Jr., personal
separating this species from C. argus are the bar-like commun., 2002).
markings on the caudal peduncle. In C. maculata, the
most posterior dark bar (usually complete) is preceded Commercial importance in native range:
and followed by pale bar-like areas, whereas in Considered an important and valuable food fish in
C. argus such pale markings are absent and the final China (Nichols, 1943; Atkinson, 1977; Hay and
dark marking is irregular, often blotch-like. Hodgkiss, 1981). It is the second most important
Commercial importance in the United States: snakehead species cultured in China with most culture
Rarely mentioned on aquarist-oriented websites. activity located in Guangdong Province, southeastern
Because it is a valuable food fish in southern China and China (Fang Fang, personal commun., 2002). Liang
Taiwan, we believed it could be available for sale in and others (1962) commented that it is only sold alive
live-food fish markets. Ralf Britz (personal commun., as a food fish within its introduced range in Taiwan.
2002) confirmed that this species was one of the two
Environmental concerns: This species is known
snakeheads purchased from a live-food fish market in
to be a thrust predator feeding on large invertebrates and
Boston, Massachusetts, in November 2001. A reexami-
nation of the second specimen confirmed that it was fishes. Moreover, its temperature tolerance indicates a
also Channa maculata. We also noted from Federal species that could live in subtropical to temperate areas
records that imports of snakeheads from Guangdong in the United States if introduced. That it is established
Province, China, had increased during 2001. Because throughout much of central Japan, an area located gener-
most culture of blotched snakeheads occurs in that ally between 34-37º N, and its southernmost range in
province, the likelihood of availability of this northern China is Hainan Island (about 19º N), is indicative of a
snakehead “look-alike” in U.S. markets was increased. species that has potential to establish from extreme
Ralf Britz (personal commun., 2002) also confirmed southern Florida to North Carolina on the Atlantic Coast
that the blotched snakehead has been present in Hawaii or central California on the Pacific Coast.
since before 1900, misidentified as C. striata, based on
specimens in the U.S. National Museum of Natural Comments: The diploid number of chromo-
History. We have learned that the species presently in somes of Channa maculata is 42 (Wu and others,
culture in Hawaii as of 2002 is C. striata (Pam Fuller, 1986).

See map on following page

79
 60°E 80°E 100°E 120°E 140°E
Mongolia
North
40°N Korea
Japan
Afghan- China
istan Nepal
Iran Bhutan South
Pakistan Korea

India PACIFIC
20°N
Myanmar Laos OCEAN
Bangladesh Vietnam
Philippines
Thailand Cambodia
Malaysia
Equator
0°
INDIAN Indonesia
Indonesia
OCEAN

EXPLANATION 0 2,000 MILES

DISTRIBUTION OF
Channa maculata 0 2,000 KILOMETERS
Native range
Scale is approximate
Introduced range

Distribution of Channa maculata in the Western Pacific basin

This species is also established on Oahu, Hawaii, and may prove to be a
more widely introduced snakehead in the Indian and Pacific Ocean basins
than previously known. See comments above and under Introduced
range in the account for Channa striata.

80
 Channa marulioides (Bleeker, 1851)
Emperor Snakehead

Reprinted with permission from P.K.L. Ng from: Lee, P.G., and P.K.L. Ng. 1991. The snakehead fishes
of the Indo-Malayan region. Nature Malaysiana 16(4):112-129. Adult; photo by J. Vierke.

After Bleeker, 1878

Original description: Ophicephalus marulioides Bleeker, 1851:424. Vijfde bijdrage tot de kennis der
ichthyologische fauna van Borneo, met bescrijving van eeinige nieuwe soorten van zoetwatervisschen. Natuurkd.
Tijdschr. Neder. Indië 2:415-442. Type locality: Sambas, Kalimantan (southern Borneo), Indonesia. Whereabouts of
holotype unknown.
Synonyms: No known synonyms (Roberts, 1989; Ng and Lim, 1990).
Common names: emperor snakehead; darkfin snakehead; ikan jaloi (Malay); toman bunga (=flower snakehead;
Malaysia).
Native range: Rivers (Musi, Hari, Indragiri, and 1994). Also recorded from southern Thailand (Malay
others) of southeastern Sumatra; Kapuas basin of Peninsula) by Herre and Myers (1937) and reported as
western Kalimantan (southern Borneo; Roberts, 1989, the only record from that country (Smith, 1945).
Kottelat, 1994); Bangka (Banka) and Belitung Kottelat and others (1993) did not list Thailand within
(Billiton) (Roberts, 1989). Peter Ng (personal its native range, although it is possible that its range
commun., 2003) collected this species in Samarinda, extends northward into extreme southern Thailand.
eastern Kalimantan, in November 1999. In peninsular Ismail (1989) included Thailand within native range
Malaysia, occurring mostly toward the center of the but added that the species was “quite rare” in
peninsular Malaysia.
peninsula in Pahang (Lee and Ng, 1994). Often
confused with Channa melanoptera (Lee and Ng, Introduced range: No introductions known.

81
 on prevomer or palatines (Smith, 1945; Kottelat and
others, 1993). These characters overlap those of
Channa melanoptera. Lee and Ng (1994) stated that
the only way to separate these two species is by colora-
tion. Channa marulioides possesses an ocellated spot
on the upper part of the caudal fin base, similar to that
in C. marulius. In live specimens, the margin of the
ocellus is orange; the margin appears white in
preserved specimens. Channa marulioides often has a
series of dark patches of scales, the posterior and
posterodorsal scales each margined by white, along the
Channa marulioides, caught and released from jungle of sides of the body, a character that is absent in
Perak State, Malaysia, January 2003. Photo courtesy of C. melanoptera (Kottelat and others, 1993; Lee and
Jean-Francois Helias, Fishing Adventures Thailand.
Ng, 1991, 1994) and C. marulius, and may disappear
with growth. The iris of the eye is orange or red as in
Size: To 65 cm (Lee and Ng, 1994). C. marulius.
Habitat preference: A riverine species (Kottelat, Commercial importance in the United States:
1994), also found in lakes, appearing to be an inland Rarely mentioned in aquarist-oriented websites. This
species (Lee and Ng, 1994). species is colorful and has perhaps been found periodi-
Temperature range: No specific information cally for sale in the aquarium fish trade. Not known
found. The native range is equatorial/tropical. from live-food fish markets.
Reproductive habits: No specific information
found. Likely a nest builder with adults guarding fertil- Commercial importance in native range: Ng
ized eggs and larvae. and Lim (1990) stated that this snakehead is sold in the
aquarium fish trade in Singapore, costing up to S$100
Characters: No patch of scales on gular region
per individual. This market likely precludes this species
of head. Dorsal fin rays 45-47; anal fin rays 30-31.
as available in live-food fish markets as a food species.
Lateral line scales 55-58; predorsal scales 13-15.
Scales between lateral line and anterior rays of dorsal Environmental concerns: Likely a thrust predator.
fin 3½. Lateral line curves downward abruptly at lateral This is an equatorial/tropical species that, if introduced,
line scales 17-20. Preopercular scales 5-7. No canines might establish only in areas with a similar climate.

100°E 120°E 140°E

China

Myanmar PACIFIC
20°N
OCEAN
Thailand Laos
Philippines
Vietnam

EXPLANATION Cambodia
DISTRIBUTION OF
Channa marulioides
Malaysia
Native range Malaysia
INDIAN Indonesia Equator
0°
OCEAN
0 500 MILES
Indonesia Indonesia
0 500 KILOMETERS
Scale is approximate

Channa marulioides

82
 Channa marulius (Hamilton, 1822)
Bullseye Snakehead

After Munro, 1955

After Hamilton, 1822; juvenile

Original description: Ophiocephalus marulius Hamilton, 1822:65, 367, pl. 17, fig. 19. An account of the fishes
found in the river Ganges and its branches. Edinburgh and London, i-vii + 1-405, pls. 1-39. Type locality: “Gangetic
provinces,” India. No types known.
Synonyms: Ophicephalus sowara Cuvier, 1831:426.
Ophicephalus grandinosus Cuvier, 1831:434.
Ophiocephalus leucopunctatus Sykes, 1839:158.
Ophiocephalus theophrasti Valenciennes, 1840:pl. 13, figs. 1-1a.
Ophicephalus pseudomarulius Günther, 1861:478.
Ophiocephalus aurolineatus Day, 1870:99.
Channa marulius ara Deraniyagala, 1945:24, pl. 24.
Common names: bullseye snakehead; giant snakehead; great snakehead; cobra snakehead; Indian snakehead; soal
(Pakistan); haal (Assam, India); sal, gajal (West Bengal, India); pumurl, bhor (Bijar, India); kubrah, sawal, dowlah
(Punjab, India); saal (Orissa, India); poomeenu, phoola-chapa, phool-mural (Andra Pradesh, India); aviri, puveral
(Tamil Nadu, India); chaeru-veraal, curuva, bral (Kerala, India); hoovina-murl, madinji, aviu (Karnataka, India); ara,
gangara, kalumaha (Sinhalese, Sri Lanka); iru viral (Tamil, Sri Lanka); ara (Sri Lanka); trey raws (Cambodia).

Smith (1945) noted that near Bankok, Thailand, the common name pla chon ngu hao is used for this species;
ngu hao means cobra. Talwar and Jhingran (1992) listed a common name of kubrah, used in Punjab, India. This
perhaps explains the use of “cobra snakehead” for this species in the U.S. aquarium fish trade.

83
 Native range: Pakistan (Kabul and Indus Rivers; Size: This species is reported to be the largest of
Mirza, 1999); many drainages of India, Sri Lanka, the family Channidae, reaching a length of 120-122 cm
Bangladesh, southern Nepal (Gandaki, Koshi, and (Bardach and others, 1972; Talwar and Jhingran, 1992).
Karnali River basins), Myanmar, Thailand, Mekong Talwar and Jhingran (1992) reported that it grows to
basin of Laos and Cambodia, and southern China (Day, 180 cm and a weight of 30 kg in Maharashtra State,
1877, Nichols, 1943; Mendis and Fernando, 1962; western India, noting that a length of 30 cm can be
Qureshi, 1965; Fernando and Indrassna, 1969; attained in 1 year. Rohan Pethiyagoda and Prachya
Pethiyagoda, 1991; Talwar and Jhingran, 1992; Musikasinthorn (personal commun., 2002) stated they
Rainboth, 1996; Kottelat, 2001a). Smith (1945) cited doubted that any snakehead would reach such a length
this species as “one of the rarest of the serpent-heads and were unaware of any specimens of that size.
found in Thailand.” It is not reported from Malaysia or Murugesan (1978) reported a maximum size of
Indonesia where it appears to be replaced by a 52.8 cm at 8 months from fish stocked at an average
somewhat lookalike species, Channa marulioides. length of 37 mm in a tank in Karnataka State, south-
Nichols (1943) recorded the species from the Yangtze western India, with lengths of 38.6 to 48 cm typical in
drainage, China. Maurice Kottelat and Tyson Roberts 1 year. He recorded growth of 2.5 to 4 mm/day for the
(personal commun. to J.D. Williams, 2001) indicated first 3 months and 0.8 to 1.3 mm/day thereafter, with an
that C. marulius, as currently recognized, is possibly a average growth rate of 7 cm/month. Johal and others
species complex. (1983) reported that bullseye snakeheads grow faster
Introduced range: Introduced into residential than Channa argus, C. punctata, or C. striata, but also
lakes and adjoining canals in Tamarac, Broward noted that growth decreases with increasing age, with
County, Florida, and established as a reproducing the greatest increase in weight occurring during the
population (Florida Fish and Wildlife Conservation second year. Their specimens were obtained from River
Commission, 2001; Howells and others, 2002; Paul L. Ghagger, Rajasthan State, northwestern India. Ahmad
Shafland, personal commun., 2002). and others (1990) also reported decreasing growth rate

Adult Channa marulius guarding young. Photographed April 23, 2003, in Tamarac, Broward County, Florida,
by Ianaré T. Sevi.

84
 Channa marulius, from Khao Laem Reservoir, Kanchanaburi Province, Thailand, June 2002. Photo courtesy
of Jean-Francois Helias, Fishing Adventures Thailand.

with increasing age in specimens from River Kali in Reproductive habits: Breder and Rosen (1966)
northern India. Wee (1982) cited C. marulius and summarized brood size from Chacko and Kuriyan
C. micropeltes as the two fastest growing snakeheads. (1947) as being about 500 young with parents guarding
them until they reached about 10 cm in length. Eggs are
Habitat preference: Lakes and rivers; deep,
pale red-yellow and 2 mm in diameter, hatch in 54 hrs
clear water with sand or rocky substrate (Talwar and
at 16-26 ºC and 30 hrs at 28-33 ºC. Parameswaran and
Jhingran, 1992); “rivers usually in the vicinity of mud
Murugesan (1976a) reported brood sizes of 357 to over
or fine sands” (Sen, 1985); “deep pools in rivers and
3,600 in swamps in Karnataka State, India. Talwar and
occasionally in lakes” (Pethiyagoda, 1991). Jhingran
Jhingran (1992) remarked that this species lays eggs in
(1984) noted presence of this species in swamps, tanks
a nest and guards them, and that breeding occurs
(=small reservoirs), and ponds, but that it also “prefers
through most of the year. Day (1875) supported this
deep, clear stretches of water with sandy or rocky
guarding habit by stating, “Colonel Puckle observes
bottom.” Rainboth (1996) listed preferred habitat as
‘that they are very savage, protecting their young with
“sluggish or standing water in canals, lakes, and
great boldness’.” Pethiyagoda (1991) noted that males
swamps from India to China, south to Thailand and
are territorial. In river basins of southern Nepal, it is
Cambodia.” He added that it is often found with
said to spawn from June until August (Shrestha, 1990).
submerged aquatic vegetation.
Parameswaran and Murugesan (1976a) indicated
Temperature range: No specific information in spawning occurs in all but the months of December and
literature. Native range of the species is from about January in Karnataka State, India, peaking during the
34º N to 7º N, indicating a species that can exist in rainy season. Jhingran (1984) stated that fecundity of
temperate to tropical conditions. Preliminary tempera- this species is “2,000-40,000 ova.” Mirza and Bhatti
ture testing on individuals from the established south- (1993) indicated a spawning period in Pakistan of April
eastern Florida population indicated the lower range to to June with parental protection of eggs and larvae for
be about 10 ºC (Paul L. Shafland, personal commun., 6 weeks. Sriramlu (1979) reported two spawning peri-
2002), suggesting that this population did not originate ods (May-June and November-December) for Channa
from northern reaches of the native range of this marulius in Andhra Pradesh, southeastern India, at
species. altitudes ranging from 548 to 670 m above sea level.

85
 A Mozambique tilapia (Oreochromis mossambicus), an introduced species
in Thailand, bitten in half by a giant snakehead (Channa micropeltes).
Photo courtesy of Jean-Francois Helias, Fishing Adventures Thailand.

Srivastava (1980) also reported two spawning peaks (1990) stated the diet of C. marulius in the River Kali,
(May and January) for this species from a lake at Gora- northern India, was more than 60 percent fishes and the
khpur, in Uttar Pradesh, northern India. This species is remainder crustaceans, gastropods, insects, and larval
one of three snakeheads known to spawn in the absence chironomids.
of vascular aquatic plants (Parameswaran and Muruge- Characters: No patch of scales on gular region
san, 1976b).
of head. Dorsal fin rays 45-55; anal fin rays 28-36;
Food preferences: Regarded as predacious pectoral rays 16-18; pelvic fin rays 6. Lateral line
(Jhingran, 1984; Talwar and Jhingran, 1992), especially scales 60-70; predorsal scales 16. Lateral line scales
on other fishes (Schmidt, 2001). The few aquarist- drop two rows between the 16th and 18th perforated
oriented websites that list this species warn that it scale. Scale rows between posterior margin of orbit and
cannot coexist in aquaria with other fishes once a preopercular angle 10. Scales on top of head moderate-
length of 25 cm is reached. Schmidt (2001) goes sized with a rosette of head scales between the orbits,
further to recommend that two individuals of the same with the frontal head scale in the center of the rosette;
species or mixed with others should not remain in the two scales between rosette and the basal head scale; 10
same aquarium after they reach 25 cm, concluding that scale rows between preopercular angle and posterior
"at that stage…they will establish a species aquarium border of orbit. Pectoral fin length about half head
in their own way." He described the species as a "thrust length. Mouth large, lower jaw with 7 to 18 canines
predator", noting that it is rarely found in the aquarium behind a single row of villiform teeth that widen to 5-6
trade and, if found, individuals are juveniles. rows at jaw symphysis. Teeth present on prevomer but
Dasgupta (2000) reported stomach contents of absent on palatines (Talwar and Jhingran, 1992), and
Channa marulius collected from several localities in Smith (1945) indicated that canine teeth were absent
West Bengal, India, as consisting primarily of fishes from both prevomer and palatines.
(40 percent), followed by crustaceans (30 percent), Talwar and Jhingran (1992) provided informa-
“macrophyte tissue” (15 percent), larval insects (10 tion on life colors for both young and adults, comment-
percent), and algae (5 percent). Ahmad and others ing that there is a “pale-edged ocellus” toward the

86
upperside of the caudal fin base that “fades with early 2002. In May 2002, law enforcement personnel of
growth,” but provided no information as to size when the California Department of Fish and Game confis-
that happens. Kottelat (2001a), however, stated that the cated a bullseye snakehead from an aquarium fish
ocellus occurs only in individuals less than 40 cm in dealer in the Los Angeles area (Richard Feeney,
standard length. Juveniles often with a series of dark personal commun. 2002).
blotches (usually five) bordered posteriorly and poster- Commercial importance in native range: This
odorsally by a series of white scales forming a white species is considered an important food fish in India,
margin to the blotches (Prachya Musikasinthorn, and it is a significant component of the freshwater fish-
personal commun., 2002). ery in Andhra Pradesh State (Talwar and Jhingran,
1992). Rao and Durve (1989) reported this species as
The only other species of Channa having an one of three snakeheads fished commercially in Lake
ocellus on the upper lobe of the caudal fin near its base Jaisamand, the oldest reservoir in India. Sriramlu
is C. marulioides. The geographic ranges of the two (1979) listed this species and Channa striata as the
species do not overlap. In contrast to C. marulius, most preferred fish in two regions of Andhra Pradesh,
C. marulioides has 55-58 lateral line scales, 13-15 India. Bardach and others (1972) noted that it was
predorsal scales, and 5-7 preopercular scales. The under culture in India and Pakistan in ponds and rice-
lateral line drops abruptly between the 17th and 20th fields, and irrigation wells that do not support other
scale in C. marulioides, and there is a two row drop of fishes. Wee (1982) listed the species as being reared in
the lateral line in C. marulius between the 16th to 18th monoculture ponds in India where it is fed tilapia.
scale. Juveniles of C. marulius may have a series of Nevertheless, Mirza and Bhatti (1993) stated that this
dark blotches along the sides, margined posteriorly and species is unsuitable for aquaculture in Pakistan
because it is piscivorous. Rainboth (1996) noted that it
posterodorsally by a series of white scales; late juve-
is “marketed fresh and sometimes alive” in Cambodia.
niles and adults of C. marulioides also have a series of
It is touted as an important sport species in Thailand
dark blotches on the sides, but the posterior and poster- (http://www.fishingasia.com).
odorsal scales of these blotches are dark, usually black, Environmental concerns: The native range of
and margined with white (Lee and Ng, 1991). this species indicates a temperate to tropical species
Commercial importance in the United States: that, if introduced, has the potential to establish into
Other than occasional mention on aquarist-oriented southern states of the contiguous United States as well
websites and Schmidt's (2001) statement that "giant as Hawaii. This species is regarded as a predator,
snakehead" are rare in the aquarium trade, we found particularly on other fishes (Schmidt, 2001). That it has
become established as a reproducing population in
that it was being marketed under the name cobra snake-
southeastern Florida indicates the likelihood that it
head as an aquarium fish in the U.S. Aquarist-oriented
could establish elsewhere if introduced into areas with
chat rooms on the Internet suggest it may be second in a suitable temperature regime.
popularity to Channa micropeltes. Its introduction into Comments: The diploid number of chromo-
southeastern Florida may have resulted from an inten- somes of Channa marulius is 44 (Donsakul and
tional release of aquarium fish by hobbyists. Neverthe- Magtoon, 1991).
less, we recently learned of its availability in live-food
fish markets in New York City (Leo Smith, personal See map on following page
commun., 2002) and, therefore, cannot rule out the
live-food fish trade as having been the source of this
introduction. Nevertheless, its popularity as a game
species in Thailand (http://www.fishingasia.com) may
have prompted someone to illegally introduce this fish
for sport purposes. Using a special permit from the
FFWCC to import restricted and prohibited aquatic
species, the USGS purchased a young bullseye snake-
head from an aquarium fish dealer in Rhode Island in

87
 60°E 80°E 100°E 120°E
North
40°N
Korea

Afghan-
Iran istan China South
Nepal Korea
Bhutan
Pakistan

Saudi India
Arabia Myanmar
20°N

Laos
Bangladesh Thailand
Vietnam

Cambodia
Thailand
Malaysia

Equator
0°

INDIAN OCEAN Indonesia

85°W 80°W
0 1,000 MILES EXPLANATION ALABAMA

AT L
GEORGIA
DISTRIBUTION OF
Channa marulius FLORIDA

AN
0 1,000 KILOMETERS 30°N

TIC
Scale is approximate Native range
GUL
Introduced range F
OF

OC
M

EAN
E
X
I
C
O

0 100 MILES

25°N 0 100 KILOMETERS

Channa marulius

88
 Channa melanopterus (Bleeker, 1855)
Blackfinned Snakehead

After Weber and de Beaufort, 1922; image reversed from original

After Bleeker, 1878

Original description: Ophicephalus melanopterus Bleeker, 1855:420. Negende bijdrage tot de kennis der
ichthylogische fauna van Borneo. Zoetwaterfischen van Pontianak en Banjdermasin. Natuurkd. Tijdschr. Neder.
Indië 9:415-430. Type locality: Kapuas River, Pontianak, Borneo, Indonesia. Location of holotype reported as
unknown, but located by Musikasinthorn (2000) as RMNH 6416.
Synonyms: No known synonyms (Ng and Lim, 1990). Often confused with Channa marulius (Ng and Lim, 1990;
P. Musikasinthorn, personal commun., 2002), and C. marulioides (Lee and Ng, 1994).
Common name: blackfinned snakehead.

Native range:Kapuas River basin of Kalimantan Temperature range: No specific information.

(western Borneo) and possibly the southern tip of Native range is equatorial to about 3º S.
Sumatra (Roberts, 1989; Lee and Ng, 1994). These Reproductive habits: No information located,
latter authors stated that records of this species from but likely a nest builder that provides parental protec-
central Sumatra are misidentifications of Channa tion of eggs and fry.
marulioides. Feeding habits: No information found. Likely a
Introduced range: No introductions known. thrust predator as other snakeheads.
Characters: No patch of scales on gular region
Size: To at least 65 cm (Lee and Ng, 1994).
of head. Dorsal fin rays 44-48; anal fin rays 28-32.
Habitat preference: No specific information, Lateral line scales 54-57; predorsal scales 13-15.
but appears to be a riverine species. Lateral line curves downward at lateral line scales 16-18.

89
Preopercular scales 5-8. Unfortunately, these characters Commercial importance in the United States:
are about equal to those of Channa marulioides. Lee Rarely mentioned in aquarist-oriented websites and
and Ng (1994) stated that the only way to separate probably has been rarely, if ever, available for sale in
these two species is by coloration. Channa marulioides the aquarium fish trade. Unknown from live-food fish
possesses an ocellated spot on the upper part of the markets.
caudal fin base, similar to that in C. marulius, and
Commercial importance in native range: No
C. melanopterus lacks this ocellus. There are also dif-
information found. Considering its moderate size, it is
ferences in coloration of live individuals (Lee and Ng,
probably available at local live-food fish markets.
1994). Channa marulioides also differs from C. melan-
opterus in that the former often (but not always) pos- Environmental concerns: Likely a thrust preda-
sesses several patches of dark scales rimmed by white tor. This is a strictly tropical species that could only
margins along the sides of the body (Lee and Ng, establish in extreme southern Florida, Hawaii, or warm
1994). thermal springs and their outflows if introduced.

100°E 120°E 140°E

China

Myanmar PACIFIC
20°N
OCEAN
Thailand Laos
Philippines
Vietnam

EXPLANATION Cambodia
DISTRIBUTION OF
Channa melanopterus Malaysia
Native range Malaysia
Possible native INDIAN Indonesia Equator
range 0°
OCEAN
0 500 MILES Indonesia
Indonesia
0 500 KILOMETERS
Scale is approximate

Channa melanopterus

90
 Channa melasoma (Bleeker, 1851)
Black Snakehead

Upper image: adult; photo by K.K.P. Lim. Lower image: subadult.

Reprinted with permission from P.K.L. Ng from Lee, P.G., and P.K.L. Ng. 1991.
The snakehead fishes of the Indo-Malayan region. Nature Malaysiana 16(4):112-129.

After Bleeker, 1878

Original description: Ophicephalus melasoma Bleeker, 1851:424. Vijfde bijdrage tot de kennis der ichthyologische
fauna van Borneo, met beschrijving van eeinige nieuwe soorten van zoetwatervisschen. Natuurkd. Tijdschr. Neder.
Indië 2:415-442. Type locality: Sambas, western Borneo. Whereabouts of holotype unknown.
Synonyms: Ophicephalus rhodotaenia Bleeker, 1851:425.
Ophicephalus mystax Bleeker, 1853:188.
Ophicephalus melanosoma Bleeker, 1856:214.
Ophiocephalus melanosoma Günther, 1861:473.
Ophiocephalus baramensis Steindachner, 1901:3435, fide Roberts, 1989, Ng and Lim, 1990, and
Rainboth, 1996. Species revalidated by Ng and others, 1996.
Note: Frequently misspelled melanosoma (Ng and Lim, 1990).
Common names: black snakehead; ikan bakak (Malay).

91
 Native range: Chao Phraya River, near Bangkok, Reproductive habits: A nest builder like other
Thailand (misidentification?); Mekong River in Cambo- channids; only one parent guards eggs and young (Ng
dia (Rainboth, 1996; misidentification?); rivers of south- and Lim, 1990).
eastern Sumatra; rivers of western Kalimantan,
Feeding habits: A nocturnal thrust predator (Ng
particularly the Kapuas basin (southern Borneo);
and Lim, 1990; Lee and Ng, 1991, 1994). Feeds on other
Bangka and Belitung (Billiton) islands; Palawan Archi-
fishes, lizards, crabs, insects and insect larvae (Inger and
pelago, Philippines (Inger and Kong, 1962; Roberts,
Kong, 1962; Ng and Lim, 1990; Rainboth, 1996).
1989; Ng and Lim, 1990; Lee and Ng, 1991). Bean and
Weed (1912) reported the species from Java, later chal- Characters: No patch of scales on gular region.
lenged by Weber and de Beaufort (1922), and there is no Dorsal fin rays 37-40; anal fin rays 22-25; pectoral rays
evidence that it occurs there (Ng and Lim, 1990). Smith 14-17. Lateral line scales 50-54; transverse scale rows
(1945) reported this species as rare in Thailand, and Lee from lateral line to dorsal fin origin 4-5½; transverse
and Ng (1994) stated that because of its preference for scale rows from lateral line to anal fin origin 7½. Color
acidic waters, the species is more common toward the in preservative dark brown above, pale below, with fins
southern part of the Malay Peninsula. It is present but dusky or dark and caudal fin usually barred. Many lateral
apparently rare in the North Selangor Peat Swamp Forest scales with dark central spots (Inger and Chin, 1962).
of the Malaysia Peninsula (Ng and others, 1992). Peter See account of Channa baramensis for characters that
Ng (personal commun., 2003) reported occurrence of separate C. melasoma from this closely related species.
this species in the Golok area of southern Thailand. Commercial importance in the United States:
Records of this species from northern Borneo (Sarawak, Rarely mentioned on aquarist-oriented websites. Not
Brunei, and Sabah) are misidentifications of the endemic known to have been imported as a live-food fish
Channa baramensis (Ng and others, 1996). species.
Introduced range: Uncertain. Presence of this
Commercial importance in native range:
species on Palawan (Kottelat, 1985; Roberts, 1989) Probably nil. This species is primarily nocturnal and
may have resulted from an introduction. The same may reported to be difficult to capture (Lee and Ng, 1991).
be true in the Chao Phraya River, Thailand, and Rainboth (1996) noted that it is not found in markets in
Mekong River, Cambodia, unless misidentified. Cambodia.
Size: To 35 cm (Inger and Kong, 1962). Environmental concerns: This species is a
Habitat preference: Prefers shaded, clear water nocturnal thrust predator, known to feed on other fishes
forest streams with mud bottom and leaf litter, but is also as well as lizards, crabs, and insects (Lee and Ng, 1991).
found in turbid waters.
Reported to burrow into mud 100°E 120°E 140°E
bottom when disturbed (Lee China
and Ng, 1991). Seems to
Myanmar PACIFIC
prefer waters of pH 5-5.3 20°N
(Lee and Ng, 1994). Also OCEAN
Thailand Laos
occurs in middle parts of Philippines
rivers. Capable of locomotion
on land (Lee and Ng, 1994). EXPLANATION Vietnam

Rainboth (1996) stated that it DISTRIBUTION OF

Channa melasoma Cambodia
prefers “sluggish or standing Native range
waters” in the Mekong River Possible Malaysia
basin. misidentification Malaysia
INDIAN Indonesia Equator
Temperature range: 0°
OCEAN
No specific information, but
native range of Channa 0 500 MILES Indonesia
melasoma is tropical Indonesia

(Borneo) to subtropical 0 500 KILOMETERS

Scale is approximate
(Thailand).
Channa melasoma

92
 Channa micropeltes (Cuvier, 1831)
Giant Snakehead

Upper image: young adult; photo by P.R. Sweet. Lower image: young juveniles. Reprinted with permission
from P.K.L. Ng from: Lee, P.G., and P.K.L. Ng. 1991. The snakehead fishes of the Indo-Malayan region.
Nature Malaysiana 16(4):112-129

After Bleeker, 1878; juvenile

Original description: Ophicephalus micropeltes Cuvier in Cuvier and Valenciennes, 1831:427. Histoire naturelle
des poissons 7:i-xxix +1-537, pls. 170-208. Type locality: Java, Indonesia. Syntypes: RMNH D2318; possible
syntypes: RMNH D1131, RMNH D1132.

93
Synonyms: Ophiocephalus serpentinus Cuvier, 1831:429.
Ophicephalus bivittatus Bleeker, 1845:519.
Ophiocephalus stevensii Bleeker, 1853:444.
Ophiocephalus diplogramma Day, 1865a:36.
Ophiocephalus diplogramme [sic] Day, 1865b:147.
Ophiocephalus studeri Volz, 1903:555.
Common names: giant snakehead; red snakehead, redline snakehead; Malabar snakehead; Mala; ikan toman,
toman (Malaysia); pla chado (Thailand), pla melang pu (Bangkok, Thailand); trey diep (juvenile) and trey chhdaur
(adult; Cambodia); toman, anak toman, gabus tobang (Kalimantan). The name red or redline snakehead is commonly
applied to juveniles of this species in the aquarium fish trade.

Native range:This snakehead has a markedly We believe that the description of Ophiocephalus
disjunctive distribution. Rivers of the Malabar (=Channa) diplogramma was based on a juvenile
(southwestern) Coast of India (restricted to Kerala C. micropeltes and that its presence in Cochin, Kerala
State, southwestern India fide Roberts, 1989; Talwar State, India, was the result of one or more introductions
and Jhingran, 1992); Myanmar (?); Thailand; Mekong from southeastern Asia that occurred prior to the mid-
basin of Laos; Vietnam; Malaysia; southeastern 1800s. Cochin has been a major trading port for several
Sumatra; Kalimantan, particularly the Kapuas basin
centuries. Moving an airbreathing snakehead, even on
(southwestern Borneo; Roberts, 1989; Rainboth,
sailing ships from its native range to a new locale some
1996); Bangka and Belitung (Billiton) islands; northern
Java (Day, 1877; Mohsin and Ambak, 1983; Roberts, 2,500 km or greater distance away, would have been an
1989; Lee and Ng, 1991, 1994; Kottelat, 1985, 2001a). easy task. The progeny from this introduction, resulting
Its presence in Myanmar (Burma) is questionable. from a small founder population, would be expected to
Ismail (1989) stated that it is “quite common in exhibit some small degree of character divergence
peninsular Malaysia.” when compared to the source population within its
We found it most puzzling that a species native range.
described from southeastern Asia (Java) could have a Introduced range: This species was reported
disjunctive distribution between a reported range in from Maine, Massachusetts, and Rhode Island, without
southwestern India (Kerala State) and Thailand, its evidence of reproduction (Courtenay and others, 1984;
westernmost confirmed range in southeastern Asia, a Fuller and others, 1999). A review of photographs of
distance of about 2,500 km. Such a distribution gap snakeheads captured from open waters of Maryland,
indicates that the form in Kerala State, India, is either a
undertaken by Maryland Department of Natural
different species or perhaps a very early introduction
Resources personnel during Summer 2002, revealed
from its native range. We believe the latter interpreta-
two specimens of giant snakeheads that had been
tion is correct.
caught on separate occasions in different drainages
Day (1865a) described Ophiocephalus diplo- during the two previous years. On September 6, 2002, a
gramma based on one specimen, about 42 mm in specimen about 56 cm long, was found struggling near
length, collected in October 1863 near the port city of
the surface in shallow water of the Inner Harbor, Balti-
Cochin in southwestern India. Day (1865b) provided a
more, Maryland, and captured by dipnet. All captures
species account for O. diplogramme [sic], his entry
based on the same juvenile specimen collected in 1863 of this species appear to be of released pet fish. Two
near the “mouth of the Cochin River.” The characters specimens, about 46 cm in length, were confiscated
he provided in that original description closely match from an aquarium fish dealer in Los Angeles, Califor-
those of Channa micropeltes, and his description of the nia, in 2000 (Camm Swift, personal commun., 2002).
life coloration appears to be that of a juvenile giant California is one of fourteen states where snakeheads
snakehead. were prohibited before summer 2002.

94
 Size: To 1 m and weight of over 20 kg (Roberts, consumes in its natural habitat. Ng and Lim (1990)
1989; Lee and Ng, 1991; Talwar and Jhingran, 1992). described the enlarged canine teeth of C. micropeltes as
Peter Ng (personal commun., 2003) noted that this being knifelike, “with two cutting edges in cross-
snakehead is known to reach a length of 1.5 m. Wee section,” the edges arranged perpendicular to the body
(1982) cited this species and Channa marulius as the axis. This allows shearing of prey.
two fastest growing snakeheads.
Beeckman and De Bont (1985) reported that the
Habitat preference: Lakes, rivers, canals, and digestive system of the giant snakehead is relatively
reservoirs (Mohsin and Ambak, 1983; Lee and Ng, short, that young fed on crustaceans and adults are
1991). Kottelat (1998) stated a preference for “deep piscivorous in the Nam Ngum reservoir, Mekong basin,
water bodies.” This species is nearly incapable of over- in northern Laos. They noted parasites in stomachs of
land movements (Ng and Lim, 1990) except for the 60 percent of all specimens captured with even more
young (Peter Ng, personal commun., 2002), but he has found in the intestines. Parasitism apparently begins
observed large individuals attempting to move on dry when this species becomes piscivorous.
land. Rainboth (1996) noted a preference for “standing
Cowx (1998) commented that escapes of
or slowly flowing waters.”
Chinese carps, tilapias, and oscar (Astronotus ocella-
Temperature range: No specific information, tus) from cage culture in Chenderoh Reservoir, Perak,
but its native range is between 20º N to about 7º S, Malaysia, failed to establish breeding populations. He
indicating a subtropical/tropical species. attributed this failure to predation by giant snakeheads.
Reproductive habits: Like other channids, Characters: A patch of scales present in the
Channa micropeltes clear a circular area of vegetation, gular area. Head depressed, somewhat pointed, and
spawn, and the pelagic eggs rise to the surface where flattened above. Mouth large, oblique, maxillary reach-
they are fiercely guarded by the parents. Guarding ing beyond posterior border of the eye. Lower jaw with
continues after hatching (Lee and Ng, 1991), probably several canine-like teeth behind a single row of villi-
until young become demersal. form teeth, the latter expanding to about 5 rows at the
jaw symphysis; large canine teeth on prevomer and
Feeding habits: Primarily a daytime feeder (Ng
palatines. Scales on top of head small; 16-17 scale rows
and Lim, 1990). Accounts of this species almost invari-
between preopercular angle and posterior border of
ably describe it as a vicious predator on other fishes.
orbit; predorsal scales 22; 95-110 scales in longitudinal
Adult and perhaps subadult Channa micropeltes feed
series, although Ismail (1989) reported 82-91 for speci-
in packs, usually in midwaters or near the surface.
mens from peninsular Malaysia. This species obviously
Parents guard their eggs and young, and are reported to
has smaller scales than other large Channa, hence the
have attacked humans that approached a nest (Smith,
species name micropeltes (=small scale). Dorsal fin
1945; Lee and Ng, 1991). Kottelat and others (1993)
rays 43-46; anal fin rays 27-30; pectoral rays 15; pelvic
stated that anglers and swimmers who got too close to
rays 6; caudal fin rounded. Pelvic fin about 50 percent
young were attacked, some seriously wounded, and
of pectoral fin length (Talwar and Jhingran, 1992;
that there have been fatalities. The report of fatalities
Musikasinthorn and Taki, 2001).
was from local fisheries officials (Maurice Kottelat,
personal commun., 2003). Peter Ng (personal Commercial importance in the United States:
commun., 2002) commented that he knew of one This species frequently appears on aquarist-oriented
instance where a man was nearly castrated by an websites. Although several sites warn that Channa
attacking giant snakehead. Prey includes other fishes, micropeltes is only suitable for aquarium purposes as
frogs, and birds (Lee and Ng, 1994). Lee and Ng juveniles, require substantial amounts of animal food,
(1991) commented that authorities at the Singapore and should not be kept with other fishes, it has been
Botanic Gardens planned to remove C. micropeltes available in aquarium fish retail stores in states where
from ponds at that facility because this snakehead was snakeheads are not prohibited. One retailer who sells
feeding on cygnets. Ng and Lim (1990) referred to this snakeheads via the Internet reported that he refused to
species as the “most ravenous” of snakeheads, and sell this species to individual hobbyists (Ken Arnold,
they, Mohsin and Ambak (1983), and Roberts (1989) personal commun., 2002). Releases of this species into
noted that it is known to kill more fishes than it waters of Maine, Maryland, Massachusetts, and Rhode

95
Island were likely made to dispose of fish that grew second most important fish group under culture
too large for aquaria. We also know of two instances, (Pantulu, 1976; Wee, 1982). FAO (1994) listed a
one in 2001 and another in 2002, when juvenile giant production of 1,490 metric tons of this species in
snakeheads were confiscated from pet shops in southern Thailand in 1992, up from 386 tons in 1986. Rainboth
California. Because it is an esteemed food fish in (1996) noted that it is sold fresh and sometimes as
southeastern Asia, it has possibly been available live fish in Cambodian markets, and that it is cultured
periodically in live-food fish markets. in cages.
Commercial importance in native range: In the Kapuas River of western Kalimantan,
Talwar and Jhingran (1992) remarked that this species there is a hook fishery for Channa micropeltes,
is only of minor interest to fisheries in Kerala State, particularly within the Danau Sentarum Wildlife
India. Conversely, Lee and Ng (1991) stated that it is a Reserve. About 40 percent of the fishes caught with
highly prized food fish in Malaysia, and grown in cage large (sizes 5-8) hooks and 60 percent taken with
culture where it is fed tilapia. It is also grown in cage medium (sizes 9-11) hooks were giant snakehead.
culture in the lower Mekong basin of southern Vietnam In recent years, cage culture has developed for this
where it and Channa striata collectively comprise the species in the Reserve where large numbers of

Mark Sabaj (left) purchasing specimens of giant snakehead for the Academy of Natural Sciences, Philadelphia,
caught from Khao Laem Reservoir, an impoundment of Mae Nam Khwae Noi River, in a market in Thong Pha
Phum, Thailand, in 2001. Photo by Mike Hardman, Los Angeles County Museum of Natural History.

96
juveniles are caught with cast nets, raised in wooden Environmental concerns: Channa
cages, and fed other wild caught fish until they reach micropeltes has a reputation of being the most
market weight of 0.8-1.5 kg. This earns about predacious of all snakeheads, known to kill more
$700,000 for the communities and represents one-third fishes than it consumes. Its greatest threat in the U.S.
of fish-related income in the Reserve. Recently recog- would be in southern Florida and Hawaii where this
nized problems with this activity include fear of dimin- subtropical/tropical species would likely establish if
ished populations of giant snakehead due to removal of introduced.
young from the wild for culture purposes and the Comments: The diploid number of chromo-
estimated 4,000 tons of river-caught fish needed somes of Channa micropeltes is 44 (Donsakul and
annually to feed them (Dudley, 2000). Magtoon, 1991).
See map on following page

Channa micropeltes, caught by angler John Oatley; Khao Laem Reservoir, Kanchanaburi Province,
Thailand, 2002. Photo courtesy of Jean-Francois Helias, Fishing Adventures Thailand.

97
 60°E 80°E 100°E 120°E
North
40°N
Korea

Afghan-
Iran istan China South
Nepal Korea
Bhutan
Pakistan

Saudi India
Arabia Myanmar
20°N

Laos
Bangladesh Thailand
Vietnam

Cambodia
Thailand
Malaysia

Equator
0°

INDIAN OCEAN Indonesia

EXPLANATION 0 1,000 MILES

DISTRIBUTION OF
Channa micropeltes 0 1,000 KILOMETERS
Native range Scale is approximate
Possible native range
Introduced range

Channa micropeltes

98
 Channa nox Zhang, Musikasinthorn, and Watanabe, 2002
Night Snakehead

Purchased from a live-food fish market at Mong Kok, Hong Kong. Photograph by Heok Hui Tan.

Original description: Channa nox Zhang, Musikasinthorn, and Watanabe, 2002:140-146. Channa nox, a new
channid fish lacking a pelvic fin from Guangxi, China. Ichthyological Research 49(2):140-146. Type locality:
Nanliu River basin, vicinity of Hepu, Guangxi Province, China. Holotype: IOZCAS (Institute of Zoology, Chinese
Academy of Sciences) 70028. Paratypes: IOZCAS 69848; IOZCAS 69849; IOZCAS 70029; IOZCAS 70039; and
IOZCAS 70042.
Synonyms: No synonyms.
Common names: None in English. We propose night snakehead based on the dark coloration of the fish.

Native range: Southern China, near Hepu, Characters: Gular region of head with one to
Guangzi Province, specifically the lower Nanlui Jiang two scales on either side of underside of lower jaw.
River, where its range overlaps that of its nearest Pelvic fins absent. Head small, rounded. Dorsal fin rays
congener, Channa asiatica. Peter Ng (personal 47-51; anal fin rays 31-33. Lateral line scales 55-63;
commun., 2003) found this snakehead in live-food fish 5.5-6.5 scales above lateral line; check scales 9-13.
markets in Guangzhou, China, and Hong Kong in Total vertebrae 53-55. Large ocellus, black with white
July 2000. rim, on caudal peduncle. This species is most closely
Introduced range: No introductions known. related to Channa asiatica which also lacks pelvic fins
Size: Reported to almost 20 cm (Zhang and and has 49-53 vertebrae, 41-47 dorsal fin rays and 28-
others, 2002), but probably attains a larger size. 32 anal fin rays. Channa nox also differs in having a
Habitat preference: Zhang and others (2002) shorter predorsal length (26.9-28.4 percent) and snout
reported the species from humid rainforest conditions. length (3.6-5.1 percent). Live coloration also differs
Appears to be a riverine species. from C. asiatica in that C. nox is dark on the upper half
Temperature range: No information available. of the body and the ocellus on the caudal peduncle
Nevertheless, distribution within native range (21-32º N) lacks a white rim.
indicates a warm temperate to subtropical species. Commercial importance in the United
Reproductive habits: No information available States: None.
concerning reproduction in natural habitats, but likely a Commercial importance in native range:
nest builder like most snakeheads. There are indica- Available in local markets in Hepu, Guangxi Province,
tions that the male of the most closely related species, southern China (Zhang and others, 2002). Also
Channa asiatica is perhaps a mouthbrooder. Whether reported as available in markets in Ghangzhou and
such behavior applies to C. nox is unknown. Hong Kong (Peter Ng, personal commun., 2003).
Feeding habits: No information available, but Environmental concerns: Likely a thrust
likely a thrust predator as is its closest relative, Channa predator on other fishes and invertebrates like other
asiatica. snakeheads.

99
 100°E 120°E 140°E
China

Myanmar PACIFIC
20°N
OCEAN
Laos
Thailand Philippines
Vietnam

EXPLANATION Cambodia
DISTRIBUTION OF
Channa nox Malaysia
Malaysia
Native range
INDIAN Indonesia Equator
0°
OCEAN
0 500 MILES
Indonesia Indonesia
0 500 KILOMETERS
Scale is approximate

Channa nox

100
 Channa orientalis Schneider, 1801
Ceylon Snakehead
Type species of Channa Scopoli, 1777

Reprinted with permission from Rohan Pethiyagoda from Pethiyagoda, R. 1991. Freshwater fishes of
Sri Lanka. Wildlife Heritage Trust of Sri Lanka, Colombo, Sri Lanka.

After Munro, 1955; female

Original description: Channa orientalis Schneider, in Bloch and Schneider, 1801:496, pl. 90. M.E. Blochii,
Systema Ichthyologiae iconibus cx illustratum. Post obitum auctoris opus inchoatum absoluit, correxit, interpolavit
Jo.Gottlob Schneider, saxo. Sumtibut Austoris Impressum et Bigliopolio Sanderiano Commissum, Beroloni. i-lx +
1-584, pls. 1-110. Type locality: “Habitat in India orientale” (=east India). No types known (Paepke, 1993). Paepke
(1993) noted that Schneider’s description was based on an illustration in Gronovius (1763).
Synonyms: There has been much confusion regarding the taxonomy of the Ceylon snakehead. Many have considered
this species to be part of what we term the Channa gachua complex as reflected in the following synonymy, all of which
are synonyms of C. gachua and are invalid for C. orientalis (Ralf Britz, personal commun., 2003).
Ophiocephalus auranticus Hamilton, 1822:69, 368, pl. 23, fig. 22.
Ophicephalus marginatus Cuvier, 1829:230, fide Menon, 1999:275.
Ophicephalus coramota Cuvier, 1831:414, fide Menon, 1999:275.
Ophicephalus fuscus Cuvier, 1831:414, fide Menon, 1999:275.
Ophicephalus limbatus Cuvier, 1831, no p., pl. 201, fide Menon, 1999:275.

101
 Ophicephalus marginatus Cuvier, 1831, no p., pl. 201, fide Menon, 1999:275.
Ophicephalus montanus McClelland & Griffith, 1842:583, fide Menon, 1999:275.
Philypnoides surakartensis Bleeker, 1849:19, fide Menon, 1999:275.
Ophiocephalus apus Canestrini, 1861:77, pl. 4, fig. 7.
Ophiocephalus kelaartii Günther, 1861:472, fide Talwar and Jhingran, 1991:1019, and Menon, 1999:275.
(?)Channa burmanica Chaudhuri, 1919:284, pl. 22, fig. 4.
Ophiocephalus gachua kelaarti Günther: Munro, 1955:100.
Note: Although Channa burmanica is listed as a synonym, some authors recognize this species as valid
(Peter Ng, personal commun., in Vierke, 1991b; Ralf Britz, personal commun., 2003).

Common names: Ceylon snakehead; Asiatic the C. gachua complex. Pethiyagoda (1991) noted a
snakehead; smooth breasted snakehead (Munro, 1955; possible introduction in the Mahaweli basin of Sri
Pethiyagoda, 1991); green snakehead (Ettrich, 1989); Lanka.
kola kanaya (Pethiyagoda, 1991). Size: Sometimes cited as the smallest species of
Native range: Channa orientalis is endemic to snakehead (Ismail, 1989; Talwar and Jhingran, 1992), a
southwestern Sri Lanka (Pethiyagoda, 1991; Ralf Britz, comment often also used for Channa gachua. Pethiya-
personal commun., 2003), specifically to the “wet goda (1991) noted that this snakehead typically does
zone” and “lower south western hills” (Pethiyagoda, not exceed 10 cm and, therefore, is significantly
1991). Mendis (1954) included both C. orientalis and smaller than C. gachua.
Ophicephalus gachua from Sri Lanka, commenting
Habitat preference: Deraniyagala (1929) and
that the former may or may not possess pelvic fins. The
Munro (1955) cited “clean freshwater pools close to
Ceylon snakehead, however, lacks pelvic fins (Deraniy-
agala, 1929; Pethiyagoda, 1991; Ralf Britz, personal streams” as the preferred habitat. Pethiyagoda (1991)
commun., 2003). Talwar and Jhingran (1992) noted stated that it occurs in “shaded, clear, flowing water
that C. orientalis lacking pelvic fins occurs in Sri with a silt or gravel substrate” and “shallow rivulets
Lanka, Myanmar, and Java; nevertheless, reports of this barely deeper than its own body.” He also predicted
fish from Myanmar and Java refer to members of the pollution and destruction of rainforest habitat in Sri
C. gachua complex. DeWitt (1960) considered absence Lanka would likely negatively affect populations of
of pelvic fins in snakeheads as an anomalous character; this species.
this conclusion, however, is not supported by Temperature range: No specific information
C. orientalis or other snakeheads that lack pelvics found. Because the species is endemic to Sri Lanka
(C. asiatica, C. bleheri, C. burmanica, apparently (unlikely), it is strictly tropical.
some members of the C. gachua complex, and C. nox).
Reproductive habits: Ettrich (1989) reported
Munro (1955) correctly identified the snakehead this species as a mouthbrooder. He described the basic
species lacking pelvic fins in Sri Lanka as Channa body color as brown, remarking that the flanks of
orientalis and treated the taxon with pelvics as males sometimes are dove gray with a violet cast,
C. gachua kelaarti. Lim and others (1990) noted that becoming paler ventrally. The dorsal and caudal fins of
Myers and Shapovalov (1932) synonymized C. gachua males are “sky-blue,” black and orange, with a blue
with C. orientalis, but suggested these species are sepa- anal fin margined in black and white with black rays,
rate with the latter lacking pelvic fins. Because and the eye red. Reproductively active females change
C. orientalis is endemic to Sri Lanka, records from from brown to shades of blue. During spawning, the
southern India and elsewhere are erroneous. male wraps itself around the female near the surface,
Introduced range: No introductions known, after which the male broods the eggs in its oral cavity,
although its presence in the Greater Sunda Islands typically remaining in an upper corner of an aquarium.
(Kottelat, 1985) would represent an introduction if the Fry remain in the male parent’s mouth until able to
species is indeed Channa orientalis. More than likely, survive on their own. Females sometimes retrieve stray
however, Kottelat’s (1985) reference is to a species of fry into their oral cavities, returning the young to the

102
parental care of the male. Fry are never ejected via the preopercle 5-6. Lateral line scales 36-42. Anal fin rays
mouth but rather leave via the gill openings. This is 20-22; dorsal fin rays 30-34. Pectoral fins with 13-15
somewhat similar to behavior observed in Channa rays. Ocellated spot at posterior end of dorsal fin,
bleheri (Vierke, 1991b). Both parents provide parental above caudal peduncle, in juveniles and females.
care with the female defending territory. It is unknown Lower jaw with 10-20 canines behind single row of
if the Ceylon snakehead is a mouthbrooder only in villiform teeth, the latter expanding to about 7 rows at
aquaria (Ettrich, 1989). jaw symphysis; prevomer and palatines with canine-
Ettrich (1989) assumed that there were two like teeth.
forms of Channa orientalis in Sri Lanka, one with and Commercial importance in the United States:
one without pelvic fins, but remarked that there were Sometimes listed on aquarist-oriented websites, but
no hybrids known between these forms. The one with often misidentified.
pelvic fins, however, is part of the C. gachua complex Commercial importance in native range:
(Deraniyagala, 1929; Pethiyagoda, 1991; Ralf Britz, Pethiyagoda (1991) stated, “Small numbers are used by
personal commun., 2003). the aquarium fish export trade.”
Feeding habits: Moyle and Senanayake (1984) Environmental concerns: Perhaps a thrust
determined that most gut contents consisted of terres- predator like other snakeheads. Because this species is
trial insects, the remainder comprised of trichopterans endemic to Sri Lanka, its ability to establish in North
and a few fishes. Pethiyagoda (1991) noted that in America, if introduced, would be limited to subtropical
aquaria, chopped steak is readily accepted by this areas (extreme southern Florida, Hawaii) or warm
snakehead. thermal springs.
Characters: Gular part of head without patch of
scales. Pelvic fins absent. Predorsal scales 6-7; scales
from posterior border of orbit to posterior edge of See map on following page

103
 80°E 100°E
Nepal
Bhutan China

India
EXPLANATION Myanmar Laos
DISTRIBUTION OF 20°N
Channa orientalis Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate Sri
Lanka Malaysia

Equator Indonesia
0°

Channa orientalis

104
 Channa panaw Musikasinthorn, 1998
Panaw Snakehead

Upper image: holotype, KUMF 3050, 151.7 mm standard length. Lower image: ventral view of head showing
enlarged scales on lower jaw; paratype, KUMF 3060. Reprinted with permission from Prachya Musikasinthorn,
author, and Tomoki Sunobe, Secretary of the Ichthyological Society of Japan, from: Musikasinthorn, Prachya.
1998. Channa panaw, a new channid fish from the Irrawaddy and Sittang River basins, Myanmar. Ichthyol. Res.
45(4):355-362.

Original description: Channa panaw Musikasinthorn, 1998:356. Channa panaw, a new channid fish from the
Irrawaddy and Sitang River basins, Myanmar. Ichthyological Research 45(4):355-362, 7 figs. Type locality: Yangon
fish market, Yangon, Myanmar. Holotype: KUMF 3050. Paratypes: KUMF 3051; KUMF 3060; KUMF 3061;
KUMF 3062; NSMT-P 36121; NSMT-P 36129; KUMF 3052; KUMF 3053; KUMF 3054; NRM 27421; and ANSP
77016.
Synonyms: No synonyms.
Common names: panaw snakehead; nga panaw (Myanmar).

Native range: Ayeyarwaddy (=Irrawaddy) and Feeding habits: No information. Nevertheless,

Sittang River basins, Myanmar (Musikasinthorn, Musikasinthorn (1998) stated this species is most
1998). closely related to Channa punctata, which suggests
Introduced range: No introductions known. adults feed primarily on other fishes and insects.
Size: No specific information in literature, but Characters: Gular part of head without patch of
known to grow to at least 17 cm (Musikasinthorn, scales. One large scale on either underside of lower
1998). jaw, rarely two on one side of jaw. Lateral line scales
39-41; predorsal scales 14-17. Dorsal fin rays 32-35;
Habitat preference: No specific information,
anal fin rays 32-35; pectoral rays 17-20. Pelvic fin
but appears to prefer rivers.
length always more than 50 percent of pectoral fin
Temperature range: No specific information. length. Most similar to Channa punctata, but differs in
Native range (about 16-24º N) indicates this species to having a narrow, pointed snout, pelvic fins 50 percent
be subtropical to tropical. or more longer than the pectorals, and there is one large
Reproductive habits: No information, but scale (rarely two on one side) on either underside of the
probably a nest builder as are most other snakeheads. lower jaw.

105
 Commercial importance in the United States: Environmental concerns: Like the closely
None known. Not listed on aquarist-oriented websites. related Channa punctata, this species is probably a
Commercial importance in native range: No thrust predator on other fishes and insects.
information, but its presence in markets in Myanmar
(Musikasinthorn, 1998) indicates that it is fished
commercially.

80°E 100°E
Nepal
Bhutan China

India
EXPLANATION Myanmar Laos
DISTRIBUTION OF 20°N
Channa panaw Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate
Malaysia

Equator Indonesia
0°

Channa panaw

106
 Channa pleurophthalma (Bleeker, 1851)
Ocellated Snakehead

After Bleeker, 1878

Original description: Ophicephalus pleurophthalmus Bleeker, 1851:270. Nieuwe bijdrage tot de kennis der
ichthyologische fauna van Borneo mit beschrijving van eenige nieuwe soorten van zoetwatervisschen. Natuurkd.
Tijdschr. Neder. Indië 1:259-275. Type locality: Bandjarmasin, Borneo, Indonesia. Syntype and/or Bleeker
specimen: BMNH 1880.4.21.123.
Synonyms: Ophicephalus urophthalmus Bleeker, 1852:578.
Ophicephalus spiritalis Fowler, 1904:530, pl. 9, fide Roberts, 1989:170.
Common names: ocellated snakehead; eyespot snakehead; kerandana (Kalimantan).

Native range: Padang (?) and rivers (Hari and Characters: Patch of scales on gular part of
Musi basins) of southeastern Sumatra; Kapuas and head. Dorsal fin with 40-43 rays; anal fin with 28-31
Barito basins of Kalimantan (southern and rays. Lateral line scales 57-58; scale rows between
southwestern Borneo; Roberts, 1989; Kottelat and lateral line and anterior base of dorsal fin 5½. Single
others, 1993). Absent from peninsular Malaysia (Ng
row of canine-like teeth on prevomer; single row of
and Lim, 1990).
small teeth and single row of 4-5 canines on palatines.
Introduced range: Introductions unknown.
Four to five prominent ocelli on sides of body, anterior-
Size: To 40 cm.
most ocellus on opercula, usually 3 to 4 on body
Habitat preferences: No information found, but (Kottelat and others, 1993).
probably a riverine species.
Temperature range: No information found. Commercial importance in the United States:
Nevertheless, native range is equatorial, indicating a Often listed on aquarist-oriented websites. This species
solely tropical species. has attractive markings (red to yellow ocelli) on the
Reproductive habits: No information found, but body that has made it of interest to aquarists. Ng and
likely a nest builder as are most other snakeheads. Lim (1990) indicated it to be among the two highest-
Feeding habits: No specific information, but priced aquarium fishes. Not known for sale in live-food
likely a predator as are other snakeheads. fish markets.

107
 Commercial importance in native range: Environmental concerns: Ng and Lim (1990)
Kottelat and others (1993) indicated Channa stated that the aquarium fish trade is growing in
pleurophthalma as an important food fish in Indonesia Indonesia and Malaysia, indicating that this species,
and Sumatra. Lee and Ng (1991) noted that it is sold in among others, may have become more readily available
markets of Sumatra and Kalimantan. Dudley (2000) for sale in the U.S. Like other snakeheads, this species
reported it in the fishery of Danau Sentarum Wildlife is probably a thrust predator. Nevertheless, it is an
equatorial species that could probably survive in very
Reserve, Kapuas River, Kalimantan.
few waters (thermal springs, perhaps extreme southern
peninsular Florida, and Hawaii) if introduced into
the U.S.

100°E 120°E 140°E

China

Myanmar PACIFIC
20°N
OCEAN
Thailand Laos
Philippines

EXPLANATION Vietnam
DISTRIBUTION OF Cambodia
Channa pleuropthalma
Native range
Malaysia
Malaysia
INDIAN Indonesia Equator
0°
OCEAN
0 500 MILES Indonesia
Indonesia
0 500 KILOMETERS
Scale is approximate

Channa pleurophthalma

108
 Channa punctata (Bloch, 1793)
Spotted Snakehead

After Bloch, 1793; images reversed from original pl. 358

After Munro, 1955

After Bleeker, 1878

Original description: Ophicephalus punctatus Bloch, 1793:139, pl. 358. Naturgeschichte der Ausländischen
Fische, 7:i-xiv + 1-144, pls. 325-360. Type locality: rivers and lakes of Malabar coast, southwestern India. Syntype:
ZMB 1394.
Synonyms: Ophicephalus karruwey Lacepède, 1801:551-552.
Ophicephalus lata Hamilton, 1822:63.
Ophicephalus indicus McClelland, 1842:583.
Ophiocephalus affinis Günther, 1861:470.
Common names: spotted snakehead; green snakehead; dolla or daula (Pakistan); taki, lata (West Bengal, India);
phool-dhok (Bihar, India); duloora, daula (Punjab, India); soal (Jammu, India); gorissa or godissa (Orissa, India);
matta-gudisa, burada-matta (Andhra Pradesh, India); korava-patti (Tamil Nadu, India); kayichal, arracan (Kerala,
India); korava, juchi, belikkorava (Karnataka, India); (Talwar and Jhingran, 1992); cheng (northern Bengal, India;
Shaw and Shebbeare, 1938); mada kanaya (Sri Lanka; Pethiyagoda, 1991).

109
 Native range: Kabol (Kabul) River basin, north-central India. This species is reported to tolerate
Afghanistan, eastward through Khyber Pass into Indus a wide range of pH levels with 100 percent survival
River basin, Pakistan; rivers of the plains of India; Sri over 72 hours from pH 4.25 to 9.4 (Varma, 1979). It is
Lanka; southern Nepal (Edds, 1986a,b, 1993; Shrestha, unable to tolerate salinities above 6 ppm (Mansuri and
1990; Pethiyagoda, 1991); Bangladesh; Myanmar; others, 1979). Nevertheless, Khora and Rao (1994)
eastward to Yunnan Province, southwestern China recorded this snakehead as present and of commercial
(Coad, 1981; Talwar and Jhingran, 1992). Musikas- importance in Bahuda Estuary, Ganjam District,
inthorn (1998) reported that this species is not present Orissa, India; this estuary drains into the Bay of
in Myanmar (replaced by Channa panaw in the Bengal.
Ayeyarwaddy (=Irrawaddy) and Sittang River basins), This airbreather can live indefinitely without
and reports from Yunnan Province, China, are probably rising to the surface if water is well oxygenated (6.0
misidentifications. He further stated that the eastern ml/L and above), but will die within 2-3 hours at an O2
terminus of the range of C. punctata is the Ganges- level of 2.79 ml/L if access to the surface is prevented
Brahmaputra River basin. Jhingran (1984) lists this (Pandey and Chanchal, 1977).
species as absent from Sri Lanka, but Pethiyagoda Temperature range: The native range extends
(1991) and Devi (1992) included Sri Lanka within its from about 34º N southward to about 7º N and altitudes
native range. up to 1,830 m, suggesting the species can exist in
Introduced range: Smith (1950) reported this temperate (for example, Kabol River basin of Afghani-
species from the vicinity of Delagoa Bay, Maputo, stan) to tropical regions. Jain and Garg (1984) provided
southern Mosambique, considered an introduction an upper lethal limit of 40 ºC and lower limit of 9 ºC
from Asia (Teugels and others, 1986). Jim Cambray, for fish acclimated to 30 ºC, but found seasonal differ-
Albany Museum, Grahamstown, South Africa ences. Those collected during summer months toler-
(personal commun., 2001) located the specimen ated a high of 40 ºC and a low of 2 ºC, whereas those
(AM/G3714) and provided a digital photograph to us. captured in winter showed an upper limit of 36 ºC and a
It appeared to be a specimen of Channa striata rather low of 4 ºC.
than C. punctata. We borrowed the specimen through Reproductive habits: Dehadrai and others
Jim Cambray and confirmed that it is C. punctata. Paul (1973) noted that snakeheads in India are sexually
Skelton (Rhodes University, Grahamstown, South dichromic and dimorphic, and during the breeding
Africa; personal commun., 2001) reported the speci- season, males and females of Channa punctata become
men as having come from the Museum in Lourenco deep yellow ventrally up to the lateral line, with many
Marques, Mozambique, and added that no other snake- small melanophores in the ventral area of males and
head has been found or reported from that area since black blotches in females. Moreover, females have a
the Smith (1950) record. This indicates that snakeheads circular genital opening whereas in males the opening
are not established in southern Africa. is elongated.
Size: To 30-31 cm (Bardach and others, 1972; Said to be a prolific breeder, with pairs spawning
Talwar and Jhingran, 1992). (in India) throughout the year, peaking before and
Habitat preference: Stagnant waters and muddy during monsoon months (Jhingran, 1984; Talwar and
streams on plains of Pakistan and India, and abundant Jhingran, 1992). Reaches sexual maturity in 1 year.
in muddy waters up to 600 m (Talwar and Jhingran, Like many other snakeheads, this species builds circu-
1992). Shaw and Shebbeare (1938) recorded the lar nests in nearshore vegetation, and the eggs are
species from “muddy or clear streams and ponds from pelagic and guarded by both parents. Most initial
2,000 feet” (610 m) “downwards.” Quayyum and guarding appears to be by the female parent. Should the
Qasim (1962) described preferred habitat in India as young be disturbed, the female follows the young and
ponds with "a swampy bottom" and abundant aquatic the male parent joins her, sometimes charging an
vegetation, as well as ponds with sand or gravel intruder and sometimes leaving and swimming into
substrate and no vegetation; they added that "large deeper water, at which point the female attacks and
numbers can live together in a small body of water." drives away the intruder. Feeding by parents continues
Kumar and Mittal (1993) found habitat preferences during protection of the young, and when one parent
included open water to very dense masses of vegetation leaves the young, the other guards (Quayyum and
in Keoladeo National Park, Bharatpur, Rajasthan, Qasim, 1962). The nest, described as cup-shaped

110
(Chacko and Kuriyan, 1947), is about 22-23 cm in worm. No algae were found, but parts of leaves and
diameter. Breeding typically occurs at night, and takes seeds were occasionally observed. By far, the most
place twice each year (Raj, 1916). Kahn (1924) stated common food item was fishes (Reddy, 1980). In
that this species builds elaborate tunnels to the nest contrast, in polluted Hussainsagar Lake, Hyderabad,
through surrounding vegetation. Jhingran (1984) indi- Andra Pradesh State, insects and their larvae were
cated that reproduction in ponds occurs through most preferred (59.5 percent), followed by fishes and fish
of the year. The larger the parents, the more offspring larvae (12.5 percent), annelids, algae, leaves, crusta-
they produce. Fecundity is between 2,300 to 29,600 ceans, amphibians, and gastropods (Reddy and Rao,
eggs, with egg diameter peaking at slightly less than 1990). In ponds, tanks, and canals in the central delta
0.5 mm. This information is in contrast to that area of River Godavari, Andra Pradesh, only 24 percent
presented by Kahn (1924) who recorded egg size as of the diet consisted of fishes, but 41.6 percent were
2 mm. Khan (1924) also stated that hatching occurred crustaceans. Dutta (1994) found insects dominant (up
in 54 hrs at 16-26 ºC and 30 hrs at 28-33 ºC. Protec- to 100 percent) in young (3.1-4.5 cm in length)
tion of young continues for 15-20 days until juveniles followed by crustaceans (up to 25 percent) and fishes,
become demersal (Quayyum and Qasim, 1962). except for the months of June, July, and November
Spawning in southern Nepal occurs from June until when no fishes were found. All individuals above a
August (Shrestha, 1990). Lowe-McConnell (1987) length of 5 cm contained fishes. This study was
gave April to July as the spawning period in Punjab conducted on specimens captured from Gadigarh
Province, India, citing brood size as up to 500 individu- Stream, Jammu, northern India.
als; guarding occurs for up to a month or until young Wee (1982) cited a study by Panday and Dwivedi
are 10 cm long. Reddy (1979) stated that spawning (1974) in which it was shown that Channa punctata has
occurs once per pair during July to October, with maxi- well-developed olfactory organs in the nasal sacs and
mal spawning between July and August in Andhra taste buds extending into the esophagus, concluding
Pradesh Province, southeastern India. This agrees with that this species locates food by odor. He also cited
Bhuiyan and Rahman (1984) who reported a single Gerald (1976b) as reporting maximal feeding activity
annual spawning between April and August in Bang- of this species occurs at 28 ºC. Food absorption effi-
ladesh. It appears that spawning season is largely corre- ciency was reported as 95.5 percent (Gerald, 1976a).
lated to active monsoonal periods. This is one of three Larger fish have a lower feeding rate than young
species known to spawn in ponds lacking vascular (Gerald, 1976a).
aquatic plants (Parameswaran and Murugesan, 1976b).
Characters: Body elongated, mostly rounded; 4
Joshi and Sathyanesan (1981) reported finding
or 5 scales between orbit and angle of preopercle, 12 or
stage I oöcytes in testicular tissue of 2 of over 100
13 predorsal scales; pelvic fins more than half the
specimens of Channa punctata, all collected in Decem-
length of pectoral fins, extending to anal fin; pectoral
ber when the species is reproductively inactive.
fins plain, no vertical bands, 15-16 rays, about 75
Oöcytes were scattered through the testis and the
percent of pectoral fin length; anal fin rays 28-37;
gonads appeared to be a normal testis externally.
dorsal fin rays 28-32, rarely 33; caudal fin rounded.
Feeding habits: Young (1.5-3.0 cm) feed prima- Mouth large; lower jaw with 3 to 6 canines behind a
rily on zooplankton, with rotifers, insect, and crusta- single row of villiform teeth that widen to 5 or 6 rows
cean larvae constituting most of the diet. Adults at jaw symphysis. Predorsal scales 12; scales in lateral
consume fishes, insects, and aquatic vegetation series 37-40; scales on top of head large, arranged in a
(Quayyum and Qasim, 1962), the latter probably rosette between orbits, the frontal scale of which has an
ingested in the process of capturing animal prey. open lateralis pit, forming the center of the rosette
The species is an opportunistic feeder. In canals (Talwar and Jhingran, 1992; Jayaram, 1999). Life
and irrigation ponds near Guntur, Andra Pradesh State, colors vary from black to pale green on dorsum and
India, stomach contents consisted of 13 species of sides, ventral sides white to pale yellow, sometimes
small fishes with young of Channa punctata (one each) with red tinge; several dark blotches on lower sides;
found in only three individuals, indicating that canni- occasionally black spots on body and dorsal, anal, and
balism is rare. Fish bones, scales, fin rays, etc., were caudal fins. Dorsal, anal, and caudal fins dark gray,
common. Insects comprised the second tier of ingested sometimes with reddish edge; pectoral and pelvic fins
food, followed by crustaceans, tadpoles, and an annelid pale orange (Talwar and Jhingran, 1992).

111
 Commercial importance in the United States: They are considered to be a delicacy and demand high
Although occasionally listed on aquarist-oriented prices (Talwar and Jhingran, 1992). Pethiyagoda
websites and in aquarium fish books, this species does (1991) noted that it is popular as a food fish in Sri
not appear to have been important to the aquarium fish Lanka and is also used as bait for catching larger snake-
trade in North America. It is not known to have been heads.
imported for culture or live-food fish market sale. Environmental concerns: Because of their
voracious, carnivorous feeding habits, snakeheads are
Commercial importance in native range: regarded as pests in India due to their devastation of
Snakehead fishes in general are regarded as important other fishes (Talwar and Jhingran, 1992), apparently in
fishery resources in India and elsewhere, fished pond or culture situations where other desired species
commercially and some species utilized in aquaculture. exist.
Quayyum and Qasim (1962) reported this species as Comments: Banerjee and others (1988)
"the main bulk of pond fishery in the plains of northern recorded the diploid chromosome number of Channa
India." Rao and Durve (1989) reported Channa punc- punctata as 32. Dhar and Chatterjee (1984), however,
tata as one of three snakehead species fished commer- found two groups, one with 32 and another with 34,
cially in Lake Jaisamand, the oldest reservoir in India. indicating a species complex.

60°E 80°E 100°E 120°E

North
40°N
Korea

Afghan-
Iran istan China South
Nepal Korea
Bhutan
Pakistan

Saudi India
Arabia Myanmar
20°N

Laos
Bangladesh Thailand
Vietnam

Cambodia
Thailand
Malaysia

Equator
0°

INDIAN OCEAN Indonesia

EXPLANATION 0 1,000 MILES

DISTRIBUTION OF
Channa punctata 0 1,000 KILOMETERS
Native range Scale is approximate

Channa punctata

112
 Channa stewartii (Playfair, 1867)
Golden Snakehead

After Playfair, 1867

Original description: Ophiocephalus stewartii Playfair, 1867:14, pl. 3. On the fishes of Cachar. Proc. Zool. Soc.
Lond. 1867(1):14-17, pl. 3. Type locality: Cachar, Assam, India. Syntypes: BMNH 1867.2.14.19-20.
Synonyms: None known.
Common names: golden snakehead; Assamese snakehead; sengalee (Assam, India; Talwar and Jhingran, 1992);
helae (Nepal).

Native range: Endemic to Brahmaputra (upper, Feeding habits: No specific information, but
middle, lower) River basin of India and Bangladesh, likely a thrust predator like other snakeheads.
and the Ganges River basin from southern Nepal
Characters: No patch of scales on gular region
southeastward (Sen, 1985; Talwar and Jhingran, 1992;
of head. Dorsal fin with 39-40 rays; anal fin rays 27.
Musikasinthorn, 2000). In southern Nepal, it occurs in
Pelvic fin about one-third as long as pectoral. Lateral
the Kamala, Bagmati, Koshi, Gandaki, and Karnali
line scales 47-50; scale rows between preopercular
River basins (Shrestha, 1990).
angle and posterior border of orbit 4-5; predorsal scales
Introduced range: Introductions unknown. 13. Dorsal fin originates above base of pectoral fin.
Size: To 25.4 cm (Sen, 1985). Black spots on many body scales.
Habitat preference: Flowing and standing water Commercial importance in the United States:
(Talwar and Jhingran, 1992); marshes and estuaries, Rarely mentioned in aquarist-oriented websites but
and tolerant of limited salinity (Sen, 1985). Sen and available through aquarium fish trade. Not known to be
Dey (1984) recorded this species from altitudes above available in live-food fish markets.
1,500 m in Meghalaya, a plateau north of Bangladesh
and south of the Brahmaputra River, India. Commercial importance in native range:
Talwar and Jhingran (1992) stated that this species is of
Temperature range: No specific information. only minor fishery interest.
Native range is about 22-27º N, indicating a warm
temperate to subtropical species. Environmental concerns: Likely a thrust predator,
feeding on other fishes and invertebrates.
Reproductive habits: No specific information,
but probably a nest builder providing eggs and young Comments: The diploid chromosome number of
with parental care. Channa stewartii is 104 (Rishi and Haobam, 1984).

113
 80°E 100°E
Nepal
Bhutan China

India
EXPLANATION Myanmar Laos
DISTRIBUTION OF 20°N
Channa stewartii Bangladesh
Native range Bay of
Bengal Thailand

0 1,000 MILES

Vietnam
0 1,000 KILOMETERS Thailand Cambodia
Scale is approximate
Malaysia

Equator Indonesia
0°

Channa stewartii

114
 Channa striata (Bloch, 1793)
Chevron Snakehead

After Bloch, 1793; image reversed from original pl. 359

After Munro, 1955; juvenile

Note “pseudo-ocellus” on posterior lobe of dorsal fin, a juvenile characteristic (Lee and Ng, 1991).

After Bleeker, 1879

Original description: Ophicephalus striatus Bloch, 1793:141, pl. 359. Naturgeschichte der Ausländischen Fische,
7:I-xiv + 1-144, pls. 325-360. Type locality: Malabar, southwestern India. Syntypes: ZMB 1400; ZMB 6522.
Synonyms: Ophiocephalus wrahl Lacepède, 1801:552.
Ophiocephalus wrahl Hamilton, 1822:60, 367, pl. 31.
Ophiocephalus chena Hamilton, 1822:62, 367.
Ophicephalus planiceps Cuvier, 1831:424.
Ophicephalus sowarah Bleeker, 1845.
Ophiocephalus vagus Peters, 1868:260.
Ophiocephalus philippinus Peters, 1868:262.
(?)Ophiocephalus melanopterus Smith and Seale, 1906.

115
Common names: chevron snakehead; striped snakehead; banded snakehead; common snakehead; soali (Pakistan);
murrel (India); haal, shawl, shol (Assam, India); shol (West Bengal, India); morrul, morl, soura (Bihar, India); sowl,
dhoali, carrodh (Punjab, India); dolla (Jammu, India); sola (Orissa, India); korramennu, korra-matta (Andhra
Pradesh, India); sowrah, veralu, kaunan (Kerala, India); pooli-kuchi, koochinamarl (Karnataka, India); sohr, dekhu
(Mararashtra, India); hal path maha, lulla (Sinhalese, Sri Lanka); viral (Tamil, Sri Lanka); pla chon or pla chorn
(Thailand); trey phtuok (juveniles) and trey raws (adults; Cambodia); ikan aruan, haruan, ruan, tomam paya
(Malaysia); gabus (Java); delak, gabus, telak (Kalimantan), cá lóc (Vietnam); dalag, dalak (Tagalog or Moro,
Philippines); bakule or bulig (young; Tagalog or Moro, Philippines); pongee (Hawaii, although that name is a general
name for snakeheads; Mike Yamamoto, personal commun., 2003).

Native range: Pakistan (Indus River basin; Its introduction into the Philippines probably occurred
Mirza, 1975), most drainages of India, southern Nepal in the early to mid-1800s, indicated by two synonyms
(Koshi, Gandaki, and Karnali River basins; Shrestha, (Ophiocephalus vagus and O. philippinus) described
1990), Sri Lanka (Mendis and Fernando, 1962; from the Philippines by Peters (1868). Although
Fernando and Indrassna, 1969; Pethyagoda, 1991); Jayaram (1999) included Borneo in the native range of
Bangladesh, Myanmar, Thailand, Cambodia, southern this species, Roberts (1989) hinted that its presence in
China, Malay Archipelago including Malaysia, western Borneo may have resulted from introductions.
Sumatra, Borneo (Pethiyagoda, 1991; Rainboth, 1996; Ralf Britz (personal commun., 2002) has advised
Jayaram, 1999); Sabah (Inger and Kong, 1962); that reports of this species from Madagascar are in
western Java (Giltay, 1933; Roberts, 1993); Vietnam, error, the result of misidentification of the blotched
Laos (Yên and others, 1992; Kottelat, 2001a,b). This is snakehead, Channa maculata. He also examined a
an amazingly extensive "native" distribution for any specimen labeled as C. striata (USNM 126588),
freshwater fish, indicating that Channa striata is quite collected by Jordan and Evermann on Oahu, Hawaii, in
probably a species complex. 1901, that was also C. maculata. Two specimens we
borrowed from the Bernice P. Bishop Museum in
Introduced range: Channa striata has been
Honolulu (BPBM 1759 and BPBM 3798), collected in
considered the most widely introduced species of
the early 1900s on Oahu and labeled as C. striata, are
snakehead. Various reports indicate it was released into
C. maculata. Specimens borrowed from the California
Hawaii before 1900, established (Jordan and Ever- Academy of Sciences (CAS 17710, 1 specimen; CAS
mann, 1903; Cobb, 1905; Smith, 1907; Tinker, 1944; 108133, 3 specimens) collected on Oahu also proved to
Brock, 1952, 1960); Madagascar, in 1978, established be C. maculata and not C. striata. We are convinced
(Raminosoa, 1987; Reinthal and Stiassny, 1991; that all early records of C. striata from Hawaii are
Stiassny and Raminosoa, 1994; Lévêque, 1998); misidentifications of C. maculata. Although most
Philippines, date unknown (Seale, 1908; Herre, 1924, records for Hawaii cite establishment only on Oahu,
1934; Conlu, 1986); Vogelkop Peninsula, Papua, Indo- Morita (1981) reported that it also occurred on Kauai.
nesia, probably during 1970s or 1980s, established Channa striata is now present and established on Oahu
(Allen, 1991) and identification confirmed by photo- but confined to a fish culture facility. The first imports
graphs provided by Gerald Allen (personal commun., of this species occurred in the early 1990s. It is likely
2002); Sundaland, Sulawesi, Lesser Sundas, Moluccas, that many identifications of C. striata on islands of the
date unknown, established (Welcomme, 1981; Kotellat Indian and Pacific Oceans are in error and that
and others, 1993; Lever, 1996); Fiji, establishment C. maculata may prove to be more widely introduced
questionable (Maciolek, 1984; Eldredge, 1994); than previously thought.
Mauritius, established (Parameswaran and Goorah, Following publicity accompanying discovery of
1981; Welcomme, 1988; Lever, 1996); New Caledonia, an established population in Maryland of northern
establishment questionable (Maciolek, 1984); Guam, snakehead, Channa argus, in June-July 2002, the lead
introduction unsuccessful (Maciolek, 1984; Eldredge, investigator received an email message from an indi-
1994). Herre (1924) recorded the source of introduc- vidual (Clifford Faik) in East Sepik Province, Papua
tion into Hawaii as southern China. Kottelat and New Guinea, who had seen the northern snakehead
others (1993) reported some populations in China to report on the CNN website. He stated that a similar
have been introduced but gave no specific locations. snakehead was now present in rivers of Sepik Province.

116
Temperature regimes at that locality would preclude through the gills and skin of almost 42 percent once
presence of northern snakehead, but is likely indicative bimodal respiration began. Pandian (1982) reported that
that the chevron snakehead is the species involved, fingerlings of this species spend up to 15 percent of the
probably introduced from populations established in time in surfacing and related activities.
Papua, Indonesia. If so, this would be the first record of Varma (1979) recorded a pH range for Channa
a snakehead from Papua New Guinea. striata of 4.25 to 9.40 with 100 percent survival over
Size: To 90 cm (Bardach and others, 1972); 72 hours, and 90 percent survival at pH 3.10 for the
91.4 cm (Sen, 1985). Can attain a length of 30-36 cm same period.
in 1 year (Bhatt, 1970). Talwar and Jhingran (1992) Temperature range: No specific information,
stated that this species is sexually mature at 30 cm, but but native range lies between about 32º N and 7º N,
added that 2 years were required to reach that size. indicative of a fish that is temperate to tropical.
Murugesan (1978), however, recorded a growth rate
Reproductive habits: Lee and Ng (1991)
of 1.3 to 3.0 mm/day for the first 3 months, slowing to
indicated the species as solitary except during spawning
0.3 to 0.9 mm/day thereafter. He also reported lengths
seasons. In India, pairs breed during most months of the
of 25-27 cm in 13½ months and 23.4-31.7 cm in 9½
year, laying a few hundred to more than 1,000 amber-
months in Kerala State, India; 32 cm in 2 years in West
colored eggs (Parameswaran and Murugesan, 1976a;
Bengal; and 30.5l cm in 2 years in Madras. In rivers of
Talwar and Jhingran, 1992). Peak spawning coincides
Uttar Pradesh, chevron snakeheads grew to 32 cm in
with peak rainfall (Parameswaran and Murugesan,
2 years.
1976a). Howell (1913) said the eggs average about
Habitat preference: Freshwater ponds and 1.25 mm and are nonadhesive, hatching in 1 to 3 days.
streams, usually in stagnant muddy waters; primarily Females mature about 30 cm in length at about 2 years
found on plains in India (Talwar and Jhingran, 1992). of age (Talwar and Jhingran, 1992; Ali, 1999). Parents
It occurs in reservoirs in Sri Lanka (Fernando and clear a shallow depression by biting off aquatic vegeta-
Indrassna, 1969). Nevertheless, in Malaysia this tion (Ling, 1977). Nevertheless, Alikunhi (1953)
species is reported to exist in rivers, lakes, swamps, remarked that Channa striata will spawn in the absence
rice paddies, mining pools, and roadside ditches of vegetation. Eggs float to the surface after fertilization
(Mohsin and Ambak, 1983; Lee and Ng, 1991). Ng (Lee and Ng, 1991). The pelagic eggs are guarded by
and Lim (1990) listed this species from “open country both parents in the Philippines (Lowe-McConnell,
areas,” adding that it is the primary snakehead in shal- 1987) and possibly throughout the native range of the
low waters (1 m or less) with dense vegetation. In species. Nevertheless, Herre (1924) stated that one or
India, it can be found in reservoirs and rice paddies the other parent guards the nest at all times, and that if
(Jhingran, 1984). In Keoladeo National Park, Bharat- food becomes scarce, parents become cannibalistic on
pur, Rajasthan, north-central India, it may be found in the young. He further indicated that in the Philippines,
open water to dense mats of aquatic vegetation (Kumar C. striata spawns throughout the year and that many,
and Mittal, 1993). Herre (1924), Umali (1950), and perhaps all, breed twice annually. Ali (1999) confirmed
Conlu, 1986) recorded it from lakes and lowland rivers ripe females present throughout the year in ricefields in
in the Philippines, and its introduction to two crater Perak, northwestern Malaysia. Peak spawning in south-
lakes at an altitude about 1,050 m above sea level. western Sri Lanka occurs between May and September,
Kottelat (1998) reported a preference for “standing with a secondary spawning October through December
waters.” Lee and Ng (1991) noted that this species (Kilambi, 1986). Jhingran (1984) cited fecundity as
seems to be the most adaptable snakehead, tolerating 3,000-30,000 ova. Lee and Ng (1991) stated that they
“quite foul water” and able to move overland. had collected fry without seeing parents nearby. They
Channa striata is an obligate airbreather. Vive- also said that eggs hatch in 3 days in Malaysia, the fry
kanandan (1977a,b) stated that the breathing organ is developing a deep orange color. This pattern persists
developed in about 60 days during growth from a length until the young reach a length of 15 mm when only an
of 1 to 4.5 cm at 26-28 ºC. Singh and others (1986) orange lateral stripe remains. At 40 mm in length, all
noted that at 28 ºC, this species breathes aquatically until orange color is lost but a “pseudo-ocellus” appears on
18 to 20 days following hatching when young reach a the posterior lobe of the dorsal fin, a characteristic lost
length of 1.1-1.2 cm and, thereafter, becomes a bimodal in adulthood. Mookerjee and others (1948) described
breather. They measured a decrease in oxygen uptake and illustrated early development of C. striata.

117
 Feeding habits: Carnivorous, feeding on worms, heads are reported as being imported annually for food
prawns, frogs, and especially other fishes (Mohsin and purposes into mainland United States. It is unknown
Ambak, 1983). Reported as a solitary (except during how much of that market involves C. striata.
breeding season), territorial, ambush feeder (Lee and During fiscal year 1999, the U.S. Department of
Ng, 1991). Conlu (1986) stated that young fry feed on Agriculture Small Business Innovation Research
algae and protozoans, juveniles feed on small crusta- Program funded a Phase II project to the Hawaii Fish
ceans, and “adults are highly carnivorous, dreaded Company of Waialua, Hawaii, $230,000 for 24 months,
predators of other pond fish.” She added that this fish is to develop commercial culture of C. striata. Phase I
used as a predator to control tilapias in culture ponds. research had established feasibility of rearing striped
Jhingran (1984) cited larvae as feeding “on insects, snakeheads in captivity, spawning, and studies on rearing
water fleas, and fish fry,” juveniles preferring “dipteran juveniles on artificial diets. Phase II was targeted to
larvae, zooplankton, and fish fry,” and adults as “pisciv- production of larvae and juveniles through induced
orous.” Mahan and others (1978) reported that Channa spawning, additional studies on feeding, and cost-effec-
striata (32 individuals ranging from 3.5 to 36.7 cm in tive grow-out performance to marketable size. Phase III
length) fed almost exclusively on shrimp (47 percent by was designed to result in a commercial effort to produce
volume) in a lake in central Java. Dasgupta (2000) farm-raised snakeheads for Hawaii, mainland U.S., and
found that this snakehead consumed primarily insects Canada, and was to be funded ($300,000) by the
(40 percent) followed by fishes (30 percent) and crusta- Wah Wah Seafood Company, Inc., of Honolulu. Phase
ceans (10 percent) in waters of West Bengal, India. Rao III, however, was never funded.
and others (1998) noted a preference for crustaceans In retrospect, importation of Channa striata to
and fishes from ponds and canals of East Godavari Hawaii is quite recent. Perhaps based on the belief that
District, Andhra Pradesh, southeastern India. Ng and this species had been established in Hawaii for nearly a
Lim (1990) described the enlarged canine teeth of century, a permit was issued in the early 1990s to Arlo
C. striata as “cylindrical in cross section … ideal for Fast of the University of Hawaii to import C. striata for
gripping, killing, and tearing.” culture research on Coconut Island in Kaneohe Bay.
A second permit to import C. striata was issued to
Characters: Gular region of head without patch Dr. Fast in 1995 in cooperation with the person who
of scales. Mouth large; lower jaw with 4-7 canines currently cultures the species in a rockpit area at
behind a single row of villiform teeth that widen to 6 Mokuleia. The culturist had the only permit from the
rows at the jaw symphysis; villiform teeth on prevomer Hawaii Department of Agriculture to import C. striata
and palatines. Pectoral fin about half of head length. with restrictions that sale to consumers must be of
Dorsal fin with 37-46 rays; anal fin rays 23-29; pectoral fresh-killed or cooked fish (Domingo Cravalho, Jr.,
rays 15-17; pelvic rays 6; caudal fin rounded. Scales on personal commun., 2002).
top of head large with a rosette of head scales between
This species often appears in aquarist-oriented
orbits, with frontal head scales forming central plate of websites and has been sometimes listed for sale by
rosette; 9 scale rows between preopercular angle and commercial aquarium websites. Interest in its use as an
posterior border of orbit; predorsal scales 18-20; scales aquarium fish seems to be limited due to the size it
50-57 in lateral series (Talwar and Jhingran, 1992). attains and its aggressive nature toward other fishes.
Coloration is quite variable in this species or species
On July 24, 2002, a specimen of Channa striata
complex. The dorsum is often dark brown to black,
was purchased from a market in San Diego, California,
typically obscuring the chevron-like markings dorsally.
where at least two other individuals of the same species
A distinguishing marking, however, is the dark stripe
were observed (Richard Rosenblatt and Phil Hastings,
extending from just above the maxillary posteroven-
personal commun., 2002). All three were dead, on ice.
trally toward the opercular curvature. The appearance of the specimen sent to us in a digital
Commercial importance in the United States: photograph seems to be of a freshly dead individual,
Introduced population is utilized as a food resource in with no cloudiness visible in the eye.
Hawaii (Maciolek, 1984), although the species Commercial importance in native range:
involved was not Channa striata but C. maculata, the Channa striata is reported as being cultivated in Paki-
result of century-old misidentifications. In addition to stan and India. There is a “tank fishery” for this species
live fish, several thousand metric tons of frozen snake- in Tamil Nadu, India. Tanks in India and Sri Lanka are

118
“ancient irrigation reservoirs” (Fernando and Indrassna, Cream extracts of haruan tissues contain high
1969). They stated that there were more than 1,000 levels of arachidonic acid, a precursor of prostaglandin,
tanks in Sri Lanka alone. In India, the chevron snake- essential amino acids (particularly glycine), and
head is described as a popular and highly prized fish, polyunsaturated fatty acids necessary to promote
widely distributed, and the most economically impor- prostaglandin synthesis. Treating wounds with these
tant species of the genus (Talwar and Jhingran, 1992). extracts has been demonstrated to promote synthesis of
Channa striata is one of three species of snakeheads collagen fibers better than standard use of Cetrimide,
commercially fished in Lake Jaisamand, the oldest an antimicrobial quaternary ammonium compound, thus
reservoir in India (Rao and Durve, 1989). Fernando increasing tensile strength (Baie and Sheikh, 2000).
and Inrassna (1969) stated that it is the only species of
snakehead in Sri Lanka of economic value, although Lee and Ng (1991) indicated that the flesh of
three additional species (C. gachua, C. marulius, and these larger snakeheads is rejuvenating following
C. punctata) are used as food fishes. It is also cultured illnesses, prepared by being double-boiled with herbs,
in Vietnam (Pantulu, 1976; Bard, 1991), Thailand, Java and only the soup is consumed. Nevertheless, for the
(Hofstede and others, 1953), and the Philippines (Guer- soup to be effective in recovery, it is firmly believed
rero, 2000). Bard (1991) noted that this species is the that the fish must be killed just before cooking,
most expensive fish produced by aquaculture in north- dispatched with careful but firm blows to the head with
ern Vietnam. Ali (1999) cited it as “a popular food fish a mallet. Herre (1924) reported much the same for the
in Malaysia” remarking that ricefields have provided Philippines. Conceivably, this could be a reason that
the largest source of this fish. Populations in Malaysia obtaining live snakeheads in live-food fish markets is
are reported to be depressed due, apparently, to over- considered important to some persons of southeast
fishing, raising costs for live specimens. China is Asian descent living in the United States. Seale (1908)
culturing C. striata and some of the product is being cited this species "as one of the most wholesome fishes
canned for sale in Malaysia (Wan Ahmad, personal and are given to invalids" in India.
commun., 2001).
Bard (1991) noted that Channa striata is
Lee and Ng (1991) cited this species as the most cultured in Vietnam and is a highly desired and expen-
economically important member of the snakeheads sive fish in the markets of Hanoi, with a price/kg
and noted that it is cultured throughout most of its matching that of beef.
range. Hofstede and others (1953) cited this species as
Environmental concerns: Adults of this species
bringing “the highest prices at the markets” in Indone-
are considered to be highly predacious, ambush feeders
sia. It is sold either fresh or alive in Cambodian
markets (Rainboth, 1996). In the Danau Sentarum on other fishes. In addition, their adaptability to living
Wildlife Reserve of Kalimantan, chevron snakehead in turbid or clear waters, their apparent ability to toler-
comprised 13 percent of the setline fish catch using ate subtropical to warm temperate climates, suggests
small (size 12-16) hooks from the Kapuas River the probability of establishment if introduced into
(Dudley, 2000). waters of the extreme southern U.S. The apparent north-
ern limit within its native range (32° N) is equivalent to
Ng and Lim (1990) and Lee and Ng (1991) a potential range from Savannah, Georgia, to just north
indicated that Channa striata, along with C. micropeltes of Ensenada, Baja California del Norte, México, in
and C. lucius, are utilized for medicinal purposes, North America. Temperature regimes in the southwest-
particularly in Indonesia and Malaysia. Mention was
ern U.S. would permit establishment well north of 32º
made of use in a postnatal diet and during recupera-
N, probably as far north as the Los Angeles basin, Cali-
tion from illnesses or surgery (Lee and Ng, 1991).
fornia; Phoenix, Arizona; or Las Cruces, New Mexico.
While no specifics were given as to how the fish were
used following surgery, a neighbor of one of the Comments: Banerjee and others (1988)
authors (WRC), a Malaysian by birth, said that the recorded the diploid chromosome number of Channa
oils from the “haruan” are used to greatly reduce scar- striata from India as 40, but Donsakul and Magtoon
ring. She added that she had seen the results and “it is (1991) reported a count of 44 for this species from
true” that scar tissue is dramatically reduced to a Thailand, indicating that it represents a species
minimum. complex.

119
 40°E 60°E 80°E 100°E 120°E 140°E 160°E 180°E 160°W 156°W
22°N
60°N Oahu
Russia Hawaii 20°N
Pacific Ocean
0 50 MILES
Kazakhstan
Mongolia 0 50 KILOMETERS

North
40°N Turkey Korea
Japan
Syria Afghan-
Iraq Iran Nepal China
istan
Bhutan South PACIFIC
Pakistan Korea
Saudi OCEAN
Arabia India
20°N
Myanmar Laos
Bangladesh Vietnam
Philippines
Thailand Cambodia
Malaysia
Equator
0°

INDIAN Indonesia Indonesia

New Guinea
Madagascar
OCEAN
Fuji

20°S
Australia New
Caledonia

EXPLANATION 0 2,000 MILES

DISTRIBUTION OF Channa striata

0 2,000 KILOMETERS
Native range Scale is approximate
Introduced range

Distribution of Channa striata

See comments above under Introduced range, as some reports of C. striata on Indian Ocean (for
example, Madagascar) and Pacific Islands (for example, Hawaii) are misidentifications of C. maculata that
require further investigation.

120
 Parachanna africana (Steindachner, 1879)
Niger Snakehead

After Boulenger, 1916

Original description: Ophiocephalus africanus Steindachner 1879:31. Über einige neue und seltene Fischarten aus
den zoologischen Museen zu Wien, Stuttgart und Warschau. Anz. Akad. Wiss. Wien. 16(4):29-34. Type locality:
Lagos, Nigeria. Holotype: SMNS (no number provided).
Synonyms: Channa africanus (Steindachner, 1879).
Parophiocephalus africanus (Steindachner, 1879).
Common names: Niger snakehead; African snakehead.

Native range: Southern Bénin to southern Characters: Patch of scales present in gular
Nigeria, primarily the Ouémé River and Niger basin region. No canines on prevomer or palatines. Trans-
(Bonou and Teugels, 1985; Skelton, 1988). verse scales 19-24; lateral line scales 73-83. Dorsal
Introduced range: Introductions unknown. rays 45-48; anal rays 32-35. Head slightly depressed
anteriorly and covered with large scales. Lower jaw
Size: To 32 cm.
slightly longer than upper jaw with 3 to 4 large canine
Habitat preference: Bonou and Teugels (1985) teeth. Coloration distinct among African snakeheads in
noted that there was little known of the biology of this having a series of forward-pointing chevrons on the
snakehead. Daget and Iltis (1965) considered this side of the body posterior to the pectoral fins that
species as a Guinean form that occupied waters in extend upward to the base of the dorsal fin (Bonou and
forested areas. Teugels and others (1992) commented Teugels, 1985).
that this species is limited to coastal sections of rivers. Commercial importance in the United States:
Temperature range: No specific information. Sometimes listed on aquarist-oriented websites and has
Nevertheless, the native range is equatorial, indicating been periodically sold through aquarium fish retailers.
a strictly tropical species. Unknown in live-food fish markets.
Reproductive habits: No specific information Commercial importance in native range:
located. Likely a nest builder that provides parental Unknown, but probably available in live-food fish
protection to young like other snakeheads. markets.
Feeding habits: No specific information. In Environmental concerns: Likely a thrust preda-
considering this species as a game fish, Copley (1952) tor. Native range is equatorial indicating that if intro-
remarked that it ate frogs and worms, as well as fishes. duced this species would be restricted to
Probably a thrust predator like other channid fishes. tropical/subtropical waters.

121
 20°W 0° 20°E 40°E
Mediterranean
Sea

co
oc
or
M
Algeria Libya
Egypt
Saudi
Arabia

Re
20°N
AFRICA

d
Chad

Se
a
Sudan
Burkina
EXPLANATION Nigeria
Ivory
Central Ethopia
DISTRIBUTION OF Coast
African Republic
Parachanna africana
Ghana Cameroon Dem.
Native range Togo Benin Rep.
0° of the
Equator Gabon Congo
0 1,000 MILES Congo Tanzania Indian
Atlantic Ocean
0 1,000 KILOMETERS Ocean

e
Angola Zambia

u
biq
zam
Nambia
20°S

Mo
Madagascar

South
Africa

Parachanna africana

122
 Parachanna insignis (Sauvage, 1884)
Congo Snakehead

After Sauvage, 1884

After Boulenger, 1916

Original description: Ophiocephalus insignis Sauvage, 1884:195, pl. 5, fig. 3. Notes sur des poissons de
Franceville, Haut Ogooué. Bull. Soc. Zool. France 90, 1884:193-198. Type locality: upper Ogooué River, Gabon.
Syntypes: MNHN 1844-0301 to 0305.
Synonyms: Ophiocephalus obscurus Poll, 1942.
Parophiocephalus insignis Daget and Stauch, 1963.
Ophiocephalus obscurus Matthes, 1964.
Common names: Congo snakehead; light African snakehead; ifoïfoli (juvenile; Zaire), foli (adult; Zaire).

Native range: Ogooué River basin, Gabon; Zaire Temperature preference: No specific informa-
River basin, Congo and Zaire. Bonou and Teugels tion. Nevertheless, the native range of this species is
(1985) recorded this species from above Boyoma equatorial/tropical.
(Stanley) Falls in the Lualaba (Congo) River. Guy Reproductive habits: Gosse (1963) commented
Teugels (personal commun., 2002) stated that he that little is known of the reproduction of this snake-
doubted this distribution to be the result of an head. Nevertheless, he had observed juveniles (“de 30 à
introduction into the upper Lualaba. 40 mm”) in groups, guarded by a large adult. He mis-
Introduced range: No introductions known. identified this species as Ophiocephalus obscurus in
Size: To about 45 cm. central Zaire.
Habitat preference: Bonou and Teugels (1985) Feeding habits: Gosse (1963) described this
noted that no specific studies of the biology of this species as “un ichthyophage typique.” He noted that
species had been undertaken. General information indi- stomach contents of nine specimens contained prima-
cates this snakehead occupies creeks, rivers, lakes, and rily fishes (Pelmatochromis, Tilapia, Hemichromis,
lagoons, and sometimes in deep waters in calm areas. Xenomystus, cyprinodonts and fish remains).

123
 Characters: Patch of scales present in gular through these blotches. Dark stripe on side of head
region. No canine teeth on prevomer or palatines. extending from posterior rim of orbit to upper part of
Scales in a transverse line 25-33 with 7-10 above the operculum.
lateral line and 16-22 below; lateral line scales 73-86. Commercial importance in the United States:
Dorsal fin rays 40-44; anal fin rays 27-31. Head Rarely listed on aquarist-oriented websites. Unavail-
slightly depressed anteriorly and covered with large able in live-food fish markets.
scales. Lower jaw longer than upper jaw with 4-5 Commercial importance in native range: No
well-developed canine teeth on each side. Lateral line specific information, but probably available in live-
complete. Coloration is distinct among African chan- food fish markets in the Congo basin.
nids in that there are chevron-shaped bars across the Environmental concerns: Likely a thrust preda-
middle of the back that extend up onto the dorsal fin; tor showing a preference for fishes. If introduced, it
central area of sides contain 4-5 dark blotches that could probably survive only in tropical/subtropical
are distinct from each other; lateral line passes conditions.

20°W 0° 20°E 40°E

Mediterranean
Sea

co
oc
or
M Algeria Libya
Egypt
Saudi
Arabia

Re
20°N
AFRICA

d
Chad

Se
a
Sudan
Burkina
EXPLANATION Nigeria
Ivory
Central Ethopia
DISTRIBUTION OF Coast
African Republic
Parachanna insignis
Ghana Cameroon Dem.
Native range Togo Benin Rep.
0° of the
Equator Gabon Congo
0 1,000 MILES Congo Tanzania Indian
Atlantic Ocean
0 1,000 KILOMETERS Ocean ue

Angola Zambia
biq
zam

Nambia
20°S
Mo

Madagascar

South
Africa

Parachanna insignis

124
 Parachanna obscura (Günther, 1861)
African Snakehead
Type species of Parachanna Teugels and Daget, 1984

Reprinted with permission from Guy G. Teugels from: Bonou, C.A., and G.G. Teugels. 1985. Révision
systématique du genre Parachanna Teugels et Daget 1984 (Pisces: Channidae). Rev. Hydrobiol. Trop.
18(4):267-280.

Original description: Ophiocephalus obscurus Günther, 1861:476. Catalogue of the acanthopterygian fishes in the
collection of the British Museum, 3:i-xxxv + 1-586. Type locality: West Africa. Syntypes: BMNH 1849.12.25.2-3.
Synonyms: Ophiocephalus (Parophiocephalus) obscurus Senna, 1924.
Channa obscura Myers and Shapovalov, 1932.
Parophiocephalus obscurus Berg, 1940.
Common names: African snakehead; snakehead; fakka (Arabic); ojul (Shilluk); abioth (Dinka); jul (Nuer);
woroko (Zande; Bailey, 1994).

Native range:White Nile, specifically the Jebel flowing water, the species occupies calm areas. Lowe-
and Ghazal systems of Sudan (Bailey, 1994) and McConnell (1988) indicated this species as “widely
Gambela region of Ethiopia (Golubtsov and others, distributed” in marshy habitats and also found in bank
1995); Lake Chad basin; Zaire (Congo), Cross, Niger, vegetation of river channels. Teugels and others (1992)
and Senegal River basins (Boulenger, 1907, 1916; listed the species as common in stagnant side channels
Irvine, 1947; Teugels and Daget, 1984; Bonou and of the Cross River, Cameroon-Nigeria, and found
Teugels, 1985; Skelton, 1988; Teugels and others, among waterlogged fallen trees and leaf debris.
1992). Boeseman (1957) recorded the species from Temperature range: No specific information,
Stanley Pool, Lualaba (Congo) River. Said to be but the native range is in equatorial Africa indicating a
“widely distributed” in Ghana and noted as both useful strictly tropical species.
as food and aquarium species (Dankwa and others,
1999). Reproductive habits: Bonou and Teugels
(1985) stated that little was known of the reproductive
Introduced range: Not known to have been
behavior of Parachanna obscura. Gosse (1963) stated
introduced.
that young are guarded by a large adult. Likely a nest
Size: To 35 cm (Bailey, 1994). builder like other channids. Under monoculture condi-
Habitat preference: Bailey (1994) indicated a tions in southern Nigeria, reproductive activity was
preference for marginal vegetation and floodplain habi- greatest in October and November (Victor and
tat in the Sudan. Bonou and Teugels (1985) stated that Akpocha, 1992). Fecundity in this monoculture pond
this species occupies a wide variety of habitats includ- varied, with stages III, IV, and V ovaries containing
ing streams, rivers, lakes, lagoons and marshes. In 35-4,010 oöcytes. Moreover, fecundity was found not

125
to be correlated to length or weight, but this could Dorsal fin rays 39-45; anal fin rays 26-32. Head
result from “poor culture conditions” (Victor and depressed anteriorly, relatively long and covered with
Akpocha, 1992). large scales. Lower jaw slightly longer than upper jaw,
Feeding habits: Bonou and Teugels (1985) cited with 4-6 well-developed canines. Lateral line typically
the species as feeding on other fishes, noting that in complete, rarely discontinous. Coloration distinct
Bénin it is used to control the young of tilapias in among African snakeheads in having a series of dark
aquaculture ponds. Citing Blache and others (1964), blotches, some of which may coalesce, and no chevron-
they recorded the food of young as copepods and insect shaped bars across the middle of the back.
larvae. Adults appear to prey on other fishes (Copley, Commercial importance in the United States:
1952; Poll, 1957; Gosse, 1963; Teugels and others, Sometimes listed on aquarist-oriented websites. Not
1992). Adebisi (1981) noted that juveniles fed on known to have been available in live-food fish markets.
prawns, copepods, and aquatic insect larvae whereas Commercial importance in native range:
adults fed only on fishes in southern Nigeria. Probably available in local live-food fish markets. This
In monoculture in a Nigerian pond, young of this snakehead is being cultured in Ghana (Morrice, 1991),
species (10-16 cm standard length) fed primarily on Nigeria (Ajana, 1983; Victor and Akpocha, 1992), and
detritus and larval insects, whereas larger individuals Bénin (Jackson, 1988), and was recommended for
(16-24 cm standard length) contained “fish parts” and culture in the Central African Republic (Micha, 1974).
juveniles, with insects and fish making up the bulk of All these localities are within the native range of this
the diet (Victor and Akpocha, 1992). species.
Characters: Patch of scales present in the gular Environmental concerns: Known to be a preda-
region. No canine teeth on prevomer or palatines. tor, particularly of other fishes (Copley, 1952; Poll,
Transverse scales 19-24; lateral line scales 65-78. 1957; Gosse, 1963; Teugels and others, 1992).

20°W 0° 20°E 40°E

Mediterranean
Sea
co
oc
or
M

Algeria Libya
Egypt
Saudi
Arabia

Re
20°N
AFRICA Chad
d
Senegal Se
a
Gambia Sudan
Burkina Nigeria
EXPLANATION Guinea
Sierra Leone Ivory
Central Ethopia
DISTRIBUTION OF Coast
African Republic
Liberia
Parachanna obscura
Ghana Cameroon Dem.
Native range Togo Benin Rep.
0° of the
Equator Gabon Congo
0 1,000 MILES Congo Tanzania Indian
Atlantic Ocean
0 1,000 KILOMETERS Ocean
ue

Angola Zambia
biq
zam

Nambia
20°S
Mo

Madagascar

South
Africa

Parachanna obscura

126
 ACKNOWLEDGMENTS
We are most grateful to a large number of individuals who gener-
ously provided assistance to us in preparing this species synopsis and risk
assessment. In particular, we thank Ralf Britz of the University of Tübingen,
Germany, currently on sabbatical leave at the Division of Fishes, National
Museum of Natural History, Washington, D.C.; Maurice Kottelat, Cornol,
Switzerland; Sven Kullander, Swedish Museum of Natural History, Stock-
holm; Prachya Musikasinthorn, Faculty of Fisheries, Kasetsart University, Bangkok,
Thailand; Peter K. L. Ng, Department of Zoology and Raffles Museum, National
University of Singapore; Rohan Pethiyagoda, World Heritage Trust, Columbo, Sri
Lanka; the late Guy Teugels, Musée Royale de l’Afrique Central, Tervuren, Belgium;
and Hiroshi Ueda, Field Science Center for Northern Biosphere, Hokkaido University,
Sapporo, Japan, for providing helpful information on snakehead taxonomy, distribu-
tion, and ecology; Nina Bogutskaya of the Russian Academy of Sciences, St. Peters-
burg, for records, literature, and field collection information on channids in Russia and
the former Soviet Union; Fang Fang, Swedish Museum of Natural History, and Ping
Zhuang, Chinese Academy of Fishery Sciences, Shanghai, for their comments on
snakehead culture in China; Jim Cambray, Albany Museum, Grahamstown, South
Africa; Karsten Hartel and James Stephen Lee, Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts; Robert Howells, Texas Parks and
Wildlife Department, Heart of the Hills Research Station, Ingram, Texas; Paul
Loiselle, New York Aquarium, Brooklyn, New York; Ted Pietsch and Margarita
Reimer, School of Aquatic and Fishery Science, University of Washington, Seattle,
Washington; Richard Rosenblatt, Scripps Institute of Oceanography, La Jolla, Califor-
nia; Paul Shafland, Florida Fish and Wildlife Conservation Commission, Boca Raton,
Florida; Leo Smith, American Museum of Natural History, New York, New York;
Franklin (Buck) Snelson, Department of Biology, University of Central Florida,
Orlando, Florida; Ross B. Socolof, Bradenton, Florida; Wayne Starnes, North Caro-
lina State Museum of Natural Sciences, Raleigh, North Carolina; John Sunada, Cali-
fornia Department of Fish and Game, San Bernardino, California; Camm Swift,
Arcadia, California; and Michael Williams, U.S. Fish and Wildlife Service, Blaine,
Washington, for input on specimens collected or confiscated and/or general informa-
tion; and Gerald R. Allen, Western Australian Museum, Perth; Robert K. Hamilton,
Baltimore Sun, Baltimore, Maryland; Phil Hastings, Scripps Institute of Oceanogra-
phy; Prachya Musikasinthorn; Peter Ng; Rohan Pethiyagoda; Heok Hui Tan, National
University of Singapore; the late Guy Teugels; Mark Sabaj, Academy of Natural
Sciences, Philadelphia, Pennsylvania; and Jörg Vierke, Husum, Germany, for permis-
sion to examine and/or use photographs and illustrations. Without their help, this
effort would not have been possible.

Special thanks to Disney Publishing Worldwide, Disney Enterprises, Inc.,

for permission to use the illustrations in figure 1, and to Jean-Francois Helias of
Fishing Adventures Thailand, for use of photographs of snakeheads caught by anglers
in Thailand and Malaysia. Tom Darden, Governor’s office, Annapolis, Maryland, also
provided several photos.

127
 We also acknowledge Steve Early and Bob Lunsford, Maryland Department
of Natural Resources, Annapolis, Maryland, for keeping us advised on the northern
snakehead introduction and eradication in Crofton, Anne Arundel County.
Our thanks to many others for their assistance in providing information,
including David Casey, Kentucky Department of Fish and Wildlife, Frankfort,
Kentucky; Vi Catrow, USGS, Leetown, West Virginia; Bill Chang, National Science
Foundation, Washington, D.C.; Domingo Cravalho, Jr., Hawaii Department of
Agriculture, Honolulu, Hawaii; Richard Feeney, Los Angleles County Museum, Los
Angeles, California; Bill Foreman, Connecticut Department of Environmental
Protection, Inland Fisheries Division, Hartford, Connecticut; Pam Fuller, USGS,
Gainesville, Florida; Bob Howells; Brett Houdyshell, Kansas Department of Wildlife
and Parks, Pratt, Kansas; Keith Johnson, Idaho Department of Game and Fish, Boise,
Idaho; Deborah Koo, Canadian Food Inspection Agency, Vancouver, British
Columbia; John McCosker, California Academy of Sciences, San Francisco,
California; Tom Nesler, Colorado Division of Wildlife, Denver, Colorado; Jim
Peterson, Montana Department of Fish, Wildlife, and Parks, Great Falls, Montana;
Tyson Roberts, California Academy of Sciences; Robert H. Robins, Florida Museum
of Natural History, Gainesville, Florida; Beth Rogers, Maryland Department of
Natural Resources; Dal Schaefer, Colorado Division of Wildlife, Denver, Colorado;
Karl J. Scheidegger and Joseph Hennessy, Wisconsin Department of Natural
Resources, Madison, Wisconsin; Russell Wong, North Carolina Wildlife Resources
Commission, Raleigh, North Carolina; and Mike Yamamoto, Hawaii Department of
Land and Natural Resources, Division of Aquatic Resources, Honolulu, Hawaii.
Special thanks go to Leo Nico, USGS, Gainesville, Florida, for providing
several important references, hand-carrying snakehead specimens to us from the Bernice P.
Bishop Museum, Honolulu, Hawaii, and for his encouragement. For loans of preserved
specimens, we thank David Catania, California Academy of Sciences; Barry Chernoff,
Field Museum of Natural History, Chicago, Illinois; Ron Englund, Bernice P. Bishop
Museum; Karsten Hartel; and Mel Stiassny, American Museum of Natural History.
Support for this research was provided by Interagency Grant Agreement
94400-1-0100 from the U.S. Fish and Wildlife Service, Division of Scientific
Authority and Fisheries Management and Division of Environmental Quality, Branch
of Invasive Species, Washington, D.C. Pam Hall, Sharon Gross, and Kari Duncan,
U.S. Fish and Wildlife Service, provided snakehead import data and information of
Service policy on non-native fishes. Pingfu Chen, Natural History Museum,
University of Kansas, Lawrence, Kansas, assisted with translation of some Chinese
literature. Ken Sulak, USGS, Gainesville, Florida, and James Joeriman, Department
of Romance Languages and Literature, University of Florida, Gainesville, Florida,
provided translation assistance of some Russian literature. Sherry L. Bostick, USGS,
Gainesville, Florida, assisted greatly with illustrations, manuscript assembly, and
mapping modification; and Britton Wilson, U.S. National Park Service, Homestead,
Florida, prepared the basic distribution maps when she was with the USGS in
Gainesville. Julie Mounts, Division of Fishes, National Museum of Natural History,
Washington, D.C., assisted in obtaining original descriptions of several species of
snakeheads. Susan Trammell, Archer, Florida, created the illustration of the northern
snakehead that appears on the cover.

128
 Finally, we thank the members of the Risk Assessment and Management
Committee of the Aquatic Nuisance Species Task Force for their review and
comments on an earlier version of the manuscript: Richard Orr, Chairperson,
U.S. Department of Agriculture Animal and Plant Health Inspection Service,
Riverdale, Maryland; Jim Andreasen, U.S. Environmental Protection Agency,
Washington, D.C.; Shawn Alam, U.S. Fish and Wildlife Service, Arlington, Virginia;
Fred Kern, National Oceanic and Atmospheric Administration National Ocean
Service, Oxford, Maryland; Marshall Myers, Pet Industry Joint Advisory Council,
Washington, D.C.; Edwin Theriot, U.S. Army Corps of Engineers, Waterways
Experimental Station, Vicksburg, Mississippi; and Paul Zajicek, Florida Department
of Agriculture and Consumer Services, Tallahassee, Florida. Our gratitude is also
extended to Ralf Britz, Carter R. Gilbert, University of Florida, Robert Howells, Peter
Ng, and Hoek Hui Tan for their careful evaluation and comments on the manuscript.
Our sincere apologies to those we may have overlooked.

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