Frontispiece. Sphyraena barracuda, U.S.N.M. No. 14978, from Florida.

From
George B. Goode, “The fisheries and fishery industries of the United States,” 1884,
Section 1, Plate 178. Drawing by H. L. Todd. Cover drawing by Craig Phillips.
 Studies in Tropical Oceanography No. 1
Institute o f Marine and Atmospheric Sciences
University o f Miami

Systematics and Life History of the

Great Barracuda
Sphyraena barracuda (Walbaum)

by
D o n a ld P. d e S y lv a

University of Miami Press

Coral Gables, Florida
 This paper may be referred to as:
Stud. trop. Oceanogr. Miami, 1: viii + 1 7 9 pp.,
32 Tables, 36 Figures, October, 1963

Second printing, 1970

Printed in the United States of America

 C ontents

A bstract ................................. •........................................................................................ 1

I n t r o d u c t i o n ............................................................ ...................................................... 2
A cknow ledgm ents .....................................................................................,............ 3
S y s t e m a t i c s ...................................................................................................................... 5
Phylogenetic Relationships of the Family S p h y ra e n id a e ................. 5
Fossil Distribution of the Family S p h y ra e n id a e ..................................... 5
Recent D is tr ib u tio n ........................................................................................ 6
Delineation of the Genus S p h y r a e n a ............................................................... 7
S y n o n y m y o f Sphyraena b a rra c u d a ............................................................. 7
M ethods of M easuring and C o u n tin g ........................................................ 22
Comparisons .................................................................................................... 24
Y o u n g S ta g e s o f t h e F a m il y S p h y r a e n id a e ............................................ 29
K ey to S p e c ie s o f S p h y r a e n id a e f o r s p e c i m e n s o f 5 .5 to 1 1 .9
MM FROM THE WESTERN NORTH A T L A N T IC ............................................... 31
K e y to S p e c ie s o f S p h y r a e n id a e fo r s p e c i m e n s o f 1 2 .3 to 1 3 0
MM FROM THE WESTERN NORTH A T L A N T IC ................................. .. 33
K ey to S p e c ie s o f S p h y r a e n id a e fo r s p e c i m e n s l o n g e r t h a n 1 3 0
MM FROM THE WESTERN NORTH A T L A N T IC ............................................ 34
P o s t l a r v a l D e v e l o p m e n t o f Sphyraena b a r ra c u d a ........................... 35
D is c u s s io n .................................. ...................................................................... 42
Description and Size of the A d u l t s .......................................................... 42
R e p r o d u c t io n ................................................................................................................ 43
Sexual dimorphism ........................................................................................ 45
Criteria for Stages of Sexual D e v e lo p m e n t......................................... 47
A ge and Size at M a tu r ity .............................................................................. 47
Sex R a t i o ........................................................................................................... 49
Spawning P e r i o d ............................................................................................ 49
Frequency and Extent of S p a w n in g ........................................................ 51
F e c u n d ity ........................................................................................................... 53
E g g s .....................................................................................................................' 53
Distribution of the Postlarvae in Relation to Hydrographic and
M eteorological C o n d itio n s...................................................................... 54

v
 L e n g t h - W e ig h t R e l a t i o n .................................................................................. 57
A ge a n d G r o w t h ........................................................................................................ 59
Cause of Annulus F orm ation ............................................ .................................. 59
Methods of Scale Collection and A n a ly sis .............. ..................................... 59
Validity of the Annulus as a Y ea r-M a rk ...................... • • • • ................... 61
Criteria for the Interpretation of Annual M a r k s ....................................... 61
False A n n u li ................................................................................................................ 64
“Skipped” A n n u li ..................................................................................................... 64
Growth of the Y o u n g ................................................................................................ 65
Empirical Growth Rate in A d u lts ..................................................................... 66
Calculated Growth R a te s ...................................... . . . ......................................... 69
Body-scale Relationship ........................................................................................ 69
C alculations .................................................................................................. * . . . . . 72
Calculated Growth In crem en ts .......................................................................... 74
Length-frequency D istribu tion s .......................................................................... 74
M o r t a l it y a n d S u r v i v a l ........................................................................................ 80
L a n d in g s o f B a r r a c u d a .......................................................................................... 83
M i g r a t i o n s ......................................................................................................................... 86
E c o l o g y .............................................................................................................................. 92
Pollution ............................................ . ......................................................................... 92
Salinity .......................................................................................................................... 92
Depth .............................................................................................................................. 93
Temperature ..................................................................................................... 93
Ecology of Young B arracu da ............................................................................. 94
Ecology of Adult B arracu da ................... ................................................ .. 95
Associates and P red a to rs ....................................................................................... 96
B e h a v io r ................... ....................................................................................................... 99
Aggregating B e h a v io r ............................................................................................. 99
Peck O r d e r ................................................. .. ............................................................. 101
F e e d in g H a b i t s .............................................................................................................. 102
F o o d H a b it s ................................................................................................................... 103

Methods ........................... ........................................... ................................................. 103

Number of Items of Food per S to m a ch ......................................................... 104

vi

«
 F ood of Young from F lo r id a .................................................................... 104
Food of Young from Bim ini, B a h a m a s ................................................... 106
Food of Young from A dditional A r e a s .................................. ................ 10 9
Food of A du lts from F lo r id a .................................................................... 110
Food of A dults from Bimini, B a h a m a s ................................................... 112
Comparisons and Discussion .................................................................... 113
A t t a c k s b y B a r r a c u d a o n H u m a n s ............................................................... 121
Literature R eview of A tta c k s ....................................................................... 122
Possible Causes of A t t a c k s ....................................................................... 126
P o is o n in g o f H u m a n s b y B a r r a c u d a ............................................................ 128
General R eview of Poisonous F is h e s ........................................................ 12 8
Characteristics of Poisonous B a rra c u d a ................................................... 128
Toxin from Barracuda F le s h ....................................................................... 133
Etiology of Fish P o is o n in g ......................................................................... 133
Relation to S p a w n in g ................................................................................... 134
Relation to D i e t ............................................................................................ 13 5
Toxis Organisms in the Food Web of the B a rra c u d a ........................... 135
F ood H a b it s a n d E c o lo g ic a l R e l a t io n s o f O r g a n is m s in
the F o od W eb o f t h e B a r r a c u d a ............................................................... 138
Geographical Distribution of Toxic B a rra c u d a ..................................... 13 9
Relation of Toxic Organisms to Hydrographic and M eteorological
C o n d itio n s .................................................................................................... 1 4 0
C o n c l u s io n s ...................... .............................................................. ........................ .... 152
L it e r a t u r e C i t e d ........................................................................................................ 153
 SYSTEMATICS AND LIFE HISTORY OF
T H E GREAT BARRACUDA,
SP H Y R A E N A BARRAC UD A (WALBAUM) 1
DONALD P. d e SYLVA
Institute of Marine Science
University of Miami

A bstr ac t

The great barracuda, Sphyraena barracuda (Walbaum), family Sphy-

raenidae, is found in all tropical seas, with the exception of the eastern
Pacific. The following nominal species are considered as synonyms of
barracuda Walbaum, 1792: picuda Bloch and Schneider, 1801; becuna
Lacepede, 1803; barracuda Shaw, 1804; barracuda Cuvier, 1829; com-
mersoni Cuvier, 1829; dussumieri Cuvier, 1829; agam Riippell, 1835;
affinis Riippell, 1835; nuageuse Lienard, 1843; picuda Poey, 1866; snod-
grassi Jenkins, 1901; and akerstromi Whitley, 1947. S. kadenar Montrou-
zier, 1857, and microps Marshall, 1953, are probably synonyms.
Barracuda were collected off Miami, Florida, from Bimini, Bahamas,
and specimens from the Danish Oceanographical Expeditions in the west­
ern Atlantic were used. Ontogenetic variation in proportional measure­
ments was determined for 171 specimens from 5.5 to 1111 mm standard
length. Keys for identification are presented for barracuda, guachancho,
picudilla, borealis, and sphyraena.
Most males mature at two years and all mature by three years. Some
females mature at three years, and all mature at four. The spawning
season is from April until October off southern Florida. Spawning was
not observed but probably occurs in the open ocean. The eggs were not
identified from plankton tows. Postlarvae were taken far from shore and
near the surface, over depths of 200 to 5200 meters.
Age analyses were based upon the scale readings of 555 barracuda,
back-calculated lengths from scale measurements, and upon modes in the
length-frequency distributions of 2707 specimens from southern Florida.
It attains 10 to 12 inches at the end of the first winter, and 18 to 22 inches
at the end of its second. It reaches an age of at least 14 years. Maximum
length is about 6 feet and a weight of more than 100 pounds.
The young barracuda drifts inshore in spring, spends its first summer in
shallow nursery areas, and moves offshore to somewhat deeper water in
late fall. During the second summer it enters the mangrove habitat or the
deeper weed beds. In its third year it enters the coral-reef habitat. The
barracuda migrates seasonally along both coasts of Florida. Some indi­
viduals may drift far out to sea.
Young sometimes occur in loose aggregations, but the adults are solitary.
Peck order was observed among individuals from 12 to 16 inches long
which were confined in a tank.
l Contribution No. 220 from The Marine Laboratory, Institute of Marine Science, University
of Miami, Papers from the DANA Oceanographical Collections No. 47, and Contribution
from the Lerner Marine Laboratory, The American Museum of Natural History. Publication
of this paper through a grant from the National Science Foundation, G-24119, is grate­
fully acknowledged. This paper represents a dissertation submitted to the Faculty of the
Graduate School of Cornell University as partial fulfillment for the degree of Doctor of
Philosophy.
 Young feed upon atherinids, gobiids, and clupeids. Tetraodontiform
fishes, hemiramphids, and carangids were most commonly taken by the
adults. A change from a shallow-water habitat to that of a reef existence
is accompanied by a change in food habits.
A summary of the 29 attacks reputedly made by barracuda on humans
is presented and analyzed. Underwater experiences with barracuda are
discussed in relation to the factors which are favorable to an attack.
Poisoning in humans who have eaten fresh barracuda is due to a toxin
in the flesh and not from bacterial poisoning. In the western Atlantic,
poisonings have occurred throughout the year. There seems to be no
relation between the attainment of maturity or the spawning cycle and
the poisonous nature of barracuda flesh. Some 58 per cent of the stomachs
of adult barracuda contained fishes which are reputedly toxic, of which
20 per cent were tetraodontiform fishes. Evidence is presented for a food-
chain origin of the toxin, and mechanisms are discussed for the trans­
mission of the toxin from planktonic and benthic algae to barracuda by
way of intermediate organisms. Variations in hydrographic and meteor­
ological features are believed to result in distinct temporal and spatial
variation in the toxicity of barracuda as well as other fishes.
I n t r o d u c t io n
The barracudas, family Sphyraenidae, form a well-defined group con­
taining about 20 species, all of which belong to the genus Sphyraena. They
are wide-ranging in the tropical waters of the world. Some are found in
schools and seldom exceed 2 feet in length, while others are solitary, no­
madic species which may reach 10 feet (Biittikofer, 1890:451). Most are
used as food, and the larger species are also valued as game fishes.
Very little has been written on the biology of any of the barracudas.
Except for the two major contributions by Gudger (1918) and Walford
(1932) the content of the rather voluminous literature is limited essentially
to brief and often inadequate species listings and descriptions, and notes
on natural history. Notwithstanding the recent taxonomic revisions by
Schultz (1953), Smith (1956), and Williams (1959), the dearth of muse­
um specimens has prevented a comprehensive, world-wide taxonomic study
of this group. Because of the comparatively large size of some specimens
and the ensuing problems of preservation, many type specimens are no
longer in existence. A number of nominal species evidently are based upon
different ontogenetic stages of a single species. Adequate series represen­
tative of ontogenetic stages are seldom available, and consequently direct
comparisons of different species can seldom be made.
Most characters used to separate the species of barracudas are based
upon the number of lateral-line scales or proportional measurements
(Schultz, 1953; Smith, 1956). Since the scales are markedly deciduous,
accurate counts are often difficult to obtain. Other attributes used for
species separation, such as eye diameter, head length, maxillary lengths,
and fin lengths, show considerable ontogenetic variation. They sometimes
vary so much that different growth stadia of a species seem to represent
different species. Thus, it is clear that such proportional comparisons must
be used with discretion in the identification of specimens. Color or color
2
patterns are seldom distinctive and the numbers of fin rays or spines are
of little taxonomic value, although Williams (1959) has used color pat­
terns with success for Indian Ocean specimens. Although a world-wide
revision, while is so urgently needed in this group, must await the accumula­
tion of additional specimens, some conclusions may be drawn concerning
the present taxonomic status of some of the species to elucidate a few of
their biological problems.
Of all the sphyraenids, perhaps the most has been written about the
great barracuda, Sphyraena barracuda (Walbaum), hereafter referred to as
the barracuda. Found throughout most of the tropical and subtropical
waters of the world, it is an important food fish over much of its range.
Because of its comparatively large size and gameness, the barracuda is
sought by anglers who consider it a popular, light-tackle game fish. The
flesh of this and other sphyraenids is known to possess poisonous qualities
occasionally.
The following sections represent an attempt to collate the widely scat­
tered accounts of the barracuda supplemented with original data, and to
deduct therefrom a synthesis of the biology of this intriguing animal.

A cknow ledgm ents

The present study is an outgrowth of field work started at Pier 5,

Miami, Florida. This famous fishing dock is the center for charterboat trips
from Miami to the nearby reefs and to the Gulf Stream. It offers an excel­
lent opportunity to make observations on large, pelagic fishes which are
usually difficult to obtain. During 1952 and 1953, at the suggestion of
Prof. Luis Rene Rivas of the Department of Zoology, University of Miami,
I made frequent visits to Pier 5 to examine catches of scombroid fishes.
As a result of several outbreaks of food poisoning in man due to the
ingestion of fresh barracuda caught in Florida waters in 1954 and 1955,
the author became interested in studying the life history of the barracuda,
and in 1955 and 1956 the field work at Pier 5 was resumed. I wish to
acknowledge the complete cooperation of Mr. Dan Daniels, formerly dock-
master of Pier 5, Miami, and the numerous charterboat captains, all of
whom willingly and patiently supported the field observations. To Dr.
Edward C. Raney I am indebted for his guidance throughout the course of
the study. I also wish to thank Drs. John P. Barlow, Howard E. Evans,
and Gustav A. Swanson for many helpful suggestions and criticisms, as
well as the staff of the Depertment of Conservation, Cornell University,
for making numerous facilities available.
I wish to thank Dr. Charles M. Breder, Jr., Dr. and Mrs. Louis A.
Krumholz, presently of the University of Louisville, Miss Francesca R.
LaMonte, and Dr. Albert E. Parr for making available the facilities of
the American Museum of Natural History and the Lerner Marine Labora­
tory at Bimini, Bahamas. Drs. ‘Krumholz and Lester R. Aronson assisted
3
with seining operations. The assistance and knowledge of collecting tech-
nics of the native staff at Bimini greatly expedited the work.
Thanks are due Dr. E. Bertelsen and the late Dr. A Vedel Taning
the Carlsberg Foundation, Copenhagen, Denmark, for making available an
extensive collection of postlarval sphyraenids collected by the D A N A ex­
peditions under the leadership of the late Prof. Johannes Schmidt. Dr.
Wolfgang Klausewitz of the Senckenberg Museum at Frankfurt-am-Main,
Germany, kindly re-examined several of Riippell’s types and reported on
them in detail.
I wish to thank Dr. F. G. Walton Smith and the staff of the Institute
of Marine Science of the University of Miami for providing much of the
equipment used in this study, and for offering many useful suggestions
towards the improvement of the manuscript. Dr. John E. Randall, Univer­
sity of Puerto Rico, supplied valuable information on several barracuda
attacks and cases of poisoning.
In addition to the persons of the above-mentioned institutions, the
following individuals loaned specimens for this study: Dr. James E. Bohlke,
Academy of Natural Sciences of Philadelphia; Dr. John C. Briggs, Univer­
sity of Texas; Dr. David K. Caldwell, Los Angeles County Museum; Dr.
Bruce B. Collette, U. S. Fish and Wildlife Service; Mr. Donald S. Erdman,
Department of Agriculture and Commerce, San Juan, Puerto Rico; Dr.
Robert W. Harrington, Jr., Florida State Board of Health; Mr. Frank J.
Mather, III, W oods Hole Oceanographic Institution; Dr. Ernest A. Lach-
ner, U. S. National Museum; Dr. Giles W. Mead, Harvard University;
Prof. Luis Rene Rivas, University of Miami; Dr. Leonard P. Schultz, U. S.
National Museum; Dr. C. Lavett Smith, American Museum of Natural
History; and Dr. Royal D. Suttkus, Tulane University.
For valuable information on the incidence and toxicology of barracuda
poisoning, thanks are extended to Dr. Bruce W. Halstead, World Life
Research Institute, Colton, California; Dr. Yoshiro Hashimoto, Fisheries
Department, University of Tokyo; Dr. Masao Migita, Tokai Fisheries
Experiment Station, Tokyo; and Mr. Sidney Paetro, Broward County
(Florida) Health Department. I wish to thank the numerous victims of
barracuda poisoning who disclosed extensive information on the nature of
the poisoning and the localities where the fish were caught. Thanks are
also due to Dr. Louis F. Westbrook and his staff of the Dade County
(Florida) Health Department for bacteriological analyses of samples of
barracuda flesh.
Special appreciation is expressed to Messrs. Joseph T. Reese of Fort
Lauderdale and the late A1 Pflueger of Miami for supplying scale samples
from large barracuda and for the use of a large number of measurements
of barracuda, respectively. I wish to express my gratitude to Captain
Eddie M oore and his crew of the M /V P A N D A for their enthusiastic
support in collecting adult specimens of barracuda in the Bahamas.
The following persons have aided materially in collecting or assisting
4
the writer in collecting specimens: Dr. and Mrs. Ilias Konstantinu; Mr. and
Mrs. William H. Peterson, Jr.; Mr. and Mrs. Neil Roseman; Miss Kath­
leen Parker; Dr. Frederick A. Kalber; Dr. Allan B. Lewis; Messrs. Selwyn
J. Bein, Robert F. Burrows, Edward F. Klima, Reynolds A. Moody, Alan
Moffett, Craig Phillips, Warren F. Rathjen, James D. Regan, Durbin C.
Tabb, and Alfred V. Volpe. I wish to thank Dr. Frederick A. Kalber for
preparing the histological sections; Dr. Allan B. Lewis and Mr. Robert V.
Miller assisted in making scale impressions. Mr. Douglass M. Payne made
photomicrographs of the scales and histological sections and photographed
the illustrations. Drs. Gerald P. Cooper, University of Michigan, Louis A.
Krumholz, Ernest A. Lachner, C. Richard Robins, Gilbert L. Voss, and
F. G. Walton Smith offered valuable criticisms of the manuscript.
Finally and especially, I wish to thank my wife, Doris, for her patient
and valuable assistance during the many hours of field and laboratory work.
Sy s t e m a t ic s
Phylogenetic Relationships of the Family Sphyraenidae. — Species of the
family Sphyraenidae are characterized by their long, pointed heads^, their
pike-like appearance, and their two dorsal fins. The jaws contain well-
developed shearing teeth, and in Atlantic species there are no gill rakers.
The phylogenetic relationships of the family have been discussed in detail
by Bridge (1896), Starks (1899), Dollo (1909), Goodrich (1909),
Regan (1912), Ribeiro (1915), Jordan and Hubbs (1919), Frost (1929),
Gregory (1933), Hollister (1937), and Gosline (1962). The family is
here placed in the suborder Percesoces, together with the Mugilidae, Poly­
nemidae, and Atherinidae. Berg’s (1955) Mugiliformes is synonymous
with the Percesoces, and includes the suborder Sphyraenoidei; his order
differs from the Perciformes in the abdominal position of the pelvics and
in having the pelvic bones connected with the cleithra or postcleithra by
a ligament. Norman (1957) included the suborder Mugiloidea under the
order Percomorphi, but removed from this group the Polynemidae
which he placed in a separate suborder, the Polynemoidea, although Gos­
line (1962) has shown the Polynemidae and Sphyraenidae to be rather
close. The family Sphyraenidae represents a compact family having anato­
mical structures not greatly different from its nearest relatives.
Fossil Distribution of the Family Sphyraenidae. —Fossil sphyraenids are
widespread since the Lower Eocene and include the genera Sphyraenodus
Agassiz ( = D ictyodus Owen), Protosphyraena Leidv, Prosphyraena Pauca,
and Sphyraena Klein. Numerous fossil species of the latter genus include
the following: S. lugardi White, 1926, from the Eocene of Nigeria; 5.
fayumensis Dames, 1883, from the Tertiary of Egypt and the Eocene of
Italy (Bassani, 1899), S. malembeensis Dartevelle and Casier, 1943, from
the Lower Miocene of the Belgian Congo (see also Henriques da Silva
[1956] for a closely related form from Portugal); S. viannai Dartevelle
and Casier, 1949, from the Miocene of Celebes; S. weberi Casier, 1954,
5
from the Miocene of Java; S. (Otolithus) handsfuchsi Schubert, 1906, from
the Miocene of Voslau, Hungary; S. pannonica Weiler, 1938, from the
Middle Oligocene of Hungary; S. striata Casier, 1946, from the Ypresian
strata of Belgium; S. major Leidy, 1855, from the Pleistocene beds of
South Carolina; S. (Otolithus) oblongus Frost, 1934, from the Lower
Eocene of the London Clay; S. tyrolensis Zittel, 1890, from Austria; S.
bolcensis Agassiz, 1843, S. gracilis Agassiz, 1843, S. intermedia Wood­
ward, 1901, and S. suessi Woodward, 1901, all from the Eocene of Monte
Bolca, Italy; and S. barracuda Walbaum, 1792, from the Cretaceous beds
of London. Zittel (1890) also recorded fossil specimens of Sphyraena from
the Oligocene of Austria and Croatia, as well as teeth in “other Tertiary
formations in Europe, Egypt, Nigeria, and America.” The genus Sphy-
raenodus Leidy occurs in the Miocene of New Jersey (speciosus Leidy,
1855, and silovianus Cope, 1875), Belgium ( lerichei Casier, 1950, and
bruxelliensis Casier, 1944), and England (priscus Agassiz, 1843, and
crassidens Agassiz, 1843).
Agassiz (1843) recognized the genera Sphyraena, Sphyraenodus, and
Hypsodon, but later (1845) he removed Hypsodon from the family.
Pictet (1853) included in the Sphyraenidae the following genera: Hypso­
don, Saurocephalus, Saurodon, Pachyrhizodus, Cladocyclus, Isodus, Rham-
phognathus, and Mesogaster, most of which are Cretaceous forms. Many
of the fossil genera and species are known only from teeth, otoliths, or
jaws, as Woodward (1901) pointed out. Generic distinctions among fossil
forms are naturally difficult to evaluate in the light of such fragmentary
knowledge. However, the distribution of fossil genera and species indicates
that the family Sphyraenidae was widespread throughout the warm, shallow
waters of the Tertiary Tethys Sea (Ekman, 1953: 63-67). Agassiz’s (1843)
recognition of Sphyraena barracuda from Cretaceous beds would suggest
that this species is a relatively stable one morphologically.
Recent Distribution.— Sphyraena barracuda is presently distributed in all
tropical, Recent seas with the exception of the eastern Pacific Ocean
(Fig. 1), the great expanse of water between the Hawaiian Islands and the
western coasts of the Americas acting as an effective barrier to dispersal
(Ekman, 1953:72-77) for many tropical fishes. Although the Equatorial
Counter Current could conceivably supply a stock of the pelagic young,
evidently this expanse plus the encroachment of polar currents toward
the equator is effective in preventing any extensive colonization by western
Pacific species. The data plotted in Figure 1 include barracuda and all the
nominal species which I consider to be synonyms of barracuda.
The barracuda may occur many miles from shoal water, particularly
about the oceanic islands of the Pacific. Murphy (1914) reported this
species about 900 miles east of Bermuda in August. Subsequently, Om-
manney (1953), Murphy and Shomura (1953a, 1953b), Mather and Day
(1954), Shomura and Murphy (1955), Springer and Bullis (1956), and
6
Iversen and Yoshida (1956; 1957) have recorded barracuda over deep
water and relatively far from land. Most of these are large individuals.
Although most of these locality records have been made possible by long-
line and other offshore fishing explorations, the distribution of the barra­
cuda appears spotty due to lack of collecting in many areas, i.e., the
South Atlantic, Indian Ocean, and the northern Australian coast. Its known
distribution is within the range given by Ekman (1953:3) for the mean
location of the 20°C isotherm, which limits the poleward distribution of
tropical species. However, some barracuda evidently tolerate somewhat
lower temperatures, since Cape Cod, Massachusetts, from which speci­
mens have been taken in August, has a mean temperature of about 18.3°C
for this month (Fuglister, 1947). Also, Durban, South Africa, and Kyushu,
Japan, evidently represent the maximum poleward dispersal of barracuda
and have slightly colder temperatures than generally preferred.
Delineation of the Genus Sphyraena.— The living members of the family
Sphyraenidae are represented by a single genus, Sphyraena Klein, 1778.
Fowler (1903) proposed the subgenus Agrioposphyraena for the large-
scaled form, with 75 to 87 scales, having a flattened or slightly concave
head, with barracuda Walbaum, 1792, as the type, as opposed to the sub­
genus Sphyraena with more than 100 scales and a slightly convex head.
Smith (1956) subsequently elevated Fowler’s subgenus to full generic
rank, and described two additional genera since all species had previously
been “. . . ruthlessly pushed into the ‘catch-all’ bag of Sphyraena by other
authors.” His generic distinctions are, however, based upon what would
appear at best to be specific characters, such as the number of gill rakers
( “one or two,” as compared with “one”), length and placement of fins,
the shape of the preopercle, and the number of lateral-line scales. Yet
Smith pointed out the ontogenetic variation in the very same characters
upon which he based his distinctions. All these characters overlap when
considered on a world-wide basis, although they suffice for western Indian
Ocean sphyraenids. Whitley’s (1947) genus Australuzza, although pro­
posed for a different species ( Sphyraena novaehollandiae Gunther), also
seems to be only a case of specific differentiation; it is based upon the
position of the dorsal and pelvic fins, and upon the number of lateral-line
scales. Until a complete revision of the family is undertaken, I prefer to
retain Sphyraena as the sole generic representative of the living species
of the family Sphyraenidae.

Sy no ny m y of Sphyraena barracuda
Sphyraena barracuda was originally described as Esox barracuda in
1792 by Walbaum from Catesby’s (1743) pre-Linnaean “Umbla minor
marina maxillis longioribus” from the West Indies. Riippell (1835) showed
that Forskal (1775) had mistaken S. agam Riippell for Esox sphyraena
Linnaeus. Walbaum’s description was apparently overlooked for a number
7
 of years, and the West Indian form was again described, this time by Bloch
and Schneider in 1801 as Sphyraena sphyraena var. picuda, based upon
Parra’s (1787) non-binomial “Picuda” from Havana. Lacepede (1803)
described S. becuna from a drawing by Plumier at Martinique, which Cu­
vier (in Cuvier and Valenciennes, 1829) subsequently believed to be
identical with Parra’s “Picuda,” but he had evidently overlooked both
Walbaum’s and Bloch and Schneider’s descriptions. Shaw (1804) des­
cribed Esox barracuda after Catesby’s (1743) specimen, and Cuvier (in
Cuvier and Valenciennes, 1829) later described S. barracuda based on
Shaw’s Esox barracuda, but offered no comment; since Cuvier’s type
locality was Brazil he may have believed that because Shaw’s specimen
was not from Brazil it was necessarily a different species. Swain (1882)
identified Shaw’s barracuda with Bloch and Schneider’s picuda. Walbaum’s
original description went unnoticed until Jordan and Rutter (1897) point­
ed out that barracuda Walbaum had priority over picuda Bloch and
Schneider, 1801. It had been recognized by most authors that becuna
Lacepede, 1803 and barracuda Cuvier, 1829, were synonyms of picuda
Bloch and Schneider, 1801, although this was doubted by Ppey (1858),
and thus picuda has been in wide use since, notwithstanding Jordan and
Rutter’s (1897) statement.
Cuvier’s (in Cuvier and Valenciennes, 1829) description of S. commer-
soni was based upon Lacepede’s (1803) plate of the “variete de la Sphy-
rene Chinoise,” and has since been loosely applied to all large, Indo-Pacific
sphyraenids not referable to S. jello Cuvier. Cuvier (in Cuvier and Valen­
ciennes, 1831) described dussumieri from the Indian Ocean, which Bleeker
(1854b) suggested was identical with commersoni, and which was cor­
roborated by Day (1875-1878). In the original description, Cuvier stated
that Dussumier himself believed that the species named after him was
identical with a common species in the Red Sea, which is evidently S. bar­
racuda. Bleeker (1865) identified as S. commersoni a specimen from the
Antilles which was labeled “S. picuda” in the Leiden Museum. He com­
pared it critically with specimens from the Netherlands Indies but could
not find “the slightest difference,” and, as Weber and de Beaufort (1921)
pointed out, it is surprising that Bleeker did not place commersoni in the
synonymy of his “S. picuda.” Since Bleeker (1854b) had previously stated
that commersoni and dussumieri might be identical, and since dussumieri
was later shown to be the same as picuda by Weber and de Beaufort
(1922), it seems strange that this point has been overlooked.
The identity of commersoni with barracuda had been proposed by
Fowler (1935b) and Herre (1931, et seq.). Fowler had earlier (1928)
suspected that S. commersoni, Kner (1879), as well as S. commersoni,
Jenkins (1901) were actually barracuda Walbaum. Bleeker (1854b)
further compared commersoni with barracuda; the latter is supposedly
distinguishable by the lack of vomerine teeth, as well as by having the
[translation] “head 3Vi times the body length, the height of the head 3
times in the length, and the height of the body 7 or more times in the
length,” all of which might be attributed to ontogenetic changes. All
specimens of barracuda I have examined from the tropical western Atlantic
have fine teeth on the vomer.
Schultz (1953) tentatively placed com m ersoni in synonymy with S.
chinensis Lacepede, 1803, but com m ersoni , as understood from Cuvier’s
(1829) description of Lacepede’s Plate 8, Figure 3 ( “Variete de la
Sphyrene Chinoise”) has the posterior margin of the preopercle obtusely
rounded, and not, as Schultz asserted, produced and forming a somewhat
membranous lobe. Smith (1956) considered commersoni to be a probable
synonym of fiavicauda Rtippell, 1835, yet in Smith’s description of
fiavicauda he stated that it has a distinct skinny flap at the angle of the
opercle. Lacepede’s (1803) figure clearly shows that the preopercle is
rounded and not produced, and Cuvier’s description, based on Commer-
son’s fish, clearly stated that the preopercle is not salient and expanded
but is rounded. Hence, comm ersoni cannot be a synonym of either
chinensis or fiavicauda. Possibly Weber and de Beaufort (1922) had
recognized this since they included comm ersoni in the synonymy of
“S. picuda” { —S. barracuda).
Cuvier {in Cuvier and Valenciennes, 1829) also stated that in com m er­
soni the opercle ends in two spinous points, whereas in barracuda it ends
in one. I have found this character to be variable in S. barracuda from the
tropical western Atlantic, being subject to ontogenetic as well as individual
variation, and care must be exercised not to overlook the lower point
which may be concealed by a fleshy flap.
Smith (1956) observed that the white tips of the fins are characteristic
of barracuda. This led Bleeker (1854b), in speaking of commersoni, to
state [translation]: “In its violet caudal, anal, and dorsal fin with white
tips it makes me think of Sphyraena agam Riippell, and in actuality is a
related species, which, however, is easily recognized by its thrice cut-out
caudal fin, which has a quadrilobed appearance.” Compare this also with
Riippell’s (1835) description of the caudal fin of agam which has “an
diesem Rande bildem drei halbrunde Ausschnitte vier Spitzen . . . .” The
white tips mentioned by Bleeker are characteristic of barracuda from the
West Indies, Africa (Smith, 1956; Williams, 1956), the Red Sea (USNM
147455), and the Hawaiian Islands (Jenkins, 1901), although this char­
acter is subject to variation depending upon the size of the specimen. The
quadrilobed tail is only obvious in large specimens, but is distinctive of
barracuda (Smith, 1956, Fig. 4b, PI. I; Williams, 1956, PI. II, lower
figure), as well as jello.
Klunzinger’s (1884) specimen identified as agam, from the Red Sea,
was thought to be different from agam Riippell by Schultz (1953). Jordan
and Evermann (1905) doubted the identification of Steindachner’s (1901)
specimen as agam Riippell from Honolulu, but failed to give the reasons
for their decision. Schultz (1953) separated agam from barracuda essen­
9
tially on the lateral-line pore count, barracuda having 76 to 85, while
agam has “more than 100” pores. Smith (1956), apparently on Klun-
zinger’s (1870) statement that agam has 85 scales, as well as Weber and
de Beaufort’s (1922) inclusion of agam in the synonymy of “S. picuda”
considered that Schultz’s retention of agam as a distinct species was in
error, and believed that in Riippell’s illustration of the type of agam the
scaling is conventional and does not represent the true scale count. Dr.
Wolfgang Klausewitz has kindly examined the type of Ruppell’s agam
in the Senckenberg Museum and informs me that it has 81 lateral-line
pores; on the basis of this and other characters, it cannot be separated
from barracuda.
Sphyraena affinis Riippell, 1835, is a young specimen of agam, accord­
ing to Klunzinger’s (1870) comparison of the two types in the Sencken­
berg Museum. I examined Ruppell’s type of affinis and find it to be
inseparable from barracuda.
I have not seen the original description of S. nuageuse Lienard, 1843,
but on the authority of Sauvage (1891) that it is a synonym of agam
( = barracuda) I am placing it in the synonymy of barracuda. Sauvage
quoted Lienard’s description, but it is not possible to ascertain if the entire
description is cited. Lienard’s description of the size of nuageuse, to 5
feet, and the coloration, with blackish spots about the lateral line, the
dusky vertical fins, the black opercular membrane and base of the
pectoral fin all are charcteristic of barracuda, and, in combination with
other characters, point to the identity of nuageuse with barracuda.
S. kadenar Montrouzier, 1857, is difficult to identify from the meager
description, but the vertical bands on the back, the only distinctive part
of the description which separates it from any other sphyraenid, might pos­
sibly identify it with barracuda, although this is also characteristic of jello
Cuvier.
Poey (1864b) redescribed Bloch and Schneider’s picuda, 1801, since
he believed Bloch and Schneider were mistaken when they considered it
to be only a variety of Esox sphyraena Linnaeus, and that it was not
permissible to give proper names to varieties. Poey felt that his previous
recognition of the species (1858) and comparison of it with S. becuna
Lacepede entitled him to describe the species as new. He also mentioned
that he was thus justified since Bloqh and Schneider added nothing to
Parra’s (1787) figure. Whether or not Poey is correct in making these
assumptions is irrelevant, since picuda Bloch and Schneider is antedated
by Walbaum’s (1792) description.
Jenkins’s (1901) original description of S. snodgrassi noted that “the
tips of the second dorsal and caudal fins [are] white” and that this species
is “. . . most closely related to Sphyraena commersonii . . .” but differs
in being “. . . less slender, in having a somewhat longer pectoral fin, and
in having the black blotches of the soft dorsal and anal fins.” The black
blotches, however, are typical of “commersoni” ( = 5 . barracuda) as
10
F ig u r e 1. Distribution of the barracuda, Sphyraena barracuda (Walbaun). Each dot represents a published record of this species or a name considered as a synonym.
figured in the excellent photograph by Williams (1956). The pectoral fin
length is an allometric character, and slenderness essentially depends
upon the physiological condition of the animal and thus cannot always
be considered diagnostic for taxonomic purposes. Subsequently, Jenkins
(1904) believed his type specimen to be identical with commersoni,
although Jordan and Snyder (1907) stated that S. snodgrassi was “prob­
ably not the same as the East Indian Sphyraena com m ersoni ” Jordan
and Richardson (1908) described a 6-inch specimen of commersoni from
Aparri in the Philippines as “agreeing with S. snodgrassi in every respect
except that there is a slight tentacle at the tip of the chin.” This character
is somewhat subject to individual variation according to my observations,
but is more developed in small individuals. They added that “from S.
commersoni, as understood by Gunther [1860] and Day [1875-1878],
this specimen differs in its shorter maxillary”; this character also varies
ontogenetically, and is not diagnostic unless specimens of comparable size
are used. Schultz (1943) gave an adequate description of “snodgrassi”
from Samoa, which he believed to be possibly identical with barracuda.
Subsequently, Schultz (1953) tentatively considered the Pacific form as
identical with the West Indian form, stating that a “careful comparison
is needed” between the two. I have examined Jenkins’s type (USNM
49693) and have compared it with barracuda of comparable size from
the West Indies, and can find no differences between them.
Whitley ( 1 9 4 7 ) described S. akerstrom i from a specimen 1 4 4 5 mm in
total length from Lowendal Island, Western Australia. He stated that his
specimen, when keyed out, “comes down to the West Indian Barracuda
(S. picuda) in Weber and de Beaufort [1 9 2 2 ] . . . but differs very much
in the size of the eye and its ratios to other parts of the head, the
ventrals are much shorter than the pectorals, and the outline of the
caudal fin is different.” Although Weber and de Beaufort ( 1 9 2 2 ) do
not state the size range forming the basis for their description, they give
the eye size for “S. picuda” ( = S . barracuda) as 6 .0 to 6 .4 in head; my
own measurements on barracuda from the tropical western Atlantic show
a range of from 6 .0 to 9 .2 , indicating that for these specimens the range
is much greater than given by these authors for the Indo-Pacific specimens.
Although my largest Atlantic specimen measured 1 1 1 1 mm standard
length, perhaps larger individuals were used by Weber and de Beaufort
( 1 9 2 2 ) . The eye of Whitley’s ( 1 9 4 7 ) type of akerstrom i ( 1 2 1 0 mm) was
contained 7 .8 times in the head ( 4 .5 per cent of the standard length), and
does not differ markedly from that found for barracuda of comparable
size (Fig. 2 1 ) . The remaining characters on which akerstrom i is based
appear to vary with the size of the fish. Specimens collected in nearby
localities on subsequent dates and assigned to akerstrom i by Whitley may
represent different species, possibly jello, according to his description
of two of these. He stated that akerstrom i has 7 6 scales in the lateral
line, yet two specimens evidently included as paratypes, although this
13
is not stated, are said to have 115 scales and 120 scales, respectively.
Although Whitley considered this to be “extraordinary variation,” it
would seem that such counts are considerably removed from what is
known about the infraspecific variation for a given species of sphyraenid.
Unfortunately, descriptions of the teeth were not included for the two
aberrant specimens, and thus comparisons with jello are not possible.
Whitley’s drawing (1947, PI. 11, Fig. 1) is probably of barracuda. Both
Schultz (1953) and Smith (1956) have included it in their synonymies
of barracuda.
Marshall (1953) described S. microps from Queensland based upon
what is evidently a large (1614 mm total length) specimen of barracuda.
Although standard length was not given by him, I converted total length
to standard length. Subsequent calculation of the percentage of eye
diameter in standard length indicates that the specimen falls well within
the range of barracuda (Fig. 21). However, since Marshall made no
comparisons with other sphyraenids it is thus not possible to ascertain
upon what distinguishing characters his type was based, although his
description appears to fit barracuda well. Marshall evidently made the
same error as Whitley (1947) in subsequently describing a paratype which
was apparently a species different from the type. This seemingly caused
Smith (1956) to place microps in the synonymy of barracuda, since the
scale count given by Marshall (1953) is for the second fish only.
However, there is no reason to believe that the scale count for this second
specimen is the same as for the type. Marshall’s lateral-line scale count
of 116 scales on the paratype is far too high for barracuda or its previously
discussed synonyms. He concluded that “in the first specimen there was
probably some mutilation for in the second fish the caudal had the middle
rays somewhat produced as in S. jello” In such a large specimen it would
be expected that these middle rays would be produced, but Marshall’s
statement that there was mutilation of the caudal fin of the type would
explain this incongruity.
The synonymy presented here for Sphyraena barracuda includes only
those references which are essentially taxonomic and distributional in
nature, and does not necessarily include references on ecology, food,
ichthyosarcotoxism, attacks, etc., which are treated elsewhere in this paper.

Genus Sphyraena Klein

Sphyraena-Klein, 1778:464 (ex Artedi, based on Esox sphyraena Linnaeus).
Sphyraena-Rose, in Artedi Piscium, 1793:112 (based on Esox sphyraena
Linnaeus).
Sphyraena-Bloch and Schneider, 1801:109 (b ased on Esox sphyraena
Linnaeus).
A cws-Lacepede, 1803:327 (ex Plumier, MS).
Sphyroena-Dumeril, 1806:148, 342 (different spelling based on Bloch and
Schneider, 1801:109-110).
Sphyrena-Rafinesque, 1810:34 (different spelling based on B lo ch and
Schneider, 1801).
14
 Agrioposphyraena-Fowlev, 1903:749 (new subgenus, based on Esox barra­
cuda Walbaum, 1792). Smith, 1956:41 (elevated to generic status, after
Fowler, 1903).
Australuzza-Whitley, 147:131 (Orthotype for Sphyraena novaehollandiae
Gunther).
Sphyraenella-Smith, 1956:38 (new genus for Sphyraena flavicauda Riippell,
1835).
Indosphyraena-Smith, 1956:39 (new genus for Sphyraena africana Gilchrist
and Thompson, 1909).
Callosphyraena-Smith, 1956:42 (new genus for Sphyraena toxeuma Fowler,
1903).

Pre-Linnaean Names
Allualu or brochet-Vlaming, 1715 (6) (Indian Seas). Valentijn, 1724 (70)
(based on Vlaming). Renard, 1754, Table 40, Fig. 202 (copied from Vlaming).
Umbla minor, etc. (the barracuda)-Catesby, 1743: 1, PI. 1 (Bahamas; size,
description, voracity, poisonous qualities).

Post-Linnaean Names
Esox sphyraena-Forskkl, 1775: 16 (Zanzibar; misidentification).
Picuda-Parra, 1787: 90; Table 35, Fig. 2 (Havana; food value, poisonous
qualities).
Esox barracuda-Walbaum, 1792: 93-94 (original description after Catesby;
type locality: West Indies).
Sphyraena sphyraena var. picuda-Bloch and Schneider, 1801: 109-110, Fig.
1; Table 29 (original description; type locality: “in America australi,” after
Parra, 1787).
Variete de la Sphyrene Chinoise-Lacepede, 1803: PI. 8, Fig. 3 (original
description; type locality: Indian Ocean). Cloquet, 1827: 233 (considered the
same as Sphyraena spet Linnaeus).
Sphyraena becuna-Lacepede, 1803: 325, PI. 9, Fig. 3 (original description
after drawing by Plumier at Martinique). Shaw, 1804: 112, PI. 109, bottom
( “Becuna pike”; description from Lacepede after Plumier, MS). Cuvier in
Cuvier and Valenciennes, 1829: 340-341 (Antilles, coast of Brazil; description;
believed it to be the same as the Picuda of Parra [1787]; size, poisonous quali­
ties). Cuvier in Cuvier and Valenciennes, 1831: 507 (Gorea, Africa). Castel-
nau, 1855: 6 ( “Sphyraena becuna Schneider” non Lacepede; Rio de Janeiro,
Brazil; coloration). Poey, 1858: 398 (same as S. picuda). Poey, 1868: 275
(similar to S. vulgaris Cuvier).
Esox barracuda-Shaw, 1804: 105-106 (original description based on Catesby
[1743]; type locality: Carolinas; voracity, alleged danger to bathers, distribution
in West Indies, poisonous qualities). Castelnau, 1855: 6 (Bahia, Brazil; com­
mon name: “Solteira”; description). Poey, 1868: 277 (Havana, Cuba).
Nichols and Breder, 1927: 72 (Long Island, New York; description, distribu­
tion, size).
Sphyraena barracuda Cuvier in Cuvier and Valenciennes, 1829: 342-349
(original description based on Esox barracuda Shaw [no comment]; type local­
ity: Brazil; common names: “becune,” “picuda”; also recorded from Puerto
Rico; brief description of internal organs and skeleton, voracity, poisonous
qualities, size). Cope, 1871: 472 (St. Martin’s; counts, coloration, comparison
with Poey’s [1858] description of picuda; disagrees with Gunther’s [1860]
description).
Sphyraena commersonii Cuvier in Cuvier and Valenciennes, 1829: 352-353
(original description based on a drawing by Lacepede, the “Variete de la
Sphyrene Chinoise,” 1803: PI. 8, Fig. 3, from the Indian Ocean). Bleeker,
15
1849a: 66 (Makassar, Celebes). Bleeker, 1849b: 55-56 (Batavia, Samarang,
Surabaya, Java, Kammel, Madura Islands, Ternate, Sindangole, Halmaheira
Islands). Bleeker, 1854b; 6, 7, 16 (Batavia, Samarang, Surabaya, Java, Kam­
mel, Madura Islands, Ternate, Sindangole, Halmaheira Islands; description,
coloration, range, comparisons with other species; suggested that S. commer­
soni and S. dussumieri may be identical). Gunther, 1860: 338 ( “Javanese Sea;
Molucca Sea; Hindostan”; description, synonymy). Castelnau, 1861: 7 (South
Africa). Bleeker, 1865: 265-266 (identification of a specimen labelled “S.
picuda” from the Antilles as Sphyraena commersonii). Castelnau, 1873: 102
(Knob Island, North Australia). MacLeay, 1881: 33 (citation of Castelnau’s
[1873] reference). MacLeay, 1882: 33-34 (Knob Island, North Australia,
after Castelnau [1873]; description, counts, coloration). Kner, 1879: 44 (Viti
Islands, Oceania). Day, 1875-1878: 342-343 ( “Seas of India to the Malay
Archipelago and beyond”; counts; identity of Bleeker’s specimens of S. com­
mersoni with S. dussumieri in the British Museum; synonymy, description,
coloration, food value). Day, 1889: 335-337 (Indian Seas; description, counts,
coloration, habitat). Sauvage, 1891:522 (Madagascar; common name). Saville-
Kent, 1893: 293 (Thursday Island, Queensland, Australia; range, size).
Fowler, 1900: 501, 520 ( “Sandwich Islands”; Tahiti). Snyder, 1904: 523
(Hawaiian Islands). Jordan and Richardson, 1908: 245 (a 6-inch specimen “from
Aparri [Philippines] agreeing with S. snodgrassi, from Honolulu, Hawaii, in
every aspect except that there is a slight tentacle at the tip of the chin”; counts,
measurements, ambiguity of Cuvier’s original description). Gilchrist and
Thompson, 1909: 255-256 (Durban, South Africa). Snyder, 1912: 495
(Okinawa). Weber, 1913: 150-152 (Makassar fish market and Kangeang
Island, Dutch East Indies). Maxwell, 1921: 203 (Malaya). Fowler, 1922: 82
(Hawaii). Barnard, 1925: 312-316 (South Africa; description, synonymy,
distribution, common names; S. snodgrassi a synonym). Whitley, 1927: 4 (Fiji).
Shmidt, 1930: 2 (Riukyu Islands). Pinter, 1930: 500 (Ceylon; cestode
parasite). Fowler, 1931: 325 (S. commersoni, Schmeltz [= Kner], 1879, not
of Cuvier). Ommanney, 1953: 66-67 (Amirante Bank, Seychelles; breeding
season, food, common name). Williams, 1956: 16-18, PI. II, bottom (East
Africa; synonymy, common names, distribution in East Africa, description,
color, size, seasonal occurrence, ecology, spawning, food, potential fishery,
migrations).
Sphyraena Dussumieri-Quvier in Cuvier and Valenciennes, 1831: 508-510,
Fig. 67 (original description; type locality: Maldive Islands, Indian Ocean, and
eastern coast of Africa; food value; Dussumier believed it to be identical with
a common species of the Red Sea). Gunther, 1860: 339 (lie de France, Indian
Ocean; description). Sauvage, 1891: 522 (Madagascar, common name).
Sphyraena agam-Riippell, 1835: 99-100, PI. 25, Fig. 2 (original description;
type locality: Red Sea; food value, size, Arabic name: “agam”; S. sphyraena,
Forskal 1775, a synonym). Gunther, 1860: 341 (Red Sea; synonymy). Playfair,
1866: 78 (Zanzibar and Red Sea, based on Esox sphyraena, Forskal, 1775, not
of Linnaeus). Klunzinger, 1870: 822 (Red Sea). Klunzinger, 1884: 128-129
(Red Sea; dangerous to man). Schultz, 1953: 281 (S. agam Klunzinger, 1884,
not of Riippell, 1835). Sauvage, 1891: 410-414 (Madagascar; S. nuageuse
Lienard, 1843, a synonym; common name). Steindachner, 1901: 500 (Hono­
lulu). Jordan and Evermann, 1905: 143 (S. agam, Steindachner, 1901, not of
Riippell, 1835).
Sphyraena affinis-Riippell, 1835: 98-99 (original description; type locality:
Red Sea). Klunzinger, 1870: 822 (S. affinis Riippell a synonym of S. agam
Riippell, 1835).
Sphyraena picuda-Richardson, 1836: 5, 32, 34 (b ased on B lo ch and
Schneider, 1801; “Gulf of Mexico, sea of Brazil, and at Gorea, on the coast
16
of Africa”). Poey, 1858: 164-166; 255, 398 (Havana, Cuba; description,
synonymy; differs from S. becuna Lacepede; Echeneis sphyraenarum attached
to gills of barracuda). Gunther, 1860: 164 (Havana). Poey, 1864a: 179
(historical account of nomenclature of S. picuda). Goode, 1876: 62 (Bermuda;
breeding plentifully; believed DeKay’s borealis to be “closely allied to, if not
identical with, Sphyraena picuda”; food value, poisonous qualities). Goode,
1877: 292 (Bermuda; common name: “sennet”). Goode and Bean, 1879a:
116, 146, 342 ( “South Florida,” “West Florida,” Cuba, Bermudas). Goode
and Bean, 1879b: 146-147 ( “Wood’s Hole,” Pensacola, and Cuba; value of
lateral-line scales in identification; scale counts, range). Goode and Bean,
1882: 239 (Gulf of Mexico). Jordan and Gilbert, 1882: 412 ( “West Indies,
north to Florida”; description, size). Swain, 1882: 307 (identification of Esox
barracuda Shaw, 1804, with Sphyraena picuda Bloch and Schneider, 1801).
Jordan and Gilbert, 1883: 589 ( “rare at Charleston Harbor, South Carolina”;
scale count; 18-inch specimen from 10 fathoms). Meek and Newland, 1883:
67-70 (Havana, Key West, “Wood’s Hole, Massachusetts”; key to species of
western Atlantic Sphyraenidae, synonymy, range; description of S. picuda
based on 40 specimens; size attained, proportional measurements). Goode,
1884: 488 PI. 178 (food value, poisonous qualities, range, common names).
Jordan, 1884: 77-79 (Florida Keys; method of catching, associated species).
Jordan, 1885: 117 (Key West, Florida; food value). Bean and Dresel, 1885:
167 (Jamaica; common names, counts). Jordan, 1886: 572 ( “West Indies”).
Bean, 1889: 132, 145, PI. 2, Fig. 14 (Beesley’s Point, New Jersey; occurrence
of young, counts). Kendall, 1889: 308 (Garden Key, Tortugas, Florida).
Jordan and Bollman, 1889: 550 (Green Turtle Cay, Bahamas). Bean, 1890:
194, 205 (Cozumel, Yucatan; food value, ecological associates, food of bar­
racuda). Henshall, 1895: 213 (food fish at Key West; found in shallower
water than S. guaguanche [sic]). Jordan and Evermann, 1896a: 823 (descrip­
tion, synonymy, distribution in Atlantic, food value, danger to bathers, common
names). Murphy, 1914: 2 (off Bermuda, latitude 31° 50' N, longitude 50°
49' W; parasites). Metzelaar, 1919: 41 (Curasao; common name: “snoek”;
synonymy, range, seasonal occurrence, commercial importance). Weber and
de Beaufort, 1921: 70-71 (Indo-Pacific; believed that S. commersoni Cuvier
and S. agam Riippell are synonymous with S. picuda Bloch). Weber and
de Beaufort, 1922: 224-225 (Indo-Pacific; synonymy, description, common
names, size, coloration, range in Atlantic and Pacific). Fowler, 1925: 7 (Guam;
description, counts, coloration). Abe, 1939: 533 (Palao Islands; size, distribu­
tion). Blanc and Fourmanoir, 1953: 546-550 (Oceania: Tahiti, Hikueru,
Apataki, Niau, Takurea, Kaukura). Mendis, 1954: 31, 102, 103 (Ceylon).
Morrow, 1954: 806 (East Africa, between Kenya and Tanganyika; “several,
up to four feet”). Munro, 1955: 90-91, PI. 16, Fig. 246 (Ceylon; New Guinea;
description, color; range: “coastal”; S. commersoni considered a synonym).
Sphyraena nuageuse-Lizn&vdi, 1843: 64 (original description; type locality:
lie de France). Sauvage, 1891: 410-414 (Madagascar, based on Lienard;
habits; probably the same as S. agam Riippell).
Lucioperca kadenar-Montrouzier [MS], 1856 (original description).
Sphyroena kadenar-Thiolliere, 1856: 149-150 (based on MS of Montrouzier,
1856; Moiou [Melanesia]).
Sphyraena kadenar-Montrouzier, 1857: 119 (description; type locality:
Kanala, New Caledonia). Jouan, 1861: 302-303 (description based on Mont­
rouzier [1856]: poisonous qualities).
Sphyraena picuda-Poey, 1864a: 179 (synonymy; identification and renaming
of Bloch and Schneider’s picuda, which they considered to be a variety of
Esox sphyraena Linnaeus). Poey, 1864b: 187-188 (synonymy, reclassification,
nomenclatorial notes; comparison with S. becuna Lacepede, S. barracuda
17
Cuvier, S. guaguanche [sic] Cuvier, and S. picudilla Poey). Poey, 1868: 275,
277, 359 (Havana; description, synonymy, poisonous qualities; differs from
becuna Laeepede). Poey, 1875: 95 (Havana). Poey, 1881: 334 (Puerto Rico,
Cuba, Santo Domingo, Puerto Cabello, Brazil; San Martin, Panama; synonymy).
Sphyraena barracuda-Jordan and Evermann, 1896b: 335 (based on Wal­
baum; “West Indies and Brazil, north to Pensacola, Charleston and the Ber­
mudas”; common names; synonymy). Jordan and Rutter, 1897: 98 (barracuda
Walbaum has priority over picuda Bloch and Schneider). Smith, 1898: 94
(Woods Hole, “a rare straggler”; young from Quisset Harbor). Jordan and
Evermann, 1900: PI. 128, Fig. 349). Gilbert, 1900: 164 (Maceio, Brazil).
Evermann and Marsh, 1902: 115-116 (Puerto Rico: San Juan Market, San
Antonio Bridge, Mayagiiez, Ensenada del Boqueron, Hucares, Fajardo; food
value). Fowler, 1903: 749 (San Domingo, Puerto Rico; coloration; proposal
of subgenus Agrioposphyraena for this species on basis of the large scales and
the shape of the head). Jordan, 1904: 539 (Tortugas, Florida; young). Fowler,
1905: 221-222 (Beesley’s Point, New Jersey, after Bean, 1889). Shattuck,
1905: 300 (Bahamas; Spanish Wells, Governors Harbor, Powells Point, Clar­
ence Harbor, and Nassau, June and July, 1903). Jordan and Thompson, 1905:
236 (Bush Key, Tortugas, Florida; size). Bean, 1906: 41 (Flatt’s and Cable
Town, Bermuda). Fowler, 1906: 92 (young from Marquesas and Bdca Chica
Key, Florida; adults from Bahia Honda Key, Florida). Kendall, 1908: 68
(Woods Hole, Massachusetts). Vincent, 1910: 46 (Trinidad; taking trolling,
with seine, and with harpoon). Rosen, 1911: 52 (Nassau, Bahamas; common
in Bahamas; young from tidepools). Starks, 1913: 41 (Natal, Brazil). Sum­
ner, Osborn, and Cole, 1913: 447 (Woods Hole, Massachusetts). Fowler,
1915a: 50 (Santo Domingo, West Indies). Fowler, 1915b: 532 (possibly seen
at St. Kitts, West Indies). Ribeiro, 1915: 2 (Brazil). Fowler, 1916: 401 (Toro
Point, Colon, Canal Zone). Hildebrand, 1916: 306 (first record for Beaufort,
North Carolina, in “the past two years”). Ribeiro, 1918: 45 Brazil; synonymy,
description). Fowler, 1920: 155 (identifies young recorded from Beesley’s
Point, New Jersey as Sphyraena picuda Bloch and Schneider by Bean [1889]
as S. barracuda). Nichols, 1921: 22 (Turks Islands, “very common”). Meek
and Hildebrand, 1923: 282-283 (Panama: Toro Point, Mindi Cut, Colon Reef,
Fox Bay, Colon, Colon Market, Porto Bello; description). Breder, 1927: 8,
13, 24 (Cuba; food includes worm eels, other barracuda). Fowler, 1927: 264
(Philippines: Vigan; Ilokos Sur Province; Luzon). Fowler and Bean, 1927:
5 (Benkoelen Beach, Sumatra). Nichols and Breder, 1927: 72 (Long Island,
New York; Woods Hole, Massachusetts, “a rare straggler”). Beebe and Tee-
Van, 1928: 93-94 (Port-au-Prince, Haiti; description, measurements, colora­
tion, range, food, sexual maturity, young, habits). Fowler, 1928: 128-130
(Hawaiian Islands: Honolulu, Hilo, and Waialua; synonymy, distribution).
Fowler, 1929: 603 (brief description of head from market in Hong Kong,
China). Truitt, Bean, and Fowler, 1929: 62-63 (Worcester County, Maryland;
range, size, food value, danger to bathers). Nichols, 1929: 224 (San Juan,
Puerto Rico). Jordan, Evermann, and Clark, 1930: 255 (range, common names,
synonymy). Parr, 1930: 45 (Rum Cay and Crooked Island, Bahamas). Fowler,
1931: 325 (Honolulu, Hawaii; same as S. commersonii, Kner, 1879). Herre,
1931: 5 (Tenibuli, Solomon Islands). ?MacCoy, 1931: 17 (Massachusetts; “a
nine-inch little barracuda Sphyraena barracuda DeKay, from Menemsha Pond,
November, 1930”). Fowler, 1932a: 5 (Suva, Fiji; counts, coloration). Fowler,
1932b: 397 (Gasparee Island, Trinidad; coloration). Beebe and Tee-Van,
1933: 89 (Bermuda; diagnosis, field characters, size, coloration, distribution;
small individuals “not uncommon”). Gregory, 1933: 261-268 (relationships,
description and figures of the skull). Fowler, 1934: 399 (reference to previous
work [1932a] reporting S. barracuda from Fiji). Beebe and Hollister, 1935:
18
213 (Union Island, Grenadines, British West Indies). Fowler, 1935a: 276-277
(Hong Kong, China; size, counts; range: “Red Sea, Madagascar, Malaya, East
Indies, Philippines, Riu Kiu, Polynesia, Hawaii, also tropical America”).
Fowler, 1935b: 440 (doubts if S. barracuda really occurs in South Africa,
since Castelnau’s [1861] description could have equally applied to S. langsar
Bleeker, 1854a). Fowler, 1936: 578-579 (Niger River, Africa; based on speci­
men described by Gunther [1860: 336]; description, coloration, synonymy;
range: “tropical parts of the Atlantic”). Herre, 1936: 100 (Fiji, Solomon
Islands, Mindanao, Tinian; description, counts, color, habits; size: “at least
1800 mm”; record of a 106-pound barracuda from Mindanao). Fowler, 1938:
66, 90, 138, 160, 168, 195, 209, 219, 276 (Pacific: Marquesas, Tuamotus,
Tahiti, Huaheine, Bolabola, Christmas Island, Fanning Island, Honolulu,
Waialua, Hilo, Takaroa, Hawaii, Palmyra; size, counts, coloration). Delsman,
1941: 74 (Cay Sal, Bahamas). Fowler, 1941: 225 (Pivers River, Beaufort,
and Cape Lookout, North Carolina; “common” off Southport). Longley and
Hildebrand, 1941: 69-70, PI. 2, Fig. 2 (Tortugas, Florida; size, food, ecology,
feeding habits, color, counts, measurements). Rohl, 1942: 392 (coast of Vene­
zuela). Fowler, 1944: 443, 462 (Panama; Jamaica). Fowler, 1945: 128-129;
185, 290, Fig. 75 (Marquesas Keys, Lake Worth, Key West, Card Soilnd,
Boca Raton, Florida; Magnolia Beach, South Carolina; distribution, references).
Baughman, 1947: 280 (Point Isabel, Texas). Breder, 1948b: 114-115 ( “West
Indies straggling to Massachusetts”; danger to man; size, line drawing). Schultz,
1949: 109-110 (Punta Gorda, Venezuela; size). Cadenat, 1950: 149 (Senegal
River, Africa). Fowler, 1952: 90 (Port-au-Prince, Haiti). Mather, 1952: 120,
125-128 (Trujillo Bay, Guaimoreto Lagoon, Puerto Castillo, Honduras; Puerto
Barrios, Guatemala; barrier reef off Cay Bokel and at Glover Reef, Swan
Islands; Carmen, Mexico; sport fishing technics; “one of the most common
reef fish in the northwestern Caribbean”). Fowler, 1953: 63 (Barranquilla,
Colombia; coloration). Herre, 1953: 235-236 (Indo-Pacific localities; common
names, synonymy, range, literature). Murphy and Shomura, 1953a: 3, 44
(Kingman Reef, Palmyra Island, latitude 6° 13' N, longitude 163° 05' W, at
15-40 fathoms). Murphy and Shomura, 1953b: 29 (near Phoenix Islands, lati­
tude 01° 18' N, longitude 179° 55' E). Schultz, 1953: 279-280, 293, PL 23a
(Bikini, Eniwetok, Guam, Rota; keys, synonymy, description, coloration of
living and preserved specimens, ecology; should be compared with specimens
from the West Indies). Mather and Day, 1954: 178-188 (offshore records of
barracuda from Brazil, the Antilles, and south of Cape Cod). Reid, 1954: 31-
32 (Cedar Key, Florida; description; large specimens rare in shallow waters).
Kilby, 1955: 212 (Bayport, Florida; 48-mm specimen; temperature, salinity).
Shomura and Murphy, 1955: 39-49 (longline catches from the vicinity of
Christmas Island). Erdman, 1956: 321 (Puerto Rico; spawning, occurrence of
young). Iversen and Yoshida, 1956: 13, 19, 26 (longline catches near Christmas
Island and Palmyra Island). Springer and Bullis, 1956: 69, 109, 113, 122, 124
(Gulf of Mexico, 10 stations). Inger, 1957: 374-375 (Sandakan, North Bor­
neo; counts, measurements). Iversen and Yoshida, 1957: 21, 22, 24, 31, 32
(longline and troll catches from Christmas, Washington, and Palmyra islands,
and Kingman Reef; two offshore records). Williams, 1959: 107, PI. ID (East
Africa; synonymy, description, distribution, ecology, taxonomy, comparisons
with Smith [1956]).
Sphyraena snodgrassi-Jenkins, 1901: 387-388 (original description; type
locality: Honolulu; type specimen: USNM 49693; “most closely related to
S. commersonii”) . Fowler, 1903: 749 (Hawaiian Islands, Tahiti; same as
S. commersoni, Fowler 1900; description, color; “close to S. barracuda, ap­
parently differing in the shorter maxillary”). Jenkins, 1904: 438 (believed that
S. snodgrassi Jenkins, 1901, is the same as S. commersoni. Jordan and Ever-
19
 TABLE 1
M ea su r em en ts expressed as r a t io s of body pa r ts and m e r is t ic data of Sphyraena barracuda and sy no nym s
ASSIGNED TO IT.
barracuda barracuda barracuda barracuda akerstromi
Western Atlantic Marshall Islands Indian Ocean Red Sea Western Australia
Schultz (1953) Smith (1956) USNM 147455 Whitley (1947)
Number of specimens 41 4 4 1 2
Standard length range (mm) 213-1111 208-1240 180-1345 147 835-1210
In standard length:
First predorsal length 2.3-2.6 2.3 2.3-2.5 2.2 2.4-2.5
Second predorsal length 1.4 — 1.4-1.5 1.4 —
Prepectoral length 3.0-3.2 — 2.7-3.4 3.0 —
Prepelvic length 2.4-2.6 — 2.4-2.9 2.5 2.5-2.7
Preanal length 1.3-1.4 — 1.3-1.5 1.4 —
Head length 3.0-3.7 3.2-3.4 2.8-3.3 3.2 3.7-3.9
Pectoral fin length 8.4-9.1 — 8.3-9.1 4.9 8.4-8.9
Pelvic fin length 9.8-13.2 — 9.5-13.2 5.1 13.0-13.1
In head length:
Length of snout 2.1-2.2 2.2 2.2-2.1 2.2 2.0-2.1
Length of maxillaries 1.7-2.2 1.8-2.2 — 2.3 1.8-2.0
Interorbital width 4.5-7.1 3.6-6.4 3.7-5.3 6.6 3.3-3.6
Greatest depth 6.2-7.5 5.5-7.0 6.7-7.2 3.9 5.5-6.1
Depth caudal peduncle 4.0-4.5 4.3-5.5 4.0-4.5 4.4 3.8-4.0
Distance between
dorsal fin origins 3.6-3.7 3.6 3.5-3.7 1.3 3.3
Eye 6.0-9.2 6.0-9.0 6.6-11.2 6.0 7.8-8.3
Postorbital 2.4-2.7 2.5-2.6 1.8-2.5 2.8* 2.4-2.5
Origin pectoral base
to pelvic base 4.0-4.5 4.3-4.4 4.3-7.5 5.0 —
Eye in postorbital 2.2-3.8 23-3.1 2.7-6.1 # 2.1 3.2-3.4
Dorsal elements V+U8 V+U8 V + I,9 4 V + I,i,9 V + i,9
Anal elements I,i,7-8 U7 1,8 Li,8 i,8
Pectoral elements ii,11-12 ii,11-12 i,l 1-ii, 10 ii-12 i,13-14
Transverse scales 10/1/9 to 13/1/14 — 10/1/9 — 9 /1/15 to 12/1/14
Lateral-line scales 75-87 76-85 75-90 84 76-83
 ?

TABLE 1— (CONTINUED)
microps picuda snodgrassi agam
Queensland “Indo-Pacific,” Hawaii, Red Sea,
Marshall (1953) Weber and de Beaufort (1922) USNM 46963 Riippell (1835)3
Number of specimens 1 ? 1 1
Standard length range (mm) 1336^ ? 253 732
In standard length:
First predorsal length 2.5 2.4
Second predorsal length — — 1.4 —
Prepectoral length — — 3.2 —
Prepelvic length — — 2.6 __
Preanal length — — 1.4 __
Head length 3.5 2.4-2.8 3.2 3.5
Pectojal fin length 11.6 — 8.4 __
Pelvic fin length 13.1 — 8.9 __
In head length:
Length of snout 2.1 2.2
Length of maxillaries 2.2 2.2-2.4 2.3 __
Interorbital width — — 6.5 __
Greatest depth 6.2C 5.8-7.2 6.7 7.0
Depth caudal peduncle — — 2.7 —
Distance between
dorsal fin origins _ 3.5
Eye 10.9 6.0-6.4 6.5 8.5
Postorbital --- __ 2.6 __
Origin pectoral base
to pelvic base 4.2
Eye in postorbital --- — 4.0 __
Dorsal elements V +1,9 V + ii,8 V 4T,i,8 V + ii,8
Anal elements ii,8 ii,7 I,i,8 ii,7
Pectoral elements — 13 ii-11 i-13
Transverse scales — 13/1/14 10/1/10 11/1/13
Lateral-line scales — “± 8 0 ” 79 81
8 Personal communication from Dr. Wolfgang Klausewitz.
bConverted from total length.
c Specimen gutted.
mann, 1905: 141-143 (Hawaiian Islands). Jordan and Seale, 1906: 218
(Hawaii). Jordan and Snyder, 1907: 208 ( “probably not the same as the East
Indian S. commersonii”\ counts; young near shore and in mullet ponds).
Iordan and Jordan, 1922: 29-30 (common in Hawaii; common name: “kaku”;
size; should be compared with other large specimens of the South Seas).
Barnard, 1925: 312 (synonym of S. commersoni). Jordan and Evermann,
1926: 3, 8 (Hawaii; common name, habitat, range into fresh water). Schultz,
1943: 82-83; 256-257 (Canton Island; comparison needed with S. barracuda;
Phtheirichthys multiradiatus Schultz from S. snodgrassi) .
Sphyraena akerstromi-Whitley, 1947: 131-136, PL 11, Fig. 1 (original de­
scription; type locality: Lowendal Island, Western Australia; extensive counts
and measurements on four specimens; relationships, food, gonads; close “to
Weber and de Beaufort’s Sphyraena picuda”). Schultz, 1953: 283 (S. akerstromi
Whitley the same as S. barracuda Walbaum). Smith, 1956: 41 (same as
barracuda) .
Sphyraena ra/crops-Marshall, 1953: 54-55, Fig. 2, PI. 2 (identification as
barracuda questionable; original description; type locality: Moreton Bay, South
Queensland, Australia). Smith, 1956: 41 (S. microps Marshall is the same as
Agrioposphyraena barracuda [Walbaum]).
Agrioposphyraena barracuda (Walbaum)-Smith, 1956: 40-41, PL I, Figs.
4a-4c (new combination; Fowler’s [1903] subgenus raised to generic level,
based on Esox barracuda Walbaum, 1792; synonymy, description, keys, meas­
urements, range in Indo-Pacific, coloration, danger to man, food value,
ontogenetic changes in proportional measurements).

Apparently some confusion that stems from the use of 17 specific

names for Sphyraena barracuda (Walbaum) has arisen from a failure to
appreciate ontogenetic and individual variation in this species. This
variation, recognized by Weber and de Beaufort (1921, 1922), Beebe
and Tee-Van (1928), Schultz (1953), and Smith (1956), accounts for
some of the problems in comparing individuals of different sizes. Although
it is not intended to present detailed infraspecific comparisons in this
paper, it seems desirable to relate specimens of barracuda from the
tropical western Atlantic to those of other regions, as well as to some
nominal species (e.g., akerstromi Whitley; microps Marshall; snodgrassi
Jenkins) for which data are available and which are evidently synonyms of
barracuda. Data in the literature are often presented as the ratio of a
body part in body length, and for purposes of comparing my data with
those in the literature this type of presentation is used here (Table 1).
This method, however, leaves much to be desired since the specimens
encompass a wide range of sizes. Thus, ontogenetic variation over this
range is not appreciated unless measurements are presented for fish of
comparable sizes, or unless the measurements are depicted graphically
as regression slopes. The value of presenting original variates as a
function of length as against presenting percentages or ratios of lengths
has been discussed by Marr (1954) and Parr (1956).
Methods of Measuring and Counting.— The following methods were used
in making proportional measurements:
(1) Standard length: tip of snout with mouth closed to midpoint of hypural
22
 TABLE 2
M easu r em en ts of Sphyraena barracuda e x p r e sse d as pe r c e n t a g e s o f t h e sta n d a rd l e n g t h fr o m t h e w e st e r n
A t l a n t ic O c e a n (i n pa r en th eses) and the w e ster n I n d ia n O c e a n ( S m it h , 1956)

Standard Length (mm) 60 (61) 110 (111.5) 150 (149) 180 (182) 370 (369) 900 (861) 1345 (1111)
Depth 14 (14.4) 15 (13.0) 15 (15.8) 15 (14.0) 15 (14.9) 14 (13.7) 14 (14.9)
Head 38 (40.0) 37 (37.0) 34 (36.2) 35 (36.0) 33 (3.3) 30 (28.7) 30 (29.0)
Eye 6.5 (6.7) 5.8 (6.0) 5.3 (5.8) 5.2 (5.7) 5 (4.6) 3.6 (3.9) 2.7 (3.3)
Interorbital width 4.5 (4.4) 5 (4.6) 5 (6.7) 6.5 (5.3) 6.8 (5.4) 7.5 (7.6) 8 (6.7)
Snout length 17 (17.5) 16.5 (16.7) 16 (16.1) 15 (15.3) 15 (14.9) 11.8 (14.3) 11 (14.6)
Postorbital length 15 (15.2) 14 (13.9) 12.3 (14.3) 13 (15.1) 13 (13.0) 11.5 (10.4) 12 (14.4)
Pectoral fin length 10 (11.0) 10 (10.1) 11 (10.3) 11 (11.0) 12 (11.9) 11 (11.5) 12 (11.1)
Pelvic fin length 10 (9.7) 10 (10.8) 11 (11.3) 10.5 (10.2) 10 (8.8) 7.5 (8.6) 7.5 (7.6)
Depth caudal peduncle 6.8 (6.7) 7.8 (6.6) 8 (7.4) 8 (7.7) 8 (7.6) 7.5 (7.5) 7.3 (6.9)
First predorsal length 48 (47.5) 45 (45.3) 44 (43.9) 44 (43.5) 44 (40.1) 40 (40.8) 41 (42.1)
Second predorsal length 75 (73.0) 75 (71.8) 72 (71.1) 72 (72.5) 73 (70.2) 67 (72.1) 70 (72.5)
Prepectoral length 37 (37.2) 37 (35.5) 35 (35.1) 35 (33.8) 35 (31.2) 30 (30.9) 29 (31.2)
Prepelvic length 43 (43.8) 41 (41.6) 40 (42.1) 41 (42.3) 40 (39.3) 34 (38.3) 36 (38.7)
Preanal length 73 (74.6) 73 (72.8) 72 (71.6) 73 (75.7) 75 (72.1) 38 (73.7) 71 (74.9)
 fold, approximately beneath the base of the second branched ray of
upper caudal lobe.
(2) First predorsal length: snout tip to origin of erect first dorsal fin.
(3) Second predorsal length: tip of snout to origin of second dorsal fin.
(4) Prepectoral length: tip of snout to origin of erect pectoral fin.
(5) Prepelvic length: tip of snout to origin of erect pelvic fin.
(6) Preanal length: tip of snout to origin of anal fin.
(7) Head length: tip of snout to posteriormost part of fleshy tip of opercle.
(8) Maxillaries: tip of snout to posterior border of maxillary including the
premaxillary and maxillary bones.
(9) Snout length: tip of snout to anterior rim of fleshy orbit.
(10) Eye diameter: horizontal distance between rims of fleshy orbit.
(11) Interorbital width: least width of bony portion of skull measured be­
tween orbits.
(12) Greatest depth: greatest depth measured at origin of spinous dorsal fin
(fish with stomachs obviously distended with food are excluded).
(13) Caudal peduncle: least depth.
(14) Pectoral fin length: length from upper base of erect pectoral fin to tip
of longest ray.
(15) Pelvic fin length: length from origin of erect pelvic fin to tip of long­
est ray.
(16) Length of spinous dorsal fin: length from origin of first dorsal fin to
tip of longest depressed spine.

Scale counts were taken according to the methods of Hubbs and

Lagler (1947: 8-13). Fin-ray counts were made following the methods
of Schultz (1953: xxii-xxiii).

Comparisons.— Measurements of barracuda from the tropical western

Atlantic are compared with Smith’s (1956) data of barracuda from the
western Indian Ocean (Table 2). It is seen that in specimens of compar­
able size most measurements agree well for the two regions. Possible
sexual dimorphism, if any, has not been resolved but would presumably
lead to a misinterpretation of the data. My measurements of large specimens
in which the sex is identified are few, but at the extreme ranges those for
which I have data are females. Whitley’s akerstromi, which is shown to
be a synonym of barracuda, is represented in Figures 2 and 3 and,
fortunately, his two specimens represent both sexes. A comparison of these
limited data reveals that the body parts of the female seem to be consis­
tently though only slightly larger in relation to standard length and head
length.
Differences in proportional measurements between specimens from the
Atlantic and Indian oceans (Smith, 1956) are most noticeable for the
larger individuals (Table 2); this is evidently due to the discrepancy in
size in the largest of the specimens compared from the Atlantic (1111
mm) and the Indian oceans (1345 mm). In these two individuals minor
differences occur in the eye diameter, snout length, and postorbital length.
However, the interorbital width seems consistently smaller in the Atlantic
specimens, but possibly this reflects a difference in method of measuring,
24
 10 100 1000 10 100 1000
STANDARD LENGTH, MM. STANDARD LENGTH, MM.
2. Changes in proportional measurements with growth of Sphyraena
F ig u r e
barracuda (including synonyms akerstromi, microps, and snodgrassi). Measure­
ments are expressed as percentage of standard length. Dots are western
Atlantic specimens of barracuda; circles are Indian Ocean specimens of barra­
cuda from Smith (1956); solid triangles are akerstromi Whitley, 1947, from
Western Australia; solid square is microps Marshall, 1953, from Queensland,
Australia; cross is barracuda, USNM 147455, from Djidda, Red Sea; solid
diamond is snodgrassi Jenkins, 1901, USNM 46963, from Honolulu, Hawaii.
A. First predorsal length. B. Prepectoral length. C. Second predorsal length.
D. Interorbital width. E. Least depth caudal peduncle. F . Length of dorsal
spines. G. Head length. H. Snout length. I. Eye diameter. J. Preanal length.
K. Prepelvic length. L. Pectoral fin length. M. Length of maxillaries. N. Great­
est body depth. O. Pelvic fin length.
since Smith (1956) did not state his method. Also noticeable is the marked
discrepancy in snout length between the two largest specimens. Perhaps
this indicates a real difference between the individuals from the two
regions, since, as suggested by the plots of akerstrom i in Figures 2H and
3D, the difference does not seem to be sexual for this character.
Ratios, measurements, and counts for barracuda and its synonyms,
which have been collated from the literature, as well as from original
measurements on museum specimens, are listed in Table 1. The range in
ratios of each specimen in the standard length is given for 41 specimens
of barracuda from the tropical western Atlantic, ranging from 213 to 1111
mm standard length. This method is used here to permit comparisons with
the data of Schultz (1953) since he presented ratios but did not give
original measurements.
Differences between measurements which cannot be attributed to size
differences and in which there is no overlap in ranges occur in akerstromi
and the western Atlantic specimens of barracuda in interorbital width
and in distance between dorsal fin origins. Marshall’s (1953) type of
m icrops evidently has a longer pectoral fin than does Smith’s (1956)
barracuda of comparable size from the Indian Ocean. However, the
proportions presented here for microps should be viewed with caution,
since standard length was not given by Marshall. Total length was
converted by me to standard length by multiplying by a factor of 0.829
which was found for barracuda from the western Atlantic; this factor may
conceivably not be applicable for specimens from other geographic regions.
The ranges of meristic counts from various regions, including presumed
synonyms of barracuda , are comparable. The fin-ray counts are presented
as taken from the literature. The pelvic and spinous dorsal fin-ray counts
showed no variation, the former being consistently 1,5. Combining total
dorsal fin elements for ease of comparison (cf. de Sylva, 1955:30-31), the
count obtained is 10 or 11, with the USNM specimen from the Red Sea
having the highest count. The range for all specimens is 9 or 10 anal
elements and that for the pectoral elements varies from 12 to 15. Whitley
(1947) gave a somewhat higher pectoral fin-ray count in his description
of akerstromi. In western Atlantic specimens of barracuda , the lateral-line
scales vary from 75 to 87, although Smith (1956) listed a maximum
count of 90 for specimens from the Indian Ocean. However, since he stated
that he has counted about 90 scales on the illustration of Riippell’s
(1835) type of agam, perhaps he included this in his count. However,
I previously pointed out that the type of agam has only 81 scales, and thus
perhaps the range presented by Smith should be somewhat lower. The
ranges of transverse scales around the body appear to be comparable for
different areas.
Original measurements for barracuda from various regions have been
plotted in Figures 2 and 3. When points representing measurements for
regions other than the western Atlantic are plotted in these figures, rough,
26
 HEAD LENGTH,MM. HEAD LENGTH,MM.
3. Changes in proportional measurements with growth of Sphyraena
F ig u r e
barracuda (including synonyms akerstromi, microps, and snodgrassi). Measure­
ments are expressed as percentages of the head length. Dots are western
Atlantic specimens of barracuda; circles are Indian Ocean specimens of barra­
cuda from Smith (1956); solid triangles are akerstromi Whitley, 1947, fron
Western Australia; solid square is microps Marshall, 1953, from Queensland,
Australia; cross is barracuda, USNM 147455, from Djidda, Red Sea; solid
diamond is snodgrassi Jenkins, 1901, USNM 46963, from Honolulu, Hawaii.
A. Pelvic fin length. B. Interorbital width. C. Pectoral fin length. D. Snout
length. E. Length of maxillaries. F . Least depth caudal peduncle. G. Greatest
body depth. H. Eye diameter.
visual comparisons may be made. If these figures are compared with the
data presented in Table 1, a number of features do not become evident
unless viewed graphically. In comparison with the western Atlantic
specimens, Smith’s (1956) two largest fish have a somewhat shorter
second predorsal length, preanal length, prepelvic length, and snout length.
These comparisons are magnified when the head length is used as a base
length (Fig. 3). Although measurements for large fish are too scarce to
draw any critical conclusions, the preanal and snout lengths seem to present
real differences which are far removed from the regression slope. Possibly
Smith’s (1956) specimens of barracuda from the Indian Ocean differ
racially from those of the western Atlantic on the basis of these two
characters. A statistical comparison of specimens from the two regions
based on a larger sample from the Indian Ocean is needed. However,
until this can be done I consider the fish from the two regions identical.
Whitley’s (1947) akerstrom i, which is said to differ from picuda ( =
barracuda) in the “size of the eye and its ratios to other parts of the
head,” as well as in the length of the maxillary bones and the pectoral fin,
is seen to approximate the points for West Indian specimens very well.
The latter specimens show somewhat greater depth and interorbital width,
but as evidenced from the considerable variance of points (Figs. 2N and
3G) depth is a poor character on which to base critical comparisons.
Differences in interorbital width may be due to method of measuring.
Although Marshall’s (1953) description of m icrops is inadequate for
analysis, possibly this species does not belong in the synonymy of barra­
cuda. The specific name, m icrops , on which Marshall apparently based his
diagnostic character, does not refer to a differentiating character, as the
eye diameter falls well within the scatter of points represented by
barracuda. However, apparent distinctions from barracuda are seen in the
pectoral fin length and the length of the maxillaries. Marshall referred to
“maxillary” measurement but he probably alluded to the length from the
tip of the snout to the posterior edge of the maxillary series, including
the premaxillary, maxillary, and supramaxillary. In his illustration (PI. 2,
Fig. 2) he showed what appears to be an elongated supramaxillary,
reaching far past the eye, but with the maxillary considerably shortened.
Since the juxtaposition of the two bones as depicted by Marshall does not
occur in any sphyraenid which I have seen, it seems that the type of
m icrops may be malformed and the maxillary shorter than in the normal
situation, and the supramaxillary is normally developed; in this case
Marshall may have measured only to the end of the maxillary. If the
measurement were taken to the tip of the supramaxillary as shown (PI. 2,
Fi^. 2) the measurement would be more nearly similar to that found in
other sphyraenids at this size.
Unless Marshall’s type has a malformed pectoral fin, it is difficult to
explain the difference in the relative length of the fins of barracuda and
m icrops of comparable size, unless there is a sexual difference, or unless
28
microps can be considered as actually representing a valid species on the
basis of this single character. Tentatively, however, I follow Smith (1956)
in placing m icrops in the synonymy of barracuda.
Neither the Red Sea specimen nor the type of snodgrassi Jenkins from
Honolulu seems to differ in any respect from the western Atlantic speci­
mens of barracuda , although perhaps differences in proportional measure­
ments would not be disclosed at such a small size. Both the pectoral and
pelvic fins do appear to be somewhat longer in snodgrassi in comparison
with the western Atlantic species, but probably not significantly so.

Y oung Stages of the F a m il y S p h y r a e n id a e

The smallest postlarval (Hubbs, 1943) specimen of barracuda collected
by me measured 11.9 mm standard length and was collected off Bimini,
Bahamas. Smaller specimens collected by the DANA vessels from
1920 to 1922 in the tropical western Atlantic were made available by the
late Dr. A. Vedel Taning of the Carlsberg Foundation. The description of
the types of collecting gear and methods, including hydrographic observa­
tions, are found in Schmidt (1929) and Jespersen and Taning (1934);
data for hydrographic stations occupied where sphyraenids were collected
are summarized in Table 7.
A graded series of 171 barracuda collected from southern Florida and
Bimini, Bahamas, as well as from the DANA localities encompass a
size range of fish from 5.5 mm (to base of urostyle) to 1111 mm standard
length. In measuring specimens, an ocular micrometer was used for those
up to 28.7 mm, dial calipers reading to the nearest 0.1 mm for those
28.7 mm to 96.6 mm, and dividers and meter stick were used for longer
specimens. Original measurements are given only for standard length; the
remainder are expressed as percentages of the standard length, or of
head length (Fig. 3).
Eighty-five specimens of barracuda were taken from the tropical western
Atlantic Ocean, most of which came from the Virgin Islands. Specimens
identified as guachancho Cuvier were taken from the Bahamas, the Virgin
Islands, Jamaica, and Bermuda; two individuals tentatively identified by
me as sphyraena Linnaeus occurred in a collection from the northeastern
coast of South America. Additional specimens from these areas represent
either borealis De Kay or picudilla Poey. These closely related species pose
a difficult problem at this time, but since borealis is apparently restricted
to the mainland of North America I am therefore tentatively assuming
that this fifth species in the DANA collections is picudilla.
Through the courtesy of Dr. Ernest A. Lachner, three drawings of
young sphyraenids by Dr. Louella E. Cable have been loaned to the writer
from the files of the United States National Museum. Two of these
represent guachancho and borealis; the third drawing is of a specimen
9 mm total length which is labelled (<Sphyraena barracuda ” from Beaufort,
North Carolina. Although I have not seen the specimen, this drawing is
29
F ig u r e 4. Photographs showing pattern in three species of Sphyraena from the
western North Atlantic Ocean. A. Sphyraena borealis, 66.0 mm standard
length; collected by Sherman C. Bishop at Fire Island Inlet, Long Island, New
York, August 11, 1938, Cornell University No. 6277. B. Sphyraena barracuda,
62.7 mm standard length; collected by Donald P. and Doris D. de Sylva at
Bimini, Bahamas, July 13, 1956. C. Sphyraena guachancho, 63.3 mm standard
length; collected by Donald P. de Sylva and Nardo Zaias at Bear Cut, Key
Biscayne, Miami, Florida, September 26, 1952. Photograph by Douglass M.
Payne.
of a young specimen of S. borealis. The fleshy tip of the lower jaw separates
it from barracuda, and the row of large, distinct melanophores along the
middorsal ridge, and the lack of pigmentation at the anal and soft dorsal-
fin bases separate it from guachancho (Fig. 5). The body is too deep, the
dorsal pigmentation not sufficiently distinct, the fleshy tip of the lower
jaw is too short, and the head lacks the pigmentation of S. sphyraena of
comparable size; furthermore, the latter is a rare species in the western
North Atlantic, is doubtfully recorded from Bermuda (Barbour, 1905),
and otherwise is definitely known in the western Atlantic only from the
coast of Brazil. Slight differences in pigmentation and size of the eye are
noted between Dr. Cable’s drawing and the DANA specimens of picudilla.
If it may be inferred that the young have about the same distribution as
the adults it would be expected that this specimen represents borealis.
On the assumption that borealis and picudilla are valid, allopatric
species, a tentative key is presented for the identification of postlarval
sphyraenids for specified size groups. Only two postlarval specimens of
S. sphyraena have been identified, both of which were in poor condition;
thus, characters based on these specimens should be used with caution.
Characters used to identify the young cannot be used to separate the
adults since they may not be present in the young. The key is subdivided
roughly into fish from 5.5 to 11.9 mm, from 12.3 to 130 mm, and fish
greater than 130 mm. Since there is no pronounced metamorphosis in
this family, and because there is considerable variation in the color pattern
of specimens of comparable size, there is actually no trenchant break
between the keys; those specimens which fall near the extremes of the
ranges should be checked using both keys.

K ey to S p e c ie s of S p h y r a e n id a e fo r S p e c im e n s of

5.5 TO 11.9 MM FROM THE WESTERN NORTH ATLANTIC

la. Head comparatively long, about 34-39 per cent of the standard length;
body not heavily pigmented; distinct melanophore patterns on dorsal and
ventral ridge and along posterior part of lateral line only; fleshy tip of
lower jaw developed to a length less than half the eye diam eter............. 2
lb. Head comparatively short, less than 30 per cent of the standard length;
body heavily pigmented on posterior part of body and about supraoccipital
region; a distinct mid-lateral row of melanophores along most of the
length of the caudal peduncle; fleshy tip of lower jaw nearly equal to eye
diam eter....................................Sphyraena sphyraena, European barracuda
2a. Melanophores along middorsal ridge not noticeably larger than surround­
ing ones, scattered and not distinctly arranged serially (Figs. 5B, C);
supraoccipital region not noticeably pigmented; tip of snout with slight
pigmentation, if at a l l .........................................................................................3
2b. Middorsal line straddled by relatively large, distinct melanophores ar­
ranged serially from point above pectoral origin to end of soft dorsal-fin
base; melanophores at bases of soft dorsal and anal fins, if present, small
and diffuse (Fig. 5A ); a few large melanophores on occipital region; tip
of snout generally heavily pigmented ............................................................4
31
 5. Drawings showing pigmentary differences in the postlarvae of three
F ig u r e
species of Sphyraena from the western North Atlantic. A. Sphyraena picudilla,
dorsal aspect. B. Sphyraena barracuda, dorsal aspect. C. Sphyraena guachan-
cho, dorsal aspect. D. Sphyraena picudilla, ventral aspect. E. Sphyraena
barracuda, ventral aspect. F . Sphyraena guachancho, ventral aspect. All
specimens are about 9 mm standard length.
32
3a. A row of melanophores along midventral ridge extends posteriad only
slightly past tips of apressed pectoral fins; tip of lower jaw not noticeably
conical or projecting beyond snout; pigmentation on tip of lower jaw and
snout sparse; lower jaw obliquely rounded in dorsal view; teeth nearly
vertical in lateral view; faint pigmentation only on either side of dorsal
midline; melanophores at base of dorsal and anal fins rather large, dense
(Fig. 6A-D); preorbital pigmentation usually distinct, subtriangular in
outline; no hourglass bands on posterior section of trunk; snout rounded
above, dorsal aspect of frontal region sigmoid in lateral view, particularly
in specimens less than 7 mm long; lower posterior margin of opercle
without spines; opercle ends in a single point; maxillary series nearly
straight when viewed laterally . . . . Sphyraena barracuda, great barracuda
3b. Midventral row of melanophores extends posteriad from isthmus to vent
(Fig. 5F); tip of lower jaw projects noticeably beyond snout (as compared
to barracuda), ending in a bluntly conical, fleshy knob; lower jaw tip
heavily pigmented; snout and lower jaw tip acute in dorsal view; teeth
directed obliquely backward, rather than vertically; faint pigmentation
only on either side of middorsal line, melanophores at bases of soft dorsa!
and anal fins diffuse; two hourglass-shaped crossbands on sides in larger
specimens: the first begins about two-thirds of the distance between the
pectoral fin origin and the anal fin origin, the second commences just be­
hind rear of anal fin base; preorbital pigmentation generally not distinctive;
dorsal profile of snout nearly straight in lateral view; lower posterior
margin of opercle with three weak spines; opercle ending in two points;
maxillaries rather strongly arched in lateral view ..........................................
Sphyraena guachancho, guaguanche
4a. Preorbital pigmentation extending forward to snout tip as a line of
melanophores; dorsal profile of snout nearly straight in lateral view; caudal
base with a vertical series of melanophores; eye larger in specimens of
comparable s i z e ................................................... Sphyraena picudilla, sennet
4b. Preorbital pigmentation diffuse, scattered; dorsal profile slightly concave
in lateral view; caudal base without pigmentation; eye larger in specimens
of comparable s i z e .........................Sphyraena borealis, northern barracuda
K ey to S p e c ie s of S p h y r a e n id a e f o r S p e c im e n s f r o m
1 2 .3 to 130 mm Standard L eng th fr o m the w e st e r n
N o r t h A t l a n t ic
la. Spinous dorsal fin placed well behind origin of pectoral fin; pectoral fin
reaches origin of pelvic f i n ................................................................................2
lb. Spinous dorsal fin placed above or slightly in front of origin of pelvic fin;
pectoral fin does not reach origin of pelvic f i n ...............................................3
2a. Last rays of soft dorsal and anal fin longer than anterior rays; teeth
noticeably directed backwards; scales, when completely formed, 108 to
114 in lateral line; interorbital noticeably convex in large specimens, with
raised longitudinal frontal and parietal crests; maxillaries strongly arched
upwards; opercular margin ending in two points; lateral line raised,
distinct, comparatively wide, at least twice the width of the pigment band
running along either side of it, its scales noticeably larger than the sur­
rounding scales; circumorbital and opercular region with silvery reflections
in preserved specimens; upoer surface of tip of lower jaw blackish; tips
of middle caudal rays usually blackish with center rays darkest, stippled;
three hourglass-shaped bands on body (Fig. 4C) : the first forms a dusky
saddle on back beginning at about the last spine of the first dorsal fin
and extends posteriad to about one-third the interdorsal distance; a second,
wider saddle extends vertically to below dorsal base, becomes narrow at
 the lateral-line region, and thence again flares at region of anal base; a
third hourglass mark occurs halfway between the second dorsal fin and
the caudal fin base, similar to the first two; margins of dorsal fins blackish
with pale bases; pectoral fin dusky, other fins pale; upper margin of
opercle p a l e .......................................... Sphyraena guachancho, guaguanche
2b. Last rays of soft dorsal and anal fin equal to or shorter than anterior rays;
teeth nearly vertical in lateral view in most specimens; scales, when com­
pletely formed, 75 to 87 in lateral line; interorbital area flattish to concave,
without raised longitudinal ridges; maxillaries nearly straight in lateral
view; opercle ends in a single point; lateral line whitish, not distinctly
raised or noticeable; sides of head seldom with silvery reflections, and, if
present at all, only about opercular bones; tips of middle caudal rays
diaphanous; outer caudal fin rays dark, except for their tips; a subtri-
angular, darkly pigmented region overlaps base of middle caudal rays
and caudal base; sides with seven to 12 variable markings which form a
longitudinal band in fish less than 30 mm; these form contiguous H-shap-
ed blotches in fish to about 60 mm (Fig. 4B); in larger individuals some
of the blotches may extend to and coalesce with the dorsal pigmentation;
soft dorsal fin with white tip in large specimens of this size range; anal
fin base pale; base of pelvic fins pale, rest of fin dusky; pectoral fin pale
except at its base; upper margin of opercle d u sk y ..........................................
Sphyraena barracuda, great barracuda
3a. Scales very small, about 150 to 160 in lateral line; fleshy tip of lower jaw
well developed; pigmentation on back and sides obscure, not forming a
distinctive pattern.................... Sphyraena sphyraena, European barracuda
3b. Scales larger, from 110 to 135 in the lateral line; pigmentation on back
and sides well developed, forming a series of quadrate saddles on back,
alternating with subquadrate rectangles along sides (Fig. 4 A ) ..................4
4a. Eye large in specimens of comparable size, averaging 8.0 per cent of
standard length in specimens of about 50 mm, and 7.0 per cent in speci­
mens of about 100 mm; interorbital space convex; median ridge of frontal
groove not developed.......................................... Sphyraena picudilla, sennet
4b. Eye relatively smaller in specimens of comparable size, averaging 7.0 per
cent in specimens of 50 mm standard length, and 6.0 per cent in specimens
of 100 mm; interorbital space flattish; median ridge of frontal groove
prom inent...................................... Sphyraena borealis, northern barracuda
K e y t o t h e S p e c ie s o f S p h y r a e n id a e fo r S p e c im e n s
LONGER THAN 130 MM FROM THE WESTERN NORTH ATLANTIC
la. Origin of spinous dorsal fin definitely inserted behind origin of pelvic fin;
tip of apressed pectoral fin reaches origin of pelvic fin; tip of lower jaw
without a distinct, fleshy t i p ........................ .....................................................2
lb. Origin of spinous dorsal fin above or slightly in front of origin of pelvic
fin; tip of apressed pectoral fin falls considerably short of origin of pelvic
fin; tip of lower jaw generally with a distinct, fleshy tip ........................... 3
2a. Scales small, numbering from 108 to 114 along the lateral line from the
upper opercular cleft to the hypural fold; when fins are depressed, the last
rays of the soft dorsal and anal fin extend beyond the anterior rays;
interorbital area convex; teeth directed obliquely backward; no numerous,
oblique bars on sides and back, except that three encircling bands may
be present on the posterior section of the trunk in specimens less than
about 160 mm; lower sides lack scattered, inky blotches; color silvery
to olive, with a faint yellow to golden longitudinal stripe in life; margins
of pelvic and anal fin black; tips of middle caudal rays black; size to
about 24 inches; Woods Hole, Massachusetts, to Brazil and throughout the
34
 Gulf of Mexico and Caribbean Sea, drifting to the eastern Atlantic...........
Sphyraena guachancho, guaguanche
2b. Scales large, 75 to 87 in the lateral line; when fins are depressed, the
anterior rays of the soft dorsal and anal fin extend beyond the posterior
rays; interorbital area flattish to concave; teeth vertical; sides of smaller
specimens in this range never with encircling bands on posterior region of
trunk; adults deep green to steel gray on back, sometimes reflecting
purplish hues; mostly silvery on sides, abruptly becoming white along
ventral surface; upper sides with 18 to 22 oblique, backward-directed
dark bars in life; lower sides with several to many inky blotches which
are variable in size and position (Fig. 7D ); soft dorsal, anal, and
caudal fins black to violet, with whitish tips; pectoral fin pale except for
its upper margin, which is black; upper border of opercle dusky to black;
length to about 6 feet; Woods Hole, Massachusetts, south to Rio de
Janeiro, Brazil, and throughout the Gulf of Mexico and Caribbean Sea;
also occurring in all other tropical waters of the world except the eastern
Pacific .............................................. Sphyraena barracuda, great barracuda
3a. Scales comparatively large, 110 to 135 in the lateral line; depth of body
12.0-14.3 per cent of the standard le n g th ..................................................... 4
3b. Scales very small, from 150 to 160 in the lateral line; depth of body 10.0-
11.1 per cent of the standard length; length to about 24 inches; eastern
Atlantic from the Bay of Gascogne to Mossamedes, Angola, and the
Mediterranean and Black seas; also recorded from Bermuda and Rio de
Janeiro, B r a z il........................ Sphyraena sphyraena, European barracuda
4a. Eye large in specimens of comparable size, averaging 6.3 per cent in
standard length in specimens 150 to 200 mm, 6.0 per cent in specimens
250 mm, and 5.5 per cent in specimens 300 mm; body depth about 13.9-
14.3 per cent of standard length; scales in lateral line 107 to 116; inter­
orbital area convex; median ridge of frontal groove little developed; length
to about 18 inches; Tortugas, Florida, Bermuda, and throughout the
Antilles, south to Rio de la Plata, Uruguay. . .Sphyraena picudilla, sennet
4b. Eye smaller in specimens of comparable size, averaging 5.3 per cent in
standard length in specimens 150 mm standard length, 4.9 per cent in
specimens 200 to 250 mm, and 4.7 per cent in specimens 300 mm; body
depth about 12.0-13.0 per cent of standard length; scales in lateral line
from 118 to 135; interorbital area flattish; median ridge of frontal groove
prominent; length to about 15 inches; Woods Hole, Massachusetts, to
Miami, Florida, and throughout the Gulf of Mexico south to Panama;
also questionably recorded from Bermuda, Santo Domingo, and Rio de
Janeiro .......................................... Sphyraena borealis, northern barracuda
P o s t l a r v a l D e v e l o p m e n t o f Sphyraena barracuda
The illustration showing the postlarval development of S. sphyraena
from the Mediterranean by Lo Bianco (1937), the illustrations of S.
pinguis and an unidentified species by Uchida, et al. (1958: 46, 47, PI. 44,
Figs. 2-4, PL 47, Figs. 1-8), and Orton’s (1955) work on the develop­
ment of argentea apparently constitute the references on this subject. Illus­
trations of young specimens of barracuda are found in Fowler (1945),
Gordon (1946), and Breder (1949).
Essential to a study of the systematics of Sphyraena barracuda is a
knowledge of ontogenetic changes in proportional measurements. For
purposes of illustration, graphs showing changes in relation to the head
length (Fig. 3), as well as in relation to standard length (Fig. 2), are
35
presented. When head length is used as a base, it tends to magnify
variation, and thus it appears to be a better indicator of interspecific
differences, since, depending upon the species, different body parts grow
at relatively different rates to one another (Martin, 1949). This can

F i g u r e 6. Drawings showing changes in pigmentation and body form with

postlarval development in Sphyraena barracuda from 5.5 mm to 11.9 mm
standard length. A. 5.5 mm standard length; collected by R /S DANA,
Station 1293-V, 17°43'N, 64°56'W, April 17, 1922. B. 6.6 mm standard
length; collected by R /S DANA, Station 952, 17°55'N, 64°48'W, May 12,
1921. C. 8.6 mm standard length; collected by R /S DA N A , Station 1352-V,
35°42'N, 73°43/W, May 21, 1922. D. 11.9 mm standard length; collected by
Donald P. de Sylva, 1 mile southwest of the harbor entrance of North Bimini,
Bahamas, June 6, 1956.
36
seldom be disclosed by an examination of the regression on standard length.
Figure 2 indicates that of the proportional measurements taken, only
preanal length and pectoral fin length grow isometrically with an increase

7. Drawings showing changes in pigmentation and body form with

F ig u r e
postlarval development in Sphyraena barracuda from 17.2 mm to 790 mm
standard length. A. 17.2 mm standard length; collected by Donald P. de Sylva,
1 mile southwest of the harbor entrance of North Bimini, Bahamas, May 12,
1956. B. 23.7 mm standard length; collected by Donald P. and Doris D. de
Sylva, at beach east of Lerner Marine Laboratory, Bimini, Bahamas, July 7,
1956. C. 213 mm standard length; collected by David K. Caldwell, Spanish
Harbor Key, Monroe County, Florida, June 7, 1956; University of Florida
No. 7072. D. 790 mm standard length; collected by Doris D. de Sylva, north
of North Bimini, Bahamas, 25°48'N, 79°17'W, July 18, 1956.
37
in the length of the fish (Martin, 1949). Most pronounced allometry is
seen in changes in the first predorsal, prepectoral, and prepelvic distances,
and in the head and pelvic fin lengths. As might be expected, inflections
are most noticeable in the growth curve in the younger stages; growth
proceeds at a nearly constant rate with increase in body length over most of
the size range of the barracuda after it has reached 20 mm.
In the following discussion of the development of barracuda, reference
is made to Figures 2 to 7; body length refers to standard length unless
otherwise stated. Original measurements are presented by de Sylva (1958:
332-345).
In the smallest specimen examined, 5.5 mm (measured to the base of
the urostyle in this specimen), the yolk has evidently been absorbed for
some time, and the teeth are comparatively well developed. The stomach
of this specimen contained fish fragments, indicating that at this stage it is
already predatory. Orton (1955) noted that the California barracuda,
Sphyraena argentea, fed at an age of four days, and if early postlarval
growth is comparable for argentea and barracuda, the specimen of barra­
cuda depicted in Figure 6A might perhaps be about six or seven days old.
The snout and maxillaries are extremely short, with the latter represent­
ing 12.2 per cent of the length. The snout profile is sigmoid in outline and
the snout tip is rounded. The large eye represents 24 per cent of the head
length, which in turn is 36.6 per cent of the body length. The body is
relatively deep at this stage, and represents 42 per cent of the head length
and 15.4 per cent of the body length. Pigmentation on the head occurs as
scattered melanophores on the preorbital region and on the tip of the
lower jaw, as well as scattered dots on the nape which extend down to
the upper margin of the opercular series. The branchiostegal rays are
well developed, and are visible through the inter- and subopercle. A
fleshy pedicel forms the base of the rounded, membranous pectoral fin,
and there are no developed fin rays.
Scattered pigmentation is seen along the dorsal surface and as a series
of concentrated melanophores along either side of the anal fin base (Fig.
5E). There is evidence of a previously well-developed finfold, and at this
stage a few dorsal rays are visible. Neither the spinous dorsal nor the
pectoral fin is formed, and the anal fin appears only as a transoarent
membrane, with no traces of supporting rays. Myomeres are visible and
can be counted under strong, reflected light, their number being 14 -f- 10
= 24. The vertebral column is indistinct, except where it turns dorsad as a
urostyle to form the precursor of the hypural plate. The caudal fin bears
no resemblance to that of the next illustrated stage (Fi^. 6B) or that of
the adult (Fig. 7D ), and, in fact, is somewhat heterocercal, with the lower
lobe the longer.
At 6.6 mm (Fig. 6B) the eye has noticeably decreased in size, and
represents 19.3 per cent of the head length. Relative depth has increased
slightly, as has the pectoral fin length, the base of which is still evident
38
as a fleshy pedicel. The snout has lengthened noticeably, from 12.2 to
18.1 per cent of the body length. The maxillaries are well developed
and are expanded at the posterior margin. Preorbital pigmentation has
become still more distinct, although the supraoccipital melanophores, at
least in this specimen, are fewer than in the earlier stage. The sub- and
interopercular bones are beginning to increase in size and extend further
ventrad. Body pigmentation has increased considerably, but is still concen­
trated along the middorsal and midventral lines, and a band of melano­
phores occurs as a broken line along the posterior part of the trunk
along the lateral line. The pectoral, dorsal, and anal fin rays are fully
developed in number, and segmentation at the tip of the posteriormost rays
is now visible in the rays of the vertical fins. The bases of the second
dorsal and anal fins are enclosed in a fleshy sheath which extends about
one-third of the height of the rays. The urostyle has now become nearly
vertical and the hypural plate is fully formed, although a trace of the
upturned urostyle remains. Only a slight indentation is seen in the caudal
margin.
Little change occurs between 6.6 mm and 8.6 mm (Fig. 6C) except
that the shape of the snout is nearly straight in lateral view. The relative
decrease in depth is not as great as the figure indicates, but there is a
decrease in the size of the eye and a slight increase in the length of the
head, snout, and maxillaries. At this stage the maxillaries reach their
maximum mean length relative to the body length. The fish closely
resembles the adult in general body outline, notwithstanding the absence
of the pelvic and spinous dorsal fins, and the teeth are longer and more
robust. The anal and soft dorsal fins are relatively longer, and the fleshy
pedicel has diminished considerably and its base is widened. Preorbital
pigmentation assumes a definitely triangular shape, and a dusky, obscure
patch appears along the hyomandibular-preopercular area. Trunk pigmen­
tation is concentrated posteriorly along the lateral line, and at the
middorsal and midventral sections; the latter pigmentation has increased
and has spread anteriad. Branching and the formation of striae in the outer
rays of the caudal, soft dorsal, and anal fin have increased considerably,
and striae in the pectoral fin rays are first evident at this stage. The
urostyle is still visible through the flesh, but the increased thickness of
the flesh obscures the myomere count in the anterior trunk section.
At 11.9 mm a noticeable change has occurred and the fish has
assumed distinctive juvenile characters (Fig. 6D). The pelvic fins are
noticeable and represent 2.3 per cent of the body length; they appear
first at 8.8 to 10.1 mm.
The spinous dorsal fin first appears at between 10.1 and 11.9 mm; its
depressed spines represent 1.9 per cent of the body length when 11.9 mm.
The head is relatively longer than in previous stages, and the eye corres­
pondingly smaller. Cephalic pigmentation is marked about the snout,
supraoccipital region, and along the pre- and postocular regions, with
39
suggestions of an incipient band along the head from the snout to the
preopercle. The sub- and interopercle have completely covered the
branchiostegal tips, but the latter bones are still visible through the
opercular bones. Although middorsal pigmentation is more extensive than
at the previous stages, the conspicuous changes are seen in the anteriad
encroachment of the melanophores along the lower sides of the body which
appear as small, indistinct blotches, rather than as generalized, stellate
melanophores. Further cross striations occur in the tips of the fin, and
first sign of branching in these rays is seen; also, there is a definite
indication of a bifurcation of the caudal fin.
At 17.2 mm (Fig. 7A ), relatively few changes have occurred but it
is at this stage when the minimum depth, as a percentage of body length,
and a maximum prepectoral length are encountered. Also, there has
been a rapid lengthening of the soinous dorsal and pelvic fins, and the
rays of the latter have already begun to show striae at their tips. The
melanophores are now more diffuse along the cephalic rerion. These
coalesce along the snout to become confluent with the lateral pigmentation
of the trunk, suggesting a diffuse, longitudinal band; the contrast between
this dark band and the whitish ventral area is striking. The caudal fin
has assumed a definite bifurcate appearance with acute lobes appearing as
a transitional stage between those illustrated in Figures 6D and 7B.
At 23.7 mm (Fig. 7B), the lateral band of melanophores is beginning
to break up on the interradial membrane. The soft dorsal and anal fin
assume a juvenile shape and are well formed. Relative maxima are seen
in the lengths of the prepelvic, head, and snout. The rate of increase in
the length of the depressed dorsal spines has dropped off rather sharply.
Deep forking of the caudal fin is noticeable, and it has reverted to the
rounded-lobe situation. The teeth are longer and sturdier, and there is
evidence of specialization of the anteriormost teeth into longer canines.
No scales are visible in the fish until about 25 mm standard length is
reached, these first appearing along the sides of the caudal peduncle. Some
keeled scales are observed at a size range of about 50 to 70 mm along the
lateral-line section of the caudal peduncle, but are less noticeable than in
borealis (see Nichols and Breder, 1927:73).
Noticeable changes other than in proportional measurements and in the
extent of scalation and color pattern occur between the stage illustrated
in Figure 7B and that of the 213-mm specimen of Figure 1C. A transition
occurs in the pattern from a partially broken longitudinal band through a
series of H-shaped blotches prior to attaining the adult pattern. The
horizontal connections between these are eventually lost and form oblique,
wavy bands along the upper sides and back. The soft dorsal, anal, and
caudal lobes may show white tips, and the pectoral fin is pale to dusky.
The pelvic fin has its central rays blackish with white margins.
Between the two successive stages depicted in Figures 1C and 7D
numerous inky blotches appear on the caudal peduncle and along the
40
lower sides. These are variable in number and in placement, and are
characteristic of barracuda; generally they do not appear in fish less than
200 mm.
Substantial proportional decreases are seen in Figure 7D in the relative
size of the eye, head, prepelvic, and first and second predorsal length, with
a slight decrease in the relative length of the pelvic fin and the maxillaries.
Slight increases are noted in the relative depth of the caudal peduncle and
in the greatest depth, while the rate of change in the pectoral length and
the interorbital remains unchanged between Figures 7C and 7D. For
excellent photographs of living specimens of barracuda about 1 to 2 inches
long, see Gordon (1946) and Breder (1949).
The adult is represented by an illustration of a specimen 790 mm
standard length (Fig. 7D). The drawing was made from a color trans­
parency of a freshly caught fish which was not lying flat and too much of
the ventral aspect is visible. An authentic drawing by H. L. Todd is
seen in Goode (1884, PI. 178).
Noticeable differences between the adult and the juvenile color pattern
shown in Figures 7C and 7D, respectively, are essentially in the transition
from indistinct, wavy bands on the back to a series of 18 to 22 crossbars
which are inclined obliquely backwards. Each of these bars may appear
solid, or each may be broken up into two, nearly separate, ovoid blotches.
The white tips of the soft dorsal, anal, and caudal lobes which are
vague in the juvenile (Fig. 7C) are distinct and diagnostic in the adult
and become more marked with age. The upper margin of the opercle is
dusky to black, and the pectoral fin is dusky, with its tip and ventral margin
pale. The inky blotches are generally well defined in large specimens, but
it should be noted that the two blotches just behind the pelvics in Figure
7D are atypical, and usually do not occur on the white ventral coloration;
these are generally confined to the sides below the lateral line and above
the line dividing the lateral from the ventral coloration. Photographs in
Ackerman (1952) show the variation in the number and position of these
blotches.
Significant proportional changes which occur between the juvenile and
the adult (Figs. 2 and 3) are seen in the decreased relative lengths of the
prepelvic distance, head, prepectoral, and diameter of the eye. A somewhat
lesser rate of change with increased leneth is observed in the leneth of the
snout, length of spinous dorsal fin, and pelvic fin. Nearly isometric growth
between the two stages is observed for the length of the preanal, the depth
of the body, and caudal peduncle, whereas first and second predorsal
lengths show a relative increase in body length, with an inflection occurring
between 600 and 800 mm. An increase in the interorbital width and in
the length of the pectoral fin is observed in Figures 2D and 2L.
Noticeable in Figures 1C and 7D is the change in the tail from a forked
to a scalloped margin. Three more or less semicircular indentations are
observed which give the margin a quadrilobed appearance; this characteris­
41
tic is even more noticeable in larger specimens. This was noted by
Riippell (1835) for agam and by Bleeker (1854b) for commersoni, both
of which are synonyms of barracuda Walbaum.
An additional difference between the juvenile and adult is the increase
in the length of the maxillaries, which range in length from a vertical
line passing through the anterior border of the eye in the juvenile to a
line passing through the posterior margin of the eye in a very large adult.
In the early stages the maxillaries fall far short of the anterior border of
the eye.
Discussion .— Allometric growth, or heterauxesis (Martin, 1949), is notice­
able in barracuda , and for most characters, until a length of about 20 mm
is attained, there is a general tendency toward positive allometry (tachy-
auxesis) after which point negative allometry (bradyauxesis) is encounter­
ed. Greatest negative allometry is seen, after the 20-mm stage is reached,
in the relative length of the head, maxillaries, snout, prepectoral, first pre­
dorsal, and prepelvic, and in eye diameter; positive allometry occurs in
the body depth, interorbital width, and pectoral fin length. Isometric
growth occurs only in preanal length. By using head length as a base
length (Fig. 3) it is possible to fit to these data an assortment of parabolic,
hyperbolic, and third-degree polynomial curves, but since head length
shows negative allometry itself, such curves are difficult to visualize in
terms of body growth.
The inflections noted in the growth data may reflect changes in the life
history of the barracuda. Although ecological aspects are treated under a
subsequent heading, it seems appropriate to interject here some comments
on the distribution of the young barracuda. It appears that most spawning
occurs at a considerable distance from land, probably over deep water
between the deeper reefs and the edge of the continental slope, and that
the young are subsequently transported both by major currents to other
regions and inshore by tidal movement. Postlarvae up to about 12 mm
occur far out to sea but only barracuda 15 mm long and above evidently
occur in littoral waters. It is suggested that during this shoreward move­
ment they undergo a moderate transformation which includes a rather
sudden increase in the extent of pigmentation, the lengthening of the teeth,
the appearance of the spinous dorsal and pelvic fins, as well as a rapid
development in function and structure of the vertical fins, the pelvic, and
caudal fins. By the time it has attained a length of 18 to 22 mm, the
barracuda has generally adapted to a littoral life; it is at this transition
in habits and ecology when the graphs show an inflection. Sexual maturity,
which occurs at a fork length of 460 to 660 mm, does not appear to be
reflected in the graphs (Figs. 2 and 3).
Description and Size of the A dults. — Jordan and Evermann (1896)
presented an extensive description of the adult barracuda. Adequate,
supplementary descriptions of a qualitative nature are available for both
42
the young and adult fish in Breder (1948a, 1949), Bleeker (1854b),
Day (1875-1878, 1889), Fowler (1925), Gilchrist and Thompson (1909),
Meek and Newland (1884), Meek and Hildebrand (1923), Poey (1858),
Schultz (1943, 1953), Smith (1956), Weber and de Beaufort (1922),
and Whitley (1947). Quantitative characters are listed in Tables 1 and 2.
The maximum size attained by the adult barracuda has been one of
speculation. Most estimates of the great size attributed to this species
can be traced to the writings of early naturalists. Labat (1724) reported
barracuda measuring 18 to 20 feet long from the Antilles, and Fermin
(1769) recorded specimens 15 feet long from the coast of Surinam.
Btittikofer (1890) recorded a 10-foot specimen of S. jello from Liberia,
and Vincent (1910) measured a specimen of barracuda from Trinidad
which was 8 V2 feet long. Popular stories often relate the sighting of 7- or
8-foot barracuda in the Florida Keys, but such fish must be uncommon,
for no fish longer than 63 inches total length occurred in the records of
Mr. A1 Pflueger from 1953 through 1957.
The largest fish for which I have measurements was a female caught
off Miami Beach on March 18, 1956, which weighed 39 pounds and 9
ounces and was 55.5 inches total length. Gudger’s (1918) 38-pound
barracuda was 55 inches long, and Herre (1936) reported a specimen
of barracuda from the Philippines which was 62 inches long and weighed
25 kilograms (55 pounds). Fish weighing upwards of 50 pounds are
evidently not rare in the Florida Keys, although I have seen none under­
water, either in the Florida Keys or in the Bahamas, which would exceed
40 or 50 pounds. These giants are not often caught by anglers but
are occasionally taken by skilled spearfishermen. Fish weighing 45-50
pounds seem to represent the maximum weight entered in local fishing
tournaments in Miami during recent years.
The world-record barracuda caught on hook and line which is recog­
nized by the International Game Fish Association (LaMonte, 1952)
weighed 103Va pounds and was 66 inches in total length, with a girth of
3 1 ^ inches. Although it was only 4 inches longer than the specimen
reported by Herre (1936), it weighed nearly twice as much. This record
fish was taken at West End, Bahamas, in 1932 by Mr. C. E. Benet.
However, Herre (1936) reported a 106-pound specimen of barracuda
caught with rod and reel by General Leonard Wood, off Cape San
Agustm, Mindanao, Philippines.
R e p r o d u c t io n

Inferences on the probable time of spawning of the barracuda were

deduced from the occurrence of the young by Breder (1949) and Erdman
(1956). Malpas (1926) and Williams (1956) examined gonad samples
from S. commersoni ( =S. barracuda) from the Indian Ocean. Goode
(1876) mentioned that S. picuda (= 5 . barracuda) commonly spawned
in Bermuda, and Holder (1924) stated that barracuda in the Florida Keys
43
 8. Internal organs in situ of a ripe male specimen of Sphyraena barra­
F ig u r e
cuda, 827 mm fork length, collected from Bimini, Bahamas, on July 7, 1956.
Scale two-fifths natural size. SV = sinus venosus; A T = atrium; V = ventricle;
B A = bulbus arteriosus; L = liver; P C = pyloric caecae; G B = gall bladder;
SP = spleen; ST = stomach; SB = swim bladder; F B = fat bodies; D T = diges­
tive tract; T = testis; A O = aorta.
44
 TABLE 3
C r it e r ia o f d e v e l o p m e n t a l st a g e s in t h e g o n a d s o f Sphyraena barracuda
Females
Stage 1 “Resting”; ovary full of immature eggs only, not showing up as distinct
granules; ovary pink (young individuals) to reddish (older indivi­
duals), gelatinous and translucent; blood vessels generally not visible;
lumen easily visible.
Stage 2 Ovary ripening, full of maturing eggs; pinkness fading to a yellowish
color; ovary opaque; blood vessels beginning to appear as faint, reddish,
transverse stripes; transverse folds beginning to appear.
Stage 3 Ovary nearly ripe; small eggs are visible which are attached to the
ovarian membrane but are not extrusible from membrane with ease;
blood vessels bright red, gorged with blood; lumen completely ob­
scured; transverse folds well developed.
Stage 4 Ovary ripe; eggs larger than stage 3, easily extrusible from ovarian
membrane; blood vessels bright reddish-purple to deep purple, gorged
with blood; ovary a very opaque white; lumen completely obscured;
transverse folds well developed.
Stage 5 Ovary spent, a flaccid bag with a few mature eggs occasionally adhering
to the walls; lumen evident; slimy consistency; veins still purplish but
less turgid; transverse folds obscure; ovary has lost its whitish color
and becomes redder as it approaches stage 1; often difficult to distin­
guish between late stages of 5 and early stages of 1.

Males
Stage 1 Testes whitish-brown, slender, firm in texture; blood vessels not de­
veloped on surface; firm to the touch.
Stage 2 Testes white and thicker than stage 1, less firm in texture; a few blood
vessels developed on surface, but milt does not flow out when testes
are pressed.
Stage 3 Testes thicker than stage 2; blood vessels further developed; milt flows
out readily when ducts are pressed.
Stage 4 Spent; testes slender and flaccid;blood vessels reduced;milt not present.

spawned in the “early spring.” Fowler (1905) collected young barracuda

from New Jersey, and (1945) recorded specimens to 2 inches in length
from Key West, Florida, in December. LaMonte (1951) reported that
“the United States Fish and Wildlife Service at Beaufort [North Carolina]
reports many barracuda from six to eight inches long, at which size they
are mature off the Beaufort-Morehead City section . . . .” However, it
seems that the species referred to here is confused with another sphyraenid,
since no mature individual of S. barracuda less than 20 inches long has
been examined by me from southern Florida waters (Fig. 10). Extensive
studies on the spawning cycle have been made by Walford (1932) on
S. argentea and by Manacop (1936) on S. jello and S. obtusata from the
Philippines.
Sexual Dimorphism.— There is no sexual dimorphism in general appear­
ance and no appreciable difference in the sexes in the length-weight
45
F ig u r e 9. Histological sections of the gonads of Sphyraena barracuda. A. Sec­
tion of immature ovary (x39), 544 mm, collected July 18, 1956. B. Section of
nearly ripe (stage 3) ovary (x5.5), 1208 mm, collected July 8, 1955. C. Section
of ripe ovary (x5.3), 1149 mm, collected July 9, 1955. D. Section of ripe
testis (x35), 971 mm, collected June 8, 1956. Measurements are fork lengths.
Specimens represented in A, B, and C were collected at Bimini, Bahamas, and
specimen represented in D was collected at Cat Cay, Bahamas. Photographs
by Douglass M. Payne.
46
relations. The sexes may be identified by macroscopic examination of the
gonads. The ovaries are pinkish in color, circular in cross section at all
stages of development, and the left is always longer. When a fresh ovary
is cut transversely and placed in water, the edges tend to curl. The testis
is creamy to whitish-brown, triangular to subtriangular in cross section,
and does not curl when a cut section is placed in water. The testes are
approximately equal in length. The location of the reproductive organs is
shown in Figure 8.
Criteria for Stages of Sexual Development.— The criteria given in Table
3, modified somewhat from Walford’s (1932) study of S. argentea, have
been found to be satisfactory for identifying the various stages of gonadal
development in S. barracuda.
Gonads were preserved in Bouin’s fluid and subsequently stained with
toluidine blue and picric acid. Photographs of histological sections of a ripe
male (stage 3) and stages 1,3, and 4 of a female are presented as Figure 9.
Age and Size at Maturity.— The relation between the onset of maturity
and length of barracuda is shown in Figure 10. Despite a scarcity of
specimens smaller than about 400 mm, it is evident that the male does
not mature sexually until it attains a fork length of about 460 mm. Most
male barracuda mature by the time they have reached about 500 mm and
an age of two years. Some females first mature at three years at a length
of about 580 mm; most do not mature until an age of four years at a
length of about 660 mm. Malpas (1926) found ripe specimens of S.
commersoni ( = barracuda) from Ceylon which measured from 350 to

TABLE 4
Sex ratio o f 463 s p e c im e n s o fSphyraena barracuda l a n d e d a t P ie r 5,
M ia m i , F lo r id a , from A u g u s t , 1955, t h r o u g h A u g u s t , 1956.
Month Number Number of Number of Per cent Per cent Ratio,
examined females males females males females: males
August, 1955 16 6 10 37.5 62.5 0.6:1
September 41 26 15 63.5 36.5 1.7:1
October 30 12 18 40.0 60.0 0.7:1
November 32 20 12 62.5 37.5 1.7:1
December 26 17 9 65.4 34.6 1.9:1
January, 1956 39 18 21 46.2 53.8 0.9:1
February 14 13 1 92.9 7.1 13.0:1
March 28 24 4 85.7 14.3 6.0:1
April 14 14 — 100.0 — —
May 22 13 9 59.1 40.9 1.4:1
June 36 31 5 86.1 13.9 6.0:1
July 36 17 19 47.3 52.7 0.9:1
August 129 76 53 58.9 41.1 1.4:1
Total 463 287 176 — — —
No. per month 35.6 22.1 13.5 62.0 38.0 1.6:1

47
 FORK LENGTH, MM
0 300 500 700 900 1100
■ « 1__ i__ I__ i__ i__ i _ i _ I __ i__ — i i i I fc i

jz Q . -i=P- i_ AUG.

-R = r i SEP.

--- 1 □ arfwlrj-T- OCT.

: nov.

0300 1100
FORK LENGTH, MM
10. Relation between size and maturity of Sphyraena barracuda. Speci­
F ig u r e
mens were collected from Pier 5, Miami, Florida, from August, 1955, through
August, 1956. Blocks above horizontal axes are male fish; those below axes are
female fish. Shaded blocks represent fish in spawning condition; open blocks
represent non-spawning fish.
48
410 mm. If these specimens were indeed barracuda, they matured much
earlier than this species does in the western North Atlantic.
Sex Ratio.— The sport fishery at Pier 5, Miami, from August, 1955,
through August, 1956, yielded 287 females and 176 males, a ratio of
8 to 5 (Table 4). Monthly fluctuations in the sex ratio occurred from one
month to the next, but the paucity of data for some months precludes
analysis of the data. From February through April, five males and 51
females were taken. It is thought that this marked change in sex ratio
from that observed in January may reflect a pre-spawning migration in
the males, so that they are not available to the angler. During the months
when the males outnumbered the females the ratio was lower than
when the reverse was true.
%
Spawning Period.— The percentage of individuals from the angler catch
TABLE 5
M o n t h l y p e r c e n t a g e o f 456 s p e c i m e n s o f Sphyraena barracuda l a n d e d
a t P i e r 5, M i a m i , F l o r id a , f r o m A u g u s t , 1955, t h r o u g h A u g u s t , 1956,
w h i c h e x h i b i t e d v a r io u s s t a g e s o f r e p r o d u c t i v e a c t i v i t y .
N u m b e r o f in d iv id u a l s r e p r e s e n t e d in p a r e n t h e s e s .

Males Stage 1 2 3 (Ripe) 4 (Spent) Total

Aug. 12.5 (1) 12.5 (1) 75.0 (6) — (8)
Sept. 40.0 (6) 6.7 (1) 53.3 (8) — (15)
Oct. 35.3 (6) — 64.7 (11) — (17)
Nov. 100.0 (12) — — — (12)
Dec. 100.0 (9) — — — (9)
Jan. 100.0 (21) — — — (21)
Feb. — 100.0 (1) — — (1)
Mar. 33.3 (1) — 66.7 (2) — (3)
May — — 100.0 (9) — (9)
June — — 100.0 (4) — (4)
July — — 89.5 (17) 10.5 (2) (19)
Aug. — — 89.8 (44) 10.2 (5) (49)
Total — (56) — (3) —(102) — (7) (167)
Females Stage 1 2 3 4 (Ripe) 5 (Spent) Total
Aug. 25.0 (1) — 75.0 (3) — — (4)
Sep. 42.3 (11) 15.4 (4) 19.2 (5) 23.1 (6) — (26)
Oct. 75.0 (9) --- — 8.3 (1) 16.7 (2) (12)
Nov. 95.2 (20) 4.8 (1) — — — (21)
Dec. 100.0 (17) — — — (17)
Jan. 100.0 (18) — — — — (18)
Feb. 84.6 (11) 15.4 (2) — — — (13)
Mar. 60.7 (17) 28.6 (8) 10.7 (3) — — (28)
Apr. 64.3 (9) 7.1 (1) 21.5 (3) 7.1 (1) — (14)
May 7.1 (1) 21.5 (3) 42.8 (6) 21.5 (3) 7.1 (1) (14)
June 6.5 (2) 32.3(10) 22.6 (7) 19.3 (6) 19.3 (6) (31)
July 25.0 (4) 6.3 (1) 12.5 (2) 31.2 (5) 25.0 (4) (16)
Aug. 28.0 (21) 6.7 (5) 16.0 (12) 24.0(18) 25.3 (19) (75)
Total — (141) — (35) — (41) — (40) — (32) (289)
49
throughout the year landed at Pier 5, Miami, showing a particular stage
of reproductive activity is given in Table 5. Individuals in a spawning
condition were first observed in the samples when the mean water tempera­
ture reaches 73-74°F (Bumpus, 1957) for the Miami region (Fig. 11).
The first ripe male barracuda, measuring 1035 mm fork length, was
observed on March 31, 1956. By May, 11 per cent of the males sampled
were ripe. In July and August a few male barracuda were taken which
had recently ceased spawning (stage 5). By October, 64.7 per cent
of the males were still ripe; by November, all males had completed their
spawning activities. There appeared to be no peak time of spawning in
the males, as milt could be expressed from the testes of mature males
throughout the spawning season.
Females ripen sexually at a later date than the males, and cease to be
ripe earlier in the year (Table 5; Fig. 11). The first ripe female was
taken on April 18, 1956, measuring 1002 mm fork length, and was the only
ripe female observed in the April samples. The larger females appeared
to ripen earlier, as indicated in the data for May (Fig. 10). In May,
21.5 per cent of the females were in a spawning condition. By mid-
October, 91.7 per cent of the females had completed their spawning.
Reproductive activities stopped toward the end of October. Spawning ceased
when the water temperature had dropped from a mean maximum summer
temperature of 83.8° to 77 °F.

UJ
cr
3
S
cr
UJ
o.
2
UJ
I—
cr
UJ

UJ

F ig u r e 11. Relation between spawning season of Sphyraena barracuda and

mean water temperatures adjacent to Miami, Florida. Solid line represents per­
centage of males in spawning condition (N = 2 0 3 ); broken line represents per­
centage of females in spawning condition (N = 2 8 4 ). Dotted line is monthly
mean water temperature at Fowey Rocks Lighthouse, southeast of Miami,
Florida. Temperature data from Bumpus (1957).
50
 The peak of spawning in the female, as indicated by the greatest
percentage of ripe females, occurred in July, and secondary peaks were
observed in the samples taken in May and September. Relatively few
females in the catch are ripe at a given time. Periods of greater spawning
activities are evidently reflected in the occurrence of peaks in the growth
of the young-of-the-year.
Postlarvae were taken from December through June in the Virgin
Islands area (Table 6). Fowler (1945) recorded specimens which
measured 33, 36, and 50 mm long from Key West as late as December.
Beebe and Tee-Van (1928) reported barracuda from Haiti measuring
20 to 26 mm which were taken from March 6 to May 8; the former
date is nearly a month earlier than the date when fish of a similar size
were collected from Florida waters.
Williams (1956) found ripe individuals of S. commersoni ( ^barracuda)
in East African waters from September to May. Ommanney (1953) took
specimens of S. commersoni from the Seychelles Islands with nearly ripe
eggs in May, and ripe females occurred in November; ripe males were
taken in December.
Frequency and Extent of Spawning.— Kisselevich (1923) used the fre­
quency distribution of ova diameters to ascertain the frequency and extent
of spawning in clupeid fishes, and Walford (1932) successfully utilized
this method for Sphyraena argentea. Hickling and Rutenberg (1936)
postulated that the breeding cycle of a fish could be predicted by an
analysis of the ova diameters, and that species having a sharply dem ar­
cated spawning period could be distinguished from those spawning over
a long period. In the first instance the frequency polygons of the ova
diameters are steep and well separated from one another, whereas those
of a fish having an extended spawning period are characterized by a series
of low-peaked polygons which are not trenchantly different.
TABLE 6
L e n g t h f r e q u e n c i e s o f 73 p o s t l a r v a l s p e c i m e n s o f Sphyraena barracuda
COLLECTED BY THE DANA EXPEDITIONS FROM JUNE, 1920, THROUGH MAY,
1922, i n t h e V ir g in I s l a n d s , J a m a i c a , M a r t i n i q u e , D o m i n i c a ,
N o r t h C a r o l in a , a n d A n d r o s I s l a n d , B a h a m a s .
Standard length December January March April May June
(mm) 1921 1922 1922 1922 1922 1922
5 — — 2 3 — —
6 2 — 2 5 1 —
7 1 1 3 3 2 1
8 1 2 6 4 1 —
9 1 3 4 — 1 —
10 — 5 5 4 1 —
11
11 a
D
12 1 _ 3 1 1 __
Total 6 11 28 20 7 1
Mean length 8.4 9.1 8.8 7.5 8.3 7.0
51
 In order to determine first if the eggs in one part of the barracuda’s
ripe ovary were larger than in another part, 559 ova diameters were
measured from the ovary of a ripe female which measured 1077 mm fork
length (Fig. 12 A-D). Ova showing varying stages of maturity appear to
be similar in size, regardless of where they were taken, since the peaks
of the polygons correspond in each succeeding section. Carbine (1944)
found that the middle part of the ovary of the northern pike, Esox lucius,

OVA DIAMETERS, MM
F ig u r e 12. Frequency distribution of 559 ova diameters from different sections
of a ripe ovary (stage 4) of a 1077-mm specimen of Sphyraena barracuda
collected at Pier 5, Miami, Florida, on August 22, 1956. A. Posterior end.
B. Anterior end. C. Between anterior end and midsection. D. Midsection. E.
Sum of frequencies of ova diameters from sections A-D.
52
contained significantly larger eggs than did the anterior or posterior sections.
Although the primary and secondary oocytes are sharply delineated
(Fig. 12E), ova larger than about 0.5 mm in diameter form a series of
low peaks which are not sharply marked. This suggests an almost continu­
ous supply of gradually maturing eggs which contribute to the general
stock of eggs throughout the spawning season. Walford (1932) found
evidence that a single female of S. argentea spawned several times per
season. The extended spawning season postulated here for S. barracuda
is corroborated by an examination of the gonads throughout the year
(Table 5), and by the abundance of barracuda less than 1 inch long
from April to November from southern Florida waters (Fig. 18). Erdman
(1956) noted the presence of young barracuda over the same period
in Puerto Rican waters.
Fecundity.— Walford (1932) calculated the number of eggs produced
by a single female of S. argentea. He was able to separate mature and
immature eggs in the ovary of S. argentea, but this distinction generally
is not marked in barracuda (Fig. 12). However, in some large individuals
of barracuda, the differences among the ova diameters were sufficient to
permit a rough calculation of fecundity. The number of eggs in a sample
was counted, weighed, and the number of eggs per gram calculated. A
barracuda 895 mm fork length was estimated to contain about 560,000
mature eggs, and one measuring 1011 mm fork length held about 670,000
mature eggs. However, the assessment of fecundity in a species which
spawns several times over an extended period seems to have little
biological value unless it could be determined how many eggs were
spawned in a season.
Eggs.—Ripe, translucent eggs were found in the ovaries only twice in the
examination of 68 female barracuda from which eggs could be expressed
from the body cavity. These translucent eggs measured from 0.74 to
0.81 mm in diameter, and are considerably smaller than the mature
ovarian eggs of S. argentea which measure from 1.02 mm (Barnhart,
1927) to 1.14 to 1.6 mm in diameter (Walford, 1932; Orton, 1955).
The modes of the peaks shown in Figure 12 occur at approximately
0.14 mm, 0.30 mm, 0.48 mm, and 0.70 mm. Evidently, the ovary of this
individual either was not dead ripe, or some eggs had already been spawn­
ed, as most eggs depicted (Fig. 12) are smaller than the translucent ones.
Barnhart (1927) and Orton (1955) stated that the egg of S. argentea
is pelagic. This was also noted for Sphyraena (sphyraena?) by Nikol’skii
(1954: 401), but Delsman (1930) believed Sphyraena (species?) has
. . such big eggs in the ripe ovary that one may conclude at once that
the eggs are not pelagic.”
Numerous attempts were made to fertilize and hatch eees artifically
using the methods of Morris (1954) and Oppenheimer (1955), but all
experiments failed.
53
 Distribution of the Postlarvae in Relation to Hydrographic and Meteoro­
logical Conditions.— Hydrographic and meteorological observations at
stations occupied by the DANA vessels from 1920 to 1922 (Schmidt,
1929) at which postlarval specimens of S. barracuda were obtained are
given in Table 7. H ydrographic methods and collecting gear are

TABLE 7
H y d r o g r a p h ic and m e t e o r o l o g ic a l o b se r v a t io n s at s t a t io n s o c c u p ie d
w here po stla r v a l s p e c im e n s o f Sphyraena barracuda w ere co llected by
DANA e x p e d it io n s to the t r o p ic a l w ester n A t l a n t ic O cean fro m
1920 to 1922. D ata are s u m m a r iz e d fro m S c h m id t (1929).
SampleWater Wind
Station depth, depth, Wea- and
number Date Latitude Longitude Hour meters meters ther force
848 6/ 6/20 18°00'N 64041^ 0300 300 (4 0 0 0 )a cloudy ESE 4
952 5/12/21 17°55'N 64048^ 2000 100 (4800) squally ESE 4-5
952 5/12/21 17°55'N 64048^ 2300 200 (4800) squally ESE 4-5
952 5/13/21 17°55'N 64048^ 0200 400 (4800) squally ESE 4-5
1186X 12/ 1/21 17°59'N 64041^ 0045 50 3160 cloudy NE 3
1190 VIII 12/13/21 17°59'N 64045^ 0400 100 3160 cloudy calm
1190IX 12/13/21 17°59™ 64045^ 0400 50 3160 cloudy calm
1191III 12/14/21 17°49'N 64054^ 1850 100 (1000) cloudy ENE 3
1191VII 12/15/21 17°49'N 64054^ 0400 100 (1000) cloudy ENE 3
1192 VII 12/15/21 17043™ 64053^ 1935 100 410 squally SE
1195V 1/ 3/22 19°01'N 65023^ 0350 50 5250 cloudy NNE4
1216VII 1/29/22 18°22'N 78038^ 0100 0 (2000) cloudy NE 3
1256V 3/ 3/22 17043™ 64056^ 1920 100 (1000) cloudy ESE 6
1257V 3/ 6/22 17°43'N 64056^ 0010 100 (1000) cloudy E 4
1266bII 3/13/22 17°45'N 64055^ 1900 800 950 cloudy NE 4
1266bV 3/13/22 17°45'N 64056^ 1900 100 950 cloudy NE 4
1266VII 3/13/22 17°45'N 64056^ 2215 50 950 cloudy NE 4
1266VIII 3/13/22 17°45'N 64055^ 2215 25 950 cloudy NE 4
1270IV 3/16/22 17043'N 64056^ 1800 600 (1000) squally SE 3
1238IV 2/11/22 26°13'N 78048^ 0400 0 620 cloudy S 2
1273IV 3/24/22 17043'N 64056^ 1915 300 (1000) cloudy ENE 4
1273V 3/24/22 17043'N 64056^ 1915 100 (1000) cloudy ENE 4
1278V 3/29/22 17043'N 64056^ 1900 100 (1000) cloudy ENE 5
1280V 3/31/22 17043'N 64056^ 2015 100 (1000) cloudy ENE 4
1281X 4 / 1/22 17°43'N 64°56'W 2330 50 (1000) clear E 1
1284VI 4/ 6/22 14°38'N 6I 0I 6W 0300 0 ( 800) cloudy ENE 2
1286V 4/ 7/22 15°17'N 61029^ 2100 50 ( 200) squally ENE 3
1289X1 4/15/22 17043'N 64056^ 0400 50 (1000) squally E 3
1292VI 4/16/22 17043'N 64056^ 1930 50 (1000) cloudy ESE 3
1293V 4/17/22 17043'N 64056^ 1940 100 (1000) cloudy E 3
1293VI 4/17/22 17o43'N 64056^ 1940 50 (1000) cloudy E 3
13521 5/21/22 35042™ 73043^ 2130 300 (5000) cloudy SW 4
1352IV 5/21/22 35°42'N 75043^ 2130 100 (5000) cloudy SW 4
1352V 5/21/22 35042™ 75043^ 2130 50 (5000) cloudy SW 4
a Depths given in parentheses were estimated from U. S. Hydrographic Office
Chart 1290 and National Geographic Society “Map of the West Indies,” 1954.
54
 TABLE 7— (CONTINUED)

Sea Surface Number Other

Station and Temp., salin- Densi- 0 2 of species of
number force °C. ity, %0 ty, o-t m l/L specimens Sphyraena
848 SE swell 26.00 __ — — 1 _
952 ESE 4 26.50 — — — 3 —
952 ESE 4 26.50 — — — 3 —
952 ESE 4 26.50 — — — 1 —
1186X NE 3 27.05 34.98 22.72 4.51 1 —
1190 VIII E 3 — — — — 1 —
1190IX E 3 — — — — 2 —
1191III ENE 3 — — — — 1 —
1191VTI ENE 3 26.80 — — — 1 —
1192 VII SE — — — — 1 —
1195V NNE 4 26.03 35.46 23.39 4.46 7 guachancho (2)
1216VII NE 3 26.20 — — — 6 —
1256V SE 4 — — — — 3 picudilla (1)
1257V E 3 — — — — 1 picudilla (6)
1266bII SE swell 25.14 35.84 23.95 5.24 2 —
1266bV SE 25.14 35.84 23.95 5.24 2 picudilla (2)
1266VII SE 25.14 35.84 23.95 5.24 2 —
1266VIII SE 25.14 35.84 23.95 5.24 2 —
1270IV SE 3 — — — — 2 picudilla (2)
1238IV SE 2 23.60 — — — 1 —
1273IV ENE 3 — — — — 1 —
1273V ENE 3 — — — — 2 picudilla (1)
1278V ENE 4 24.70 — — — 4 picudilla (3)
guachancho (1)
1280V ENE 3 24.80 — — — 6 guachancho (1)
1281X E 1 24.90 — — — 11 picudilla (2)
guachancho (1)
1284VI NE swell 26.00 — — — 2 guachancho (1)
1286V E swell 26.10 — — — 1 picudilla (1)
1289X1 E 3 25.30 — — — 1 —
1292VI S swell 25.40 — — — 2 —
1293V S swell 25.40 — — — 1 —
1293VI S swell 25.40 — — — 5 —
13521 SE swell 25.60 36.08 23.99 — 3 —
1352IV SE swell 25.60 36.08 23.99 — 1 —
1352V SE swell 25.60 36.08 23.99 — 1 —

described by Schmidt (1929) and Jespersen and Taning (1934). All

tows were made at a vessel speed of 2 knots, and each lasted two hours,
with the exception of a 30-minute haul at Station 1271, a 90-minute
haul at Station 1191, and a 180-minute haul at Station 1186. All specimens
were taken with a 200-cm conical, stramin net, except for those collected
at Stations 1284-VI, 1292-VI, and 1293-VI, at which 150-cm nets were
operated, and at Station 1270-1V, at which a 150-cm silk net was operated
The maximum depth at which the net fished is given in Table 7, but since
the nets were not of the closing type, the actual depth of capture of the
55
postlarvae was not ascertained. The DANA vessels operated in the
tropical western Atlantic in June, 1920, May, 1921, and from November,
1921 through May, 1922. Postlarval specimens of barracuda were
collected during each month except November and February (Tables 6
and 7). Most were captured immediately north of St. Croix, Virgin
Islands, but a few were collected from the waters off Jamaica, Martinique,
Dominica, Andros Island, Bahamas, and Cape Hatteras, North Carolina.
When the frequency of postlarval specimens of barracuda taken by the
nets was plotted, two distinct peaks of abundance occur (Fig. 13): one
at 4 AM, the second at 7 PM. This would at first suggest a diurnal
migration. However, when the total number of stations occupied is
compared with the distribution of collections in which the barracuda occur­
red, it is apparent that the greatest number of barracuda was collected at
the hours when most collecting was undertaken. Even so, the almost
complete absence of barracuda in collections made during daylight hours
does not preclude the occurrence of a limited vertical migration. But barra­
cuda of all sizes are agile and speedy swimmers, and perhaps they are
able to dodge the approaching net in the daylight hours.
As might be expected in the open-ocean environment, only slight
differences in salinity and temperature were noted at the DANA stations
(Table 7); the surface salinity values were between 34.98 and 36.08%c.

CO
sO <
O
3
o o
y tc
oj S<

h i
K z
s 8

HOURS
13. Relation between hours at which postlarval specimens of Sphyraena
F ig u r e
barracuda were collected by plankton nets (dashed line) and hours at which
plankton nets were operated (solid line) by DANA vessels, 1920-1922, in the
tropical western Atlantic Ocean. Data are from Schmidt (1929).
56
Surface temperatures ranged from 23.60 to 26.80°C, and were typical
of the tropical western Atlantic. Dissolved oxygen values in surface
waters from the localities at which barracuda were collected, although
limited in the number of observations, were all high.
Information on the depths of water at the DANA stations is supple­
mented by U.S. Hydrographic Chart 1290, and the National Geographic
Society’s Map of the West Indies. All postlarval barracuda were taken over
depths greater than 200 meters, and the maximum depth was 5250 meters.
I collected postlarval barracuda measuring 11.9 to 17.9 mm near
Bimini, Bahamas, relatively far from shore. These were taken at the
surface by dipnet and plankton tows over depths of nearly 200 meters, at
a locality where a current rip exists between the littoral waters and the
Florida Current. Postlarval barracuda have also been taken by me from
shallow-water habitats. It is uncertain if these were spawned near the
location of their capture or had been transported there by currents and
tides. Weber (1913: 150-152) collected a 13-mm larva of Sphyraena
(species?) and four specimens of S. jello which measured 47 to 55 mm in
length, far offshore in the Halmaheira Sea, Dutch East Indies. Beebe and
Tee-Van (1928) found specimens of barracuda of 20-26 mm in length,
from March to May, which were collected several miles from Haiti. On
these bases, barracuda probably spawn over deep water near the juncture
between the oceanic and coastal circulation, so that the eggs and larvae
may be subsequently entrained in both coastal and offshore waters.
In two of the DANA Collection hauls, three sphyraenids from the
western North Atlantic occurred together; in these, picudilla and guachan-
cho were taken with barracuda , notwithstanding that the adults of these
three species have considerably different habitats and vertical distributions.
Possibly the postlarvae of each species occurred in a different stratum
or density layer in the water column and all were brought together as
the net was retrieved.
L e n g t h -w e ig h t R e l a t io n

Specimens from both Florida and Bimini taken over a 17-month period
were utilized in the calculation of the length-weight relation. Possible
racial and seasonal differences between the barracuda of these areas might
be included in the calculations. Using the lengths and weights of 59
females, 37 males, and 54 individuals of undetermined sex, the empirical
relation for the two measurements are presented in Figure 14. From these
data, the length-weight relation for the male is W = 1.56 X 10~6 L 2-84589,
and for the female is W = 2.67 X 10~6 L 2-92134. The relatively small
exponents observed here are characteristic of a slender species such as the
barracuda, which contrast with that of a chunkv, heavy-bodied fish such
as the little tuna, Euthynnus alletteratus, in which the exponent is 3.16
(de Sylva and Rathjen, 1961).
The calculated regression lines are approximately equal, except at about
57
 TOTAL LENGTH, INCHES

FORK L E N G T H , MM

F i g u r e 14. Length-weight relation of 150 specimens of Sphyraena barracuda

from Florida and Bimini, Bahamas, collected from April, 1955, through August,
1956. Open circles and solid line represent male fish (N = 3 7 ); solid dots and
broken line represent female fish (N = 5 9 ); triangles represent 54 fish of un­
determined sex, and for which the equation has not been calculated nor the
slope fitted to the points.
58
900 mm when the curves intercept. Males are heavier than the females at
the same length. The empirical data fit closely the calculated regressions
for smaller fish, although points representing fish greater than 600 mm are
more scattered about the slopes.
The following mean conversion factors have been calculated from the
length of over 200 barracuda which measured from 17 to 1111 mm
standard length from the western North Atlantic:
To Obtain: M ultiply by: To Obtain: M ultiply by:
Standard from fork 0.919 Fork from total 0.901
Standard from total 0.829 Total from standard 1.207
Fork from standard 1.089 Total from fork 1.110

A ge a n d G row th
With the exception of Walford’s (1932) study on the age of S. argentea
published data are not available on age assessment of any of the species
of Sphyraenidae. Cockerell (1914) used his brief study and description
of the scales of Sphyraena picuda ( = S. barracuda ) to draw conclusions
on the phylogenetic affinities of the Percesoces.
Cause of Annulus Formation in Barracuda.— The literature on the use of
scales and other hard parts for age determination of fishes is profuse. Most
references given by Graham (1929b) and Lagler (1956) deal with tem­
perate forms, where the theory is usually held that slackened body growth
due to decreasing water temperature is reflected in a corresponding de­
crease in the growth rate of the scale, as reflected in the relative spacing of
the circuli. In tropical and subtropical species, a marked slowing of growth
generally does not occur as a result of the smaller changes in water tem­
perature. Menon (1953) believed that temperature change was not
necessary for annulus formation in the scales of tropical fishes, but that the
periodic rhythm of spawning seemed to be sufficient per se to cause for­
mation of spawning marks on the scale. Chevey (1933) noted that a
temperature difference might cause an annulus in tropical species, although
the slowing of growth of the scale would be less pronounced than in
temperate species. He concluded that a temperature difference of at least
4 to 5°C seemed necessary to cause growth slowing and a resulting annual
check in the scale growth of tropical fishes. The barracuda is considered
a tropical species (Ekman, 1953: 46-47), and although there are seldom
sudden drops in water temperature during winter months in southern
Florida, the data of Bumpus (1957) show a mean annual temperature
change of 6.5 °C for the surface waters southeast of Miami, Florida. Thus,
if the assumptions of Chevey (1933) are applicable to the barracuda, the
annual temperature change in the waters of southern Florida would be
adequate to cause annulus formation in the scales of the barracuda.
M ethods of Scale Collection and Analysis.— Scales were collected from
barracuda caught off Miami and brought in by sport fishermen to Pier 5,
59
F ig u r e 15. Monthly increment from last annulus to anterolateral margin of
scale based on measurements of 316 specimens of Sphyraena barracuda of
age-classes I through VI from southern Florida. Circles represent mean values
for combined age-classes for each month. Dashed lines are mean rate of deposi­
tion of scale increment. Bracket A indicates probable resorption at scale margin
prior to annulus formation; bracket B represents probable period of annulus
formation.

Miami. Smaller fish which generally do not enter the anglers’ catch were
collected from the waters south of Miami, using hook and line, seine, and
rotenone. A large number of scales was collected and lengths recorded by
taxidermists Joseph T. Reese of Fort Lauderdale and the late A1 Pflueger
of Miami, Florida. A series of scales was collected from the left side of
each barracuda, midway between the lateral line and the base of the first
dorsal fin. This area has an unusually high number of regenerated scales,
but, fortunately, enough were collected so that perfectly formed scales were
usually obtained.
Because thickness and opacity preclude reading the scale itself, plastic
impressions were used (Gathman and Dawson, 1948). Butler and Smith
(1953) concluded that although there may be significant differences in
size between the original scale and its impression, errors in computing
back-calculated body lengths are within the range of errors obtained in
measuring the fish and its scales. Empirical ages of 555 fish were assigned
by scale reading irrespective of the fish length, sex, or date of capture.
All scale readings were made using a projection device (Phillips and Web­
ster, 1959). Agreement between first and second readings on a sample
of 100 scales was over 95 per cent. Using this method, annulus formation
at the posterolateral margin of the scale may be identified (Figs. 16 and
17).
The age analyses were pooled for 488 individuals collected over a
60
period of 13 months. Ages were recorded as the number of winters com­
pleted using Roman numerals, as is customary, to designate the number of
annuli present. The last day of winter was arbitrarily regarded as March 1.
Scales in which the annulus was omitted were considered as having the
annulus if adjacent annuli of other barracuda belonging to the same size
group had the annulus present. These scales were not used for growth
calculations.
Validity of the Annulus as a Year-m ark.— Lagler (1956) presented cri­
teria for the establishment of a valid annular mark. These assumptions
necessitate that only one mark is deposited per year and at about the same
same time every year.
To obtain the approximate time of annulus formation, the increment
from the last annulus to the anterolateral margin of the scale was summed
for fish belonging to age groups I through VI, and the mean obtained for
each month. The trends of the mean monthly increments were fitted by
eye in Figure 15. Annulus formation occurs sometime between January
and March; from December through January there appears to be a slight
resorption of the scale margin.
The time of annulus formation based upon scale increment is corrobor­
ated by an examination of the scales from 10 young barracuda collected
from November to June (Table 8). Young specimens collected in No­
vember and early December showed no annulus, but a check in the
growth of the scale appeared to have occurred recently in fish collected
on December 31 and on January 4. However, an individual collected on
January 7 showed no such mark. No young were available during February,
but two specimens collected on March 19 had well-defined annuli, with
numerous additional circuli laid down past annulus formation. Observa­
tions on samples of young and yearlings collected from March to Novem­
ber disclosed no annuli supplementary to those laid down between January
and March.
The relation between age and size of barracuda based upon empirical
scale analysis is given in Figure 19. Fish assigned to the same age group
in general have similar lengths, although overlap is great due to the slow
growth rate, differential growth, and the prolonged spawning period. Modes
in the length-frequericy distributions agree well with the calculated and
observed ages, although fish less than two years and older than 10 years
are difficult to detect in the distributions. For the younger age groups
there is good agreement among calculated growths of different age groups
of the same year’s collections. Calculations based upon samples taken in
August, 1955, and Aueust, 1956, agree well. In addition, young-of-the-year
do not show annuli (Table 8) before the length of the fish indicated by
the back-calculated growth (Table 11).
Criteria for the Interpretation of Annual M arks.— The criteria used for
annulus determination in fishes by Creaser (1926), Van Oosten (1921,
61
F ig u r e 16. Impressions of scales from specimens of Sphyraena barracuda
collected from southern Florida. A. Female, 756 mm fork length, showing four
annuli (x7.6); collected April 5, 1956. B. Male, 911 mm fork length, showing
eight annuli (x6.9); collected December 22, 1955.
62
F ig u r e 17. Impressions of scales from specimens of Sphyraena barracuda col­
lected from southern Florida. A. Sex undetermined, 1245 mm fork length,
showing nine annuli (x6.2); collected March 1, 1956. B. Sex undetermined,
1259 mm fork length, showing 14 annuli (x7.0); collected August 12, 1956.
63
1929), Beckman (1943), Kesteven (1950), and Lagler (1956) have
been used here.
False Annuli.— Sprugel (1954) summarized the possible causes of forma­
tion of false annuli. These were comparatively common in barracuda, and
of the 515 scales aged, 25.8 per cent had from one to four false annuli, and
nearly three-fourths had one false annulus (Table 9). False annuli seemed
to be most prevalent in the younger fish. Table 9 shows that 46.7 per
cent of the false annuli occurred in fish under two years, 75.7 per cent of
the fish were younger than four years, and no fish showed false annuli after
the eighth year of life. Because it was considered that the distribution of
the false annuli might merely reflect the abundance of the respective age
groups, the frequency of the observed ages in relation to the frequency of
false annuli was compared. There is a distinct preponderance of young
fish with false annuli, whereas the age-frequency distribution is composed
essentially of fish older than two years of age.
“Skipped” Annuli.— Sigler (1949) noted that some white bass, Roccus
chrysops, failed to form an annulus at the expected time. This was found
in barracuda also (Table 9). Slowed growth due to a lack of food does
not explain this phenomenon in the barracuda, which has an abundance of
food available at all times, and food intake throughout the year shows
little variation (Fig. 33).

TABLE 8
R esults of the e x a m in a t io n of sc a l es of 10 y o u n g -o f - t h e -y e a r
s p e c im e n s o f Sphyraena barracuda F l o r id a
from so u t h e r n to
ASCERTAIN PROBABLE PERIOD OF ANNULUS FORMATION.

Collection date Fork length (mm) Description of scale

November 15, 1955 161 No annulus formed
December 2, 1955 163 No annulus formed
December 15, 1954 153 No annulus formed
December 31, 1949 175 Annulus just being formed; crossing-
over along posterolateral margin of
scale
December 31, 1949 172 Annulus just being formed; one circu-
lus laid down beyond crossing-over
January 4, 1950 192 Annulus just being formed; crossing-
over along posterolateral margin of
scale
January 7, 1950 207 No annulus formed
March 19, 1952 164 Annulus formed, with four circuli laid
down beyond crossing-over
March 19, 1952 227 Well-developed annulus, with five cir­
culi laid down beyond crossing-over
June 13, 1955 241 Well-developed annulus, with eight cir­
culi laid down beyond crossing-over

64
 TABLE 9
A ge d is t r ib u t io n in r e l a t io n to the num ber of fa lse annuli and “s k ip p e d ”
a n n u l i o bser v ed in the scales of 519 s p e c im e n s Sphyraena barracuda
of
collected fr o m so u th er n F l o r id a fro m A u g u st, 1955, t h r o u g h A u g u s t , 1956.
No. Per cent
Age Number of Per cent of Number of Per cent of fish fish
class false annuli false annuli “skipped” annuli “skipped” annuli examined examined
O 22 13.0 16 34.0 8 1.5
I 57 33.7 10 21.3 19 3.7
II 27 16.0 9 19.2 38 7.3
III 22 13.0 7 14.9 53 10.2
IV 16 9.5 2 4.2 78 15.0
V 14 8.3 — — 67 12.9
VI 8 4.7 2 4.2 67 12.9
VII 3 1.8 1 2.2 66 12.7
VIII — — — — 47 9.1
IX — — — — 31 6.0
X — — — — 18 3.5
XI — — — 13 2.5
XII — — — — 8 1.5
XIII — — — — 4 0.8
XIV — — — — 2 0.4
Total 169 100.0 47 100.0 519 100.0

Growth of the Young .— Young barracuda were commonly observed in the

shallow waters of the Florida Keys from May until November, after which
time they move into somewhat deeper water where they become difficult
to collect. Seine collections were made as often as possible each month,
but collecting during the winter months yielded few small barracuda.
Despite their observed abundance, barracuda are difficult to collect with
seines because of their wariness.
From field and museum collections, 244 young barracuda were measur­
ed; these represent collections made from 1945 through 1956 (Fig. 18).
Variations in growth from year to year may be obscured by the groupings
of all data under a single month’s growth.
From April to October, young barracuda grow to an average length of
about 4 inches in standard length, with a range of about IV2 to 11 inches.
The variation in the observed growth of the young appears to be due
essentially to prolonged spawning throughout the season. For example,
the mean growth rate (Fig. 18) increases steadily from April through
July, then drops to a value only slightly higher than the mean value during
May. This appears due primarily to the occurrence of relatively large
numbers of very small, recently hatched fish.
The variations in length among the young in a given month are con­
siderable, and means are smoothed by a running average of threes, pre­
sented by a dashed line. The relatively abrupt slackening in the growth rate
65
 240

APR. MAY JUNE JULY AUG SER OCT NOV DEC. JAN. MAR.
(8) (II) (16) (21) (85) (5) (42) (35) (7) (5) (9)
18. Monthly length-frequency distributions of 244 young-of-the-year
F ig u r e
specimens of Sphyraena barracuda collected from southern Florida from 1945
through 1956. Dashed line represents mean growth rate obtained by moving
average of threes. Number in parentheses represents number in each month’s
samples. Horizontal bar represents mean; vertical line represents range of varia­
tion; longer rectangle represents one standard deviation on either side of the
mean; shorter rectangle represents two standard errors on either side of the
mean.

of the young fish noted by Walford (1932: 28-30) in Sphyraena argentea

was not evidenced in S. barracuda.
Those specimens of barracuda hatched later in the season grew at a
somewhat slower rate. Presumably, this reflects a decrease in abundance
of larval and postlarval fishes, on which young specimens of barracuda
feed, as the season progresses. But, generally, there is little change in the
seasonal growth rate of young specimens of barracuda on the basis of data
presented in Figure 18.
Empirical Growth Rate in Adults .— Adults as used in the arbitrary sense
here represent barracuda more than 12 months old. Empirical data are
based upon scale readings of barracuda collected at Pier 5, Miami, Florida,
from August, 1955, through August, 1956 (Fig. 19). Of 488 specimens,
151 were males and 222 were females; the sex of the remaining 115 was
not determined.
The empirical growth rate of males, females, and specimens whose sex
was not determined are shown in Figure 19 using the method of Hubbs and
66
Hubbs (1953). Data for fish older than nine years are few, but in those
less than nine years there appears to be no significant difference in the
growth rate between the sexes. The only departure in the growth rate
between the sexes is for class-III fish; however, a t-test indicates that the
difference is not significant for that class (t= 0 .2 7 , P > 0 .5 0 ).
The lengths of male and female barracuda assigned to age-class I are
not representative of the population. These fish are immature, and only
the larger individuals have developed sexually to a stage whereby the sex
can be identified by field methods. Thus, males (410 mm) and females
(400 mm) of age-class I appear to have a greater mean length than the
specimens whose sex is unknown (364 mm). When all specimens belong­
ing to age-class I are grouped, the mean fork length is 378 mm, which is
more representative of the population.
The barracuda is a relatively long-lived species; two individuals weigh-

1 . . I . . 1 .... 1. 1. __ I____ L _____ I .

1400.

1
1000-

800 -

60 0 .
I
f
400 .
MALES FEMALES COMBINED

1
200 -
0* T T~T I T
VI VII VIII IX XI XII
AGE C LA SS

F ig u r e 19. Empirical growth rate of Sphyraena barracuda from southern Flor­

ida, based on scale analysis of 488 specimens collected from August, 1955,
through August, 1956. Horizontal bar represents mean; vertical line represents
range of variation; longer rectangle represents one standard deviation on either
side of the mean; shorter rectangle represents two standard errors on either
side of the mean. In each age-class, stippled rectangles are males, obliquely
ruled rectangles are females, and solid rectangles combine males, females, plus
fish of undetermined sex.
67
 20. Relation between anterior scale radius in mm (x 26.7) and fork
F ig u r e
length of 295 specimens of Sphyraena barracuda collected from southern Flor­
ida from April, 1955, through August, 1956. Relation is expressed by the equa­
tion, y = 30.4 + 4.02 x.

68
 TABLE 10
L e n g t h o f s p e c i m e n s o f Sphyraena barracuda c o l l e c t e d f r o m so u t h e r n
F l o r id a a t t h e t i m e o f s c a l e f o r m a t i o n i n a r e a b e t w e e n base of
SPINOUS DORSAL FIN AND LATERAL LINE.

Fork length (mm) Description of area

28.0 No scales visible
30.0 No scales visible
30.1 No scales visible
30.2 Scale pockets present but no scales visible
30.4 No scales visible
31.0 Scale pockets present but no scales visible
31.8 Scales developed
32.9 Scales developed
35.0 Scales well developed
38.0 Scales well developed

ing 26 and 39 pounds had 14 annuli. Since the barracuda is known to reach
a weight of 106 pounds (Herre, 1936:100), it is probable that it lives
for much longer than 14 years.
Calculated Growth Rates .— Calculated growth rates more accurately depict
the length of the fish at each year than do empirical growth rates because
of increment in growth of the scale beyond the last annulus. Methods of
back-calculating growth from scale analysis have been studied by Graham
(1929a), Van Oosten (1929, 1953), Hile (1936, 1941), Cohen (1954),
Lagler (1956), and others.
Body-scale Relationship .— In calculating growth rates, it is assumed that
throughout life the scale grows at the same rate as the fish. Van Oosten
(1929) made use of scale magnifications without reverting to the actual
scale measurements in the back-calculation of body growth, and this
method has been followed in the present study.
It appears that the variance of the independent variable (body length)
is correlated with the dependent variable (scale length). Although this
variance could be reduced by the use of logarithmic transformations, Whit­
ney and Carlander (1956) considered it sufficiently accurate to disregard
the assumption that the variances of the body lengths are the same for
each scale length.
The body-scale relation for 295 barracuda collected in 1955 and 1956
from the waters adjacent to Miami, Florida, is shown in Figure 20. It was
assumed here that the relation is constant among the year classes within
the sample, and that there was no difference between the sexes. The
equation y = 30.4 -f 4.02 x was obtained for the general model
y = a -f bx, with a correlation coefficient (r) equal to +0.97; an r 2 value
of 0.94 suggests that linearity accounts for 94 per cent of the variance in
the fork length (Y ) due to scale length (X ). There is probably very
69
 TABLE 11
Su m m ary of m ean c a lculated fo rk lengths ba sed on scale m easurem ents of Sphyraena barracuda collected
fr o m so u th er n F l o r id a fro m A u g u st, 1955, through A u g u st, 1956.
Observed MALES
No. mean Calculated fork lengths (mm) at end of year of life3
Age of length at
class fish capture (mm) I II III IV V VI VII VIII IX X XI XII XIII
I 4 410 291
II 15 530 286 452
III 13 656 265 457 595
IV 28 715 275 448 578 675
V 17 786 260 421 552 655 734
VI 25 853 263 434 561 657 745 818
VII 18 905 267 444 569 674 758 831 892
VIII 9 940 242 419 534 631 734 811 869 918
IX 4 1040 261 438 567 706 811 891 942 1019 1050 — — — —
X 2 1060 271 394 494 608 665 729 786 832 891 953 — — —
XI 3 1106 284 442 564 662 748 834 896 956 1008 1053 1089 — —
XIII 2 1260 336 476 572 676 783 867 948 1017 1017 1113 1060 1197 1131
Total: 140
Calculated mean fork length: 269 440 566 663 747 825 890 945 1013 1042 1077 1197 +b
Growth increment: 269 171 126 97 84 78 65 55 68 29 35 + +
Calculated mean length, August: 404 525 629 712 789 864 913 973 1047 1057 1095 — +
Number of fish: 140 136 121 108 80 63 38 20 11 7 5 2 3
aOne male of age-class XIV was not included because scale was not taken from the same location on the body as others.
bData insufficient.
 TABLE 11— (CONTINUED)
Observed FEMALES
No. mean Calculated fork lengths (mm) at end of year of life
Age of length at
class fish capture (mm) I II III IV V VI VII VIII IX X XI XII XIII
I 6 400 270
II 13 510 260 418
III 23 607 263 427 561
IV 27 728 261 431 562 677
V 25 785 269 436 556 659 738
VI 27 848 277 432 556 659 749 824
VII 25 941 278 428 569 687 777 855 913
VIII 22 970 282 440 556 663 755 833 900 947
IX 15 1014 282 425 551 671 752 829 899 944 989 — — — —
X 9 1040 277 405 551 637 741 821 884 951 1000 1038 — — —
XI 9 1094 259 429 559 670 760 838 907 963 1017 1067 1096 — —
XII 4 1140 244 409 545 654 739 814 873 922 982 1034 1027 1128 —
XIII 1 1160 262 447 555 620 686 755 821 882 948 1002 1037 1098 1133
Total: 205
Calculated mean fork length: 271 429 558 667 753 833 900 947 996 1047 1072 1122 +b
Growth increment: 271 158 129 109 86 80 67 47 49 51 25 50 +
Calculated mean length, August: 409 508 623 722 796 873 934 971 1020 1073 1085 1147 +
Number of fish: 205 199 186 163 136 111 84 60 38 23 14 5 1
bData insufficient
little, if any, curvilinearity (Fig. 20), and because of the rather high
correlation coefficient and the relatively small variance there seemed to be
no need to test for linearity of regression.
The intercept value of 30.4 mm represents the fork length of the fish
when the scales were first deposited in the area of scale sampling. This
calculated observation is in good agreement with empirical observations
of young fish. Scales are first deposited on the caudal peduncle in young
barracuda at about 25 mm fork length, and are laid down progressively
anterad along the lateral-line region. Examination of 10 young barracuda
showed that scales were present between the first dorsal fin and the lateral
line, the location at which scales were taken from the adult fish, at
between 31.0 and 31.8 mm fork length (Table 10). Although no barra­
cuda less than 200 mm fork length was used in computing the body-scale
relation, empirical evidence on the approximate time of scale formation
would suggest that the relation remains approximately linear even at the
smallest stages.
Calculations .—Fork lengths at the end of each year to annulus deposition
(approximately February) were back-calculated using Fraser’s (1920)
equation:
ln = (L — C.F.) sn + C.F.,
S
where L = fork length of fish at capture,
C.F. = correction factor corresponding to fork length at which scales
were deposited in the area of sampling ( = 30.4 m m),
S = length of scale at capture (x 26.7 in this study),
sn = distance to any annulus from focus (x 26.7),
ln = fork length of fish when annulus was deposited.
The number of annuli on each scale was recorded, and measurements
were made from the focus to the anterolateral margin of the scale in mm
(x26.7) to obtain scale scale length (S), and from the focus to each
annulus (sn) along this same line. Since scales from the same area of the
barracuda show a certain amount of variability in size and shape, from
three to six scale impressions were measured on each slide and their
measurements averaged to reduce the variance about the regression line
of the body-scale relation (Fig. 20).
The same scales were used for back-calculated growth computations
as were used for empirical observations.
Calculations based on 140 males and 205 females from Miami, Florida,
are summarized in Table 11. Moderately good agreement among calculated
lengths at each annulus suggests again that these marks are yearly in
nature. Differences in calculated lengths at each age are remarkably similar
for the sexes, and are all within the expected range of error. The class-VII
females (N = 39) are calculated to have a length 10 mm greater than that
of the males (N = 38) and the class-IX males (N = 11) are 17 mm
72
 FORK LENGTH, MM
600 800 1000 1200 1400
' ------1 .....................................I.,...
I
SER 1
JM L

OCT
n W ln J ln ,
NOV.
—Tl—f^l—T1—f—1—fl—, n nn■ 1
ry i i- Irn -n prin n n n_____BE£_
JAM

-O__ CLn n f a .. FEB.

nnnn
.rn — H r - n n - , n APR. 1
MAY 1
JUNE
jfm Eh_
JULY
FLORIDA-----
BAHAMAS....

n b Mi'vHri

" " T " " ! ' " ........11 11111rji■i ■pi •■| ■i•| • 11• ■■11■111 TTTII j l l » l | »i

0 200 400 600 800 1000 I2(

1200 1400 1600
TO TAL LENGTH, MM
21. Length-frequency measurements of 1091 specimens of Sphyraena
F ig u r e
barracuda. Histograms enclosed in solid lines represent monthly samples col­
lected in southern Florida (N = 8 6 0 ) by seining (less than 300 mm long) and
from angler catches at Pier 5, Miami. Histograms enclosed in dashed lines in
July sample represent 231 barracuda collected from Bimini, Bahamas, in July,
1955, and July, 1956. Area between broken, vertical lines represents range
generally not sampled in Florida due to gear selectivity. Back-calculated mean
lengths based on scale analysis for August are indicated by arrows on length-
frequency distribution for August.
73
larger than the females (N = 29). The difference between the males and
females of age-class XII is considerably greater than in any other age
classes, but a discrepancy in the calculated growth rate of the males for
this class (N = 2) may be due to the small sample involved. The differ­
ences between the mean growth rate of the sexes of the class-VII group
was found to be non-significant (t = 0.04, P > .5 0 ); similarly, sexual
difference for class IX was not significant (t = 0.49, P > .5 0 ).
Barracuda do not exhibit contrasting regions of rapid and slow growth
from summer to winter, even though there is some evidence for this in the
length frequencies for O-class fish (see above under Growth of the Young).
Where data were adequate, slight evidence for Lee’s phenomenon, or the
apparent change calculated from different age groups (Van Oosten, 1929),
was evidenced in the back-calculated growths for the first five year classes
of male barracuda but this was not observed for the females.
Calculated Growth Increments.— The calculated growths of barracuda
from Florida are summarized in Table 11. The increments between suc­
cessive years are approximately the same for the sexes. The greatest in­
crement is during the first year, when the calculated growth for males and
females is 269 mm and 271 mm, respectively.
Length-jrequency Distributions.— The observed length frequencies of
young and adult barracuda collected from southern Florida are shown in
Figure 21. Young barracuda (i.e., those less than 12 months old) were
mostly collected by seine from 1945 through 1956; the adults were largely
from Pier-5 anglers’ catches.
In May, young are observed in the 80-mm class interval. By the follow­
ing March, the young have become too large to be collected by seining, and
their absence, indicated by vertical, broken lines in Figure 21, is construed
as being due to gear selectivity. Class-I fish are observed even in the
small frequency distribution for May, where they appear at the 280-mm
interval; age-class II is represented at the 460-mm class interval.
The sample taken in August, 1956, is the best estimate of the relative
proportion of each age group in the sample. A scale of values represent­
ing calculated mean fork lengths for each age, corrected to give lengths
for August, is superimposed above the length-frequency diagram for this
month. Agreement between the peaks in the length-frequency distribution
and the calculated lengths are remarkably similar (age-classes IV through
IX) where data are adequate.
Length frequencies of 231 barracuda collected in Bimini, Bahamas,
during July, 1955, and July, 1956, are superimposed on the distribution
of the Florida specimens for July as dashed lines in Figure 21. The rela­
tively larger size of the O-class fish from Bimini suggests either more
rapid growth rate or earlier spawning.
In the distribution for July (Fig. 21), a comoarison between the barra­
cuda taken in Bimini and Florida indicates that most of the individuals
74
 1953
VA-J---------- 1---------- 1---------- L ______ 1_______ 1___ _

JAN. L
J ______ n r. L

I n n n i— i ^ |

n n i---» P--- 1 r ' " V . r t

FISH

h v j j

i . j l F ' U „ “ " 1
OF

i „ . n

i J U N E I
NUMBER

n
^ i—i n rH rn- ' *—1 P----1 n n ’

i , „ J V i , . - r l

^ nnPs rif-^
A ______ OH__ CLD_r-1 n r -. r n ....n 1 « P

0 _a______ ___ o -o _r in ___ a___ lzj__ □__

0 1 n n n f l l — Hn i— i n n n NOV. F
m — i n M U 'n in n n D E C .1

L
r i 1 1 1 1 1
0 10 20 30 40 50 60
TOTAL LENGTH, INCHES
F ig u r e 22. Length-frequency measurements of 501 specimens of Sphyraena
barracuda from taxidermist’s records from southern Florida taken in 1953.
75
 1954
M ^ m iu fc U A iJU t 11111111111 h i

r—i fl n

JL Jk a ^ r
- " t
x 0
— 5 - 3

JD__£L
" " t
" 4

£ ol M __CL r-T T T t/K

tr
uj 0 1 n n___d3.nPrfil
CD
2 5
3 J l

AUG.
JL . / ■ V
n n fl ^ l - T J U i n , ^ ! SEP~ |
0
0
J ------------ cm---- n ib ----- 11 r—'In-n---- h
0 rArJl nP*l nn N0V' E
0
i/n n n HI r M “ S’ I
0 10 20 30 40 50 60
TOTAL LENGTH, INCHES
23. Length-frequency measurements of 682 specimens of Sphyraena
F ig u r e
barracuda from taxidermist’s records from southern Florida taken in 1954.
76
 1955

JA N.

0
__n nJVT-A-jinfi
5 FEB.
i
J ---------mD—LI____
JL n r> H
n ll nn l n» r "v_______
u " ^ ^1^1 a K

0 j __ --.0 „n ____ n JlRn, MAR^

1</> 5M
i
77 0-1 j=D____ aJL
°0 l -CL -0—n,. ^ J W M U ln n n " ^
ll_
o 1 pA p . i»j x ^ — r
DC
JULY
m—i a M t.I k i
LlI
m nr-in
AUG.

nrH.nnn
SEP.
, ] _______ rJI ■ -n n '-'^ V lU n
OCT.
1 fin.
n n nn a__an.
NOV
0 \ nn n__ n n nn j i Ol

0 J __________ o ___ o n ____ t b __ «~fJ1 r^ ~ h f___ n ..PEC

Mfi'i'i i ri yi n i| 11 n i m i | i m n i 8 i|ii ii i i ii i [i i i i i m i | i n i
0 10 20 30 40 50 60
TO TA L LEN G TH , INCHES
24. Length-frequency measurements of 672 specimens of Sphyraena
F ig u r e
barracuda from taxidermist’s records from southern Florida taken in 1955,
exclusive of Pier-5 samples.
77
 1956

i
fiiJ i 11111 >>11>>>a **ii I ah i JLi j l ii In i,u u u l ii ii ii 111 Ii i

5
0
jn-rx.
5
0
i n_ ■
n
5 p/UI MAR
0 .n...nn j=£L d±£2 £ a
APR.
5
n o nj y J W
xco t5 n n

u.
**■ 0 i JQ____ O__ D J 1 Q I im = L
MAY

0 o
JULY
c 5
m 0 £L nn J V N S n
5
AUG.
1 10
5
0 ax
SEP.
5
0 JD___ C 3 - H n -EL cQfloN o ,
0 nnn n n 0C T -

0 -EL
5 DEC.
0 ^TTir| 111111r ti p n n 111 J1........
0 10 20 30 40 50 60
TOTAL LE N G TH , INCHES
25. Length-frequency measurements of 852 specimens of Sphyraena
F ig u r e
barracuda from taxidermist’s records from southern Florida taken in 1956,
exclusive of Pier-5 samples.
78
 TOTAL LENGTH, INCHES
10 20 30 40 50 60
H h — '— •— i— i__ 1__ i__ 1-----1---- 1-----1-----1---- fc
A JAN.

>-] FEB. J W r l l 1]

j MAR.

24J APR.
12 J ,
0
1 _ rtn ~ r'u’a _ ____ L
X
CO 1 MAY .
0
Li. 1 JUNE
U_ 0
1 JULY
UJ
1 —On
CD

1 -
t II tltu s

12 H
I SEP.
! !
I I EESnZTCEEXXI

0
1 0CT„„ F
1 NOV.
0
12i DEC..
4//■■ r ... r - 9
0 200 400 600 800 1000 1200 1400
FORK LE N G TH , MM
26. Length-frequency measurements of 2707 specimens of Sphyraena
F ig u r e
barracuda from taxidermist’s records from southern Florida taken in 1953-
1956, exclusive of Pier-5 samples. Back-calculated mean lengths based on scale
analysis for August are indicated by arrows.
79
of age-class I and many of those from age-class II are absent from the
Florida length frequencies. What may be a dominant year class can be
followed in the distribution throughout the year by retracing the modal
growth from age-classes IX (1947) and XII (1944). Age-class V also
appears to be particularly abundant in the August samples, but it is not
possible to retrace its growth.
Total lengths, measured to the nearest inch, of 2707 barracuda caught
between Palm Beach and the Florida Keys (not including those landed at
Pier 5) were made available by the late Mr. A1 Pflueger. The data were
recorded by months for 1953 through 1956, and are shown in Figures
22 to 25. The composite data for these years, added by months, are given
in Figure 26.
A comparison was made between calculated lengths and length-fre­
quency modes for the month of August, based upon Pier-5 samples.
No calculated lengths or modal values are available for O-class fish, and
ages older than XI were not obtained by back-calculations. In most cases
the modal growth rate is slightly higher than either the observed or
calculated growth rates, and the calculated growth rate is higher than
that of the observed. These discrepancies may be due in part to the pres­
ence of weak year classes in the distributions. Nevertheless, it appears that
the growth rates using the two different methods are in relatively good
agreement.
As a supplementary method for testing the validity of check marks as
annual marks, Hile (1941) stressed the importance of tracing year classes
through several years on the assumption that weak or strong year classes
should be apparent over this period. In order to examine this hypothesis,
the monthly data have been summed to obtain the annual total number
landed (Fig. 27). Empirical lengths at each age are fitted to the graph
and each age class is traced back through succeeding years. If it can be
assumed that the mean rate of growth was approximately equal for taxi­
dermist’s samples from 1953 through 1956, and that this rate is the same
in the Pier-5 samples, the modes of each succeeding year class can be
traced over a four-year period when mean calculated lengths are assigned
to each year class. It would appear that in 1944, 1946, 1947, and 1948
comparatively strong year classes were produced, whereas in 1945 and
1949 poorly represented year classes apparently occurred. This is in
moderate agreement with the Pier-5 samples (Fig. 21).

M o r t a l it y a n d S u r v iv a l

Ricker (1948) reviewed the historical aspect of the development and

interpretation of “catch curves” derived from a knowledge of the age
distribution within a population sample. Rounsefell and Everhart (1953)
presented assumptions which must be satisfied prior to the estimation of
total mortality rates. Although all of the assumptions may not be fulfilled,
80
 TOTAL LENGTH, INCHES

CALCULATED AGE
27. Succession of year-classes in length-frequency measurements of
F ig u r e
2707 specimens of Sphyraena barracuda from taxidermist’s records from
southern Florida taken in 1953-1956, exclusive of Pier-5 samples. Oblique
lines connect successive year classes in which each age class was spawned.
Back-calculated mean lengths based on scale analysis are indicated by arrows.
81
it seems feasible to attempt to construct a rough catch curve for barracuda.
The catch curve of the Pier-5 barracuda is presented in terms of the
total age classes and the logarithmic transformation of their frequency in
the sport catch (Fig. 28). The ages are based here upon empirical age
analysis, rather than on back-calculated rates. As Ricker (1948) pointed
out, the ascending (left) leg of the curve and dome (central portion)
represent age classes which are taken in less frequent proportions than are
older fish, whereas the descending (right) leg of the curve represents
fish which are vulnerable to the fishery. If the descending leg is a straight
line, a uniform rate of survival is assumed. A concave descending leg
suggests a decrease in natural mortality with age, as well as a population
which may be overfished; a convex descending leg seems to indicate an

>-

AGE C LA SS
28. “Catch curve” based on age composition of 488 specimens of
F ig u r e
Sphyraena barracuda caught on hook and line from southern Florida from
August, 1955, through August, 1956.

increase in natural mortality with age, and, conversely, an underexploited

population. Keeping in mind that all of the assumptions for mortality esti­
mates cannot be satisfied, the catch curve (Fig. 28) suggests that, since
the descending leg is convex, the barracuda population suffers little fishing
mortality; rather, it would appear that the fishery is actually being under­
exploited. However, Ricker (1948) noted that it is difficult to distinguish
between a decrease in fishing pressure upon the population, and either an
increasing mortality with age, in which the more rapidly growing fish die
off sooner, or incomplete recruitment appearing in the older age groups.
Such a decrease in fishing pressure may be reflected in the age composition
of the barracuda due to the general inability of the angler to catch these
large fish.
Survival estimates were obtained using the method outlined by Jackson
(1939). Considering fish of only age-class IV and older because these
fish are theoretically incompletely vulnerable, the total survival of fish
between age-classes IV and XIII is 81 per cent, or a mortality of 19 per
82
 TABLE 12
C a l c u l a t io n o f m o r t a l i t y a n d s u r v i v a l b a s e d o n e m p i r i c a l a g e a n a l y s is
o f sc a l e s o f 488 s p e c i m e n s o f Sphyraena barracuda, c a u g h t o n h o o k a n d
l in e , fr o m 5, M i a m i , F l o r id a , f r o m A u g u s t , 1955, t h r o u g h A u g u s t ,
P ie r
1956. A n a l y s i s b a s e d o n m e t h o d o f J a c k s o n (1939).
Annual mortality Per cent Per cent
between ages Ratio survival mortality
MI 461/478 96 4
II-III 427/459 93 7
III-IV 376/425 88 12
IV-V 303/374 81 19
V-VI 238/301 79 21
VI-VII 170/236 72 28
VII-VIII 112/168 67 33
VIII-IX 73/110 66 34
IX-X 43/71 61 39
XXI 26/41 63 37
XI-XII 13/24 54 46
XII-XIII 6/11 55 45
XIII-XIV 2 /4 50 50

cent due to both natural and fishing mortality (Table 12). Such a survival
rate seems exceedingly high, and implies that the fishery is highly under­
exploited. Although it is conceivable that total survival for all age classes
approaches this value, the estimated survival between age-classes XII
and XIII of 55 per cent does not seem plausible. The samples for younger
fish are correspondingly higher. It appears that the most important factor
responsible for this apparent high survival is the neglect of the assumption
that recruitment is constant. An examination of the descending leg of
the catch curve (Fig. 28) indicates that the points representing each age
class are somewhat irregular; this suggests a replenishment of certain age
groups at intervals, which tends to give the appearance of a high survival.
If the stock annually was overexploited in the Miami area, the recruitment
from unfished areas would supply a fresh stock of all ages.
The catch curve indicates that fish are not exploited to any degree prior
to age-class IV, and that the highest rate of exploitation, or natural
mortality, occurs in fishes older than age-class VI. Since most males are
mature at age-class II, and females mature at age-classes III and IV,
theoretically these fish have had an opportunity to spawn at least once prior
to their entrance into the sport fishery. To maintain a basic spawning stock,
it would thus appear that there is no need for the management of barracuda
in the Miami area.
L a n d in g s of B arracuda

A few barracuda are taken commercially by handlines and by trolling

with live or artificial bait; some are caught in haul seines, trammel nets,
83
and runaround gill nets. The present-day commercial catch is taken
essentially by anglers who sell their fish to dealers. The reports do not
include the many barracuda taken for personal use.
The U.S. Fish and Wildlife Service (1952-1957) reported the annual
commercial catch of barracuda, all of which is taken from Florida waters.
Some are taken by angling in the Carolinas and along the northern Gulf
of Mexico but are not given in the statistical records. The commercial
catch for selected years from 1918 through 1956 is given in Table 13.
The barracuda is an important recreational species. Along the eastern
coast of Florida, Ellis (1957) found that in the number of charterboat
trips when fish were caught, barracuda were exceeded only by dolphin
( Coryphaena hippurus ), little tuna (Euthynnus alletteratus ) , and king
mackerel (Scomberomorus cavalla). Barracuda ranked seventh in numbers
of sport fish caught.
Since 1950, the reported commercial landings of barracuda have been

TABLE 13
C o m m e r c i a l l a n d i n g s o f Sphyraena barracuda f r o m F l o r id a f o r s e l e c t e d
YEARS, 1918-1956, AND TOTAL FOOD FISH, 1950-1956. DATA FROM
U.S. D e p a r t m e n t o f C o m m e r c e (1920-1938), A n d e r s o n a n d
P o w e r (1940-1954), a n d U.S. F is h a n d W i l d l i f e S e r v ic e
(1952-1957).
Average Average
Landings of price per Total price per
barracuda, Value, pound, food fish, Value, pound,
Year pounds, dollars cents pounds3 dollars cents
1918 7775 329 4.4 b — __
1923 1600c 64 4.0 — — —
1927 1000c 20 2.0 — — —
1929 16,000 560 3.5 — — —
1932 18,720d 749 4-0 — — —
1933 4245 180 4.2 — — —
1934 4245e 180 4.2 — — —
1939 500f 5 1.0 — — —
1940 200f 4 2.0 — — —
1945 45,800 2648 5.8 — — —
1949 43,100c 2236 5.2 — — • —
1950 95,400 4940 5.2 62,723,258 7,871,379 12.6
1951 75,900 3795 5.0 71,314,300 9,172,891 12.8
1952 66,893 4013 6.0 69,855,3658 9,831,153 14.1
1953 72,800 4048 5.6 65,452,736 8,183,093 12.5
1954 41,044 2873 7.0 64,580,484 8,154,828 12.6
1955 20,305 1015 5.1 66,635,464 7,979,452 12.0
1956 14,463 723 5.0 66,780,403 7,356,316 11.1

aExcludes alewives, scads, menhaden, sharks, ladyfish and freshwater catfish.

bNot included for these years. eMonroe County only.
cWest coast only. fPalm Beach County only.
dDade County only. gExcluding fresh- and saltwater catfish.
84
decreasing (Table 13; Fig. 29), although landings of food fish is increased.
The relative catch of barracuda taken commercially by counties (Table
14) remained roughly the same until the summer of 1954, when the catch
reported in Broward County sharply decreased. The landings reported in
Palm Beach were curtailed late that year, and in April, 1955, the landings
reported for Dade County abruptly ceased. These decreases in reported
landings can be traced to a series of outbreaks of ichthyosarcotoxism
caused by barracuda which occurred along the southeastern coast of
Florida from 1954 to 1956 (Paetro, 1956; de Sylva, 1956). Because a
Dade County law which forbids the sale of barracuda has been rigidly
enforced since this period, dealers have reported few or no landings. No
such decrease was observed in the catch reported from Pinellas County on
the western coast of Florida, where ichthyosarcotoxism evidently does not
occur, and where the sale of barracuda is not prohibited by law.
During semi-weekly visits to Pier 5, Miami (Table 15), the greatest
number was taken in August (16.7 individuals per visit), and the fewest
during February (3.4 per visit).

29. Commercial monthly landings of Sphyraena barracuda from Flor­

F ig u r e
ida waters, 1950 to 1957. Data are based on U.S. Fish and Wildlife Service
“Florida Landings.”

A comparison of the commercial and sport landings of barracuda is

made (Figs. 30-31). The large number landed in the commercial catch
during December, January, and February contrasts with the relatively few
caught for trophies for the same period. Although some barracuda are
reported in commercial catches in August, their numbers are proportionally
lower than in the Pier-5 data for this month. This may reflect the dispersal
of barracuda from the Florida Keys to the Gulf Stream, as well as a
slackening of fishing pressure in the Keys due to an abundance of
mosquitos. From September through November a decrease in commercial
landings is observed, followed by an increase in the catch with the return
of the angler to the Keys
85
 30. Relative monthly catch of Sphyraena barracuda by anglers in
F ig u r e
southern Florida in 1953-1956, as reflected in number of fish from taxidermist’s
records.

M ig r a t io n s

Records from commercial catches suggest that barracuda migrate

northward in the spring and southward in the fall (Table 14). Other
factors which may enter here include: (1) an onshore movement in the
86
spring, with barracuda in the southernmost regions moving in earliest;
(2) a coastwise change in fishing pressure as the weather improves; (3)
seasonal changes in the effectiveness of fishing as a result of changes in
the type of gear.
An inshore movement of barracuda with the onset of warm weather
seems unlikely, since in the southern part of Florida the reverse has been
observed. The remaining possibilities would depend upon the intensity of
and type of fishing carried out in the northern coastal areas. Angling is
widespread in Florida throughout the year, and it would be expected that
at least a few barracuda would be reported from the northern coastline
during the winter months if they were available.
Migrations of sphyraenids have been reported by other authors. Williams
(1956) believed that along the East African coast S. commersoni (= 5 .
barracuda) made local migrations corresponding to the spawning period
during the monsoon season. Meek (1916) stated that barracuda (species?)
migrate periodically to the coast (locality?) in the summer and remain
there until the end of the year. Migrations were noted by Villadolid (1940)
for S. picuda ( = 5 . barracuda) in the Philippines, and by Nikol’skii (1954:
401) for Sphyraena (sphyraena?). Walford (1933) stated that S. argentea
makes a spring migration, evidently for spawning, as far north as Ventura
County, California. On the basis of tagging studies, Springer and McErlean
(1961) reported that, generally, only large specimens of tagged S. barra­
cuda moved any great distance in Florida waters.

fei
si
<?
ssfe 40
So

31. Relative abundance of angler-caught specimens of Sphyraena bar­

F ig u r e
racuda from southern Florida, taken in 1953-1956 (broken line), commercial
landings of S. barracuda from Florida (solid line), and commercial landings
of total food fish from Florida (dashed line), taken in 1950-1956. Angler-
caught specimens of S. barracuda are taxidermist’s records. Commercial land­
ings are from U.S. Fish and Wrildlife Service “Florida Landings”; food fish as
listed here exclude alewives, cigarfish, menhaden, sharks, ladyfish, and fresh­
water catfish.
87
 TABLE 14
C o m m e r c ia l l a n d in g s o f b a r r a c u d a , in p o u n d s , r e p o r t e d f r o m F l o r id a c o u n t ie s ,

in U.S. F ish and W il d l if e S e r v ic e “ F l o r id a L a n d i n g s / ' 1952-1956.

West coast counties East coast counties
(north) (south) (north)

St. Lucie
Broward

St. John
Brevard
Monroe
Pinellas

Volusia
Walton

Martin
Collier

Beach
Dade

Palm
Lee

1952
Jan. — 200 — 484 3229 2185 80 1100
Feb. — — — 86 3896 3483 225 950 187 — — — —

Mar. — 200 — — 1037 2576 135 1150 950 — — — —

Apr. — 300 — — 363 1100 85 1600

May — 811 — 12 968 784 480 1987 — — — — —

June — 585 — — 298 1584 180 1385 490 — 400 — 150

July — 750 — 56 585 2420 300 1017 — 12 350 75 —

Aug. — — — — 1229 900 548 1283 — — — 119 —

Sept. — — — — 413 1542 420 530

Oct. — — 22 — 570 2361 255 600
Nov. — — — — 295 1873 270 300
Dec. — — — — 3652 5121 638 600
1953
Jan. — — .— — 1774 4076 180 1100
Feb. — — — 278 3229 3943 335 500 315 — — — —

Mar. — — 12 11 3203 2571 445 900 80 — — — —

Apr. — 545 — — 829 2700 685 400 335 — — — —

May 108 778 — — 1251 3212 150 497 195 589 — — —

June — — — — — 1400 140 794 — 40 — 37 —

July — 23 — 16 480 1596 290 2345 — 46 — 9 —

Aug. — 249 — — 827 1411 200 618 — 68 — — —

Sept. — — — — 815 1000 200 840 — 20 — — —

Oct. — — — — 306 580 12 1200

Nov. — — — — 173 1001 — — — 300 — — —

Dec. — 500 — — 492 1470 410 400 11,226 — — — —

1954
Jan. — — — — 4103 945 690 — — — — — —

Feb. — — — — 1836 609 310 72 29 23 — — —

Mar. — 521 — — 1549 765 565 10

Apr. — 2118 — — 510 414 769 1719 39 — — — —

May 674 652 — — 829 183 380 1589

June — 1196 — 31 598 714 445 2777 — 74 — 61 —

July — 244 — 92 703 1282 175 962 — 213 250 99 —

Aug. — 475 — 55 703 170 160 762 — 118 250 20 —

Sept. — 52 — 36 707 300 — 220

Oct. 22 — — — 616 457 — 100 — — — — —

Nov. — — — 15 1444 494 350 150

Dec. — — — — 1424 — 90 — — — — — —

(continued)
88
 TABLE 14 (continued)

West coast counties East coast counties

(north) (south) (north)

St. Lucie
Broward

Brevard

St. John
Monroe
Pinellas

Volusia
Walton

Martin
Collier

Beach
Dade

Palm
Lee

1955
Jan. — — — 837 950 — — — — — — —

Feb. — — — — 1456 1179

Mar. — — — — 1283 71 50 85
Apr. — — — — 453 — — — — — — — —

May — — — — 452 — — 58 — — — — —

June — 1923 — — 206 — — 576 — — — — —

July — 1693 — — 585 — — 356 — 20 — — —

Aug. 28 782 — 9 256 — — 193 — — — — —

Sept. — 1897 — — 298 — — 88 — — — — —

Oct. — — — — 397 — — 29 — — — — —

Nov. — 1075 — 15 1000 — 370 111 — — — — —

Dec. — 425 — 56 1029 — — 14 — — — — —

1956
Jan. — — — 18 17 — — . 82 — — — — —

Feb. — — — 38 156 — — 26 — — — — —

Mar. — 1300 — — 37 — — 12 — — — — —

Apr. — 1581
May — 1642 — 10 20 — — 20 — — — — —

June — 1733 — 60 103 — — 31 — — — — —

July — 569 — 46 140 — — — — 77 — 24 —

Aug. — 1133 — — 442 — — 90 — 84 — 45 —

Sept. — — — — 127 — — — — — — 39 —

Oct. — 146 — 18 2 —

Nov. — 363 — 114 54

Dec. — 186

Insight into the possible causes of the northward migration of the

barracuda may be gained by referring to Figure 32. Since barracuda are
caught in quantity from Palm Beach southward throughout the year
(Table 14), counties to the north which report only seasonal catches are
represented. The catch from 1952 through 1956 is plotted by month for
each county north of Palm Beach in which barracuda were reported by the
U.S. Fish and Wildlife Service in “Florida Landings” (1952-1957).
Arbitrarily using the midpoint of each county on the east coast where
barracuda were reported, I have represented on the left vertical axis its
distance from a southern baseline, the Monroe County line. The area
included between the dashed lines represents regions north of Palm Beach
which reported barracuda during the month indicated. Superimposed on
the graph are the mean monthly isotherms for the coastal waters along the
89
eastern coast, the values for which were taken from Fuglister (1947).
Barracuda are taken in Martin County in February, and in April and May
they move northward and appear in St. John County, a distance of about
330 miles north of Monroe County. Shortly thereafter they begin a gradual
drift southward, and by September they have moved as far south as St.
Lucie County. On the southward migration they are found in Martin
County until December, with some remaining near the southern boundary
of this county until January. Barracuda move northward (or offshore?)
out of the Palm Beach region in June and return in September. These
migrations probably include only a limited segment of the population.
This suggests that barracuda prefer temperatures higher than 74°F.,
as no fish were reported in the commercial catch north of this isotherm.
In May, the migration seems to follow the 779 isotherm to St. John
County where the barracuda appear in June. Before the temperature has
reached 80° in July, some of the barracuda are already moving southward.
By September they have moved as far south as St. Lucie County, where
the mean isotherm for this month is 80°. Since there is a movement of

TABLE 15
N um ber of s p e c im e n s of Sphyraena barracuda l a n d e d a t P ie r 5, M i a m i ,
F l o r id a , fro m A u g u st, 1955, t h r o u g h A u g u s t , 1956. ( C o l l e c t i o n s w e r e
n o t m ade fro m A p r il 19 t o M a y 11, a n d f r o m J u l y 2 t o J u l y 21, 1956.)
Date No. Date No. Date No. Date No.
visited landed visited landed visited landed visited landed
1955
Aug. 18 13 Nov. 28 8 Feb. 23 4 June 21 9
22 14 Dec. 1 4 Mar. 1 14 23 18
24 6 8 2 3 1 July 1 9
Sept. 7 7 16 1 5 2 22 5
10 15 20 6 17 1 25 6
13 1 22 10 20 7 29 20
14 2 26 2 21 9 Aug. 2 8
17 1 27 7 29 15 6 9
18 5 1956 31 7 9 32
21 5 Jan. 1 1 Apr. 5 8 12 19
26 19 3 0 11 1 16 1
Oct. 3 6 4 7 12 1 19 12
6 12 5 2 14 6 22 34
13 1 12 12 18 6 25 20
18 4 16 26 May 12 13 29 16
20 1 20 10 18 5 Total 649
25 10 22 3 22 4 No. trips 83
26 1 25 4 25 3 No. per
28 11 27 5 29 8 trip 7.8
Nov. 9 17 Feb. 7 1 31 1
16 11 13 5 June 9 9
23 1 16 4 13 10
27 8 21 3 18 2

90
 ME AN I S O T H ER MS (°F)
58 66 74
° ° ° 80 80
° ° 74
° 58
°

32. North-south drift of Sphyraena barracuda in relation to annual

F ig u r e
temperature changes in Florida east-coast waters, inferred from northernmost
locality in which specimens of S. barracuda were reported in commercial
landings by U.S. Fish and Wildlife Service “Florida Landings.” Mean monthly
isotherms taken from Fuglister (1947). Vertical axis at right shows principal
counties north of Palm Beach where S. barracuda is landed.

some of the fish out of the Palm Beach area in June to Martin County
when the temperature is above 82°, adult barracuda may avoid tempera­
tures much higher than this figure.
A similar but less well-marked migration is noted for barracuda along
the western coast of Florida (Table 14). During most of the year, they
are landed in Pinellas County (St. Petersburg area), but these landings
are generally not extensive prior to March or April, even though a number
of individuals seem to be resident there throughout the year. During May,
August, and September, barracuda have been reported as far north as
Walton County at the northern Florida “panhandle,” approximately 600
miles north of Monroe County. Large barracuda migrate into the northern
Gulf of Mexico during the summer months (Fig. 1), and are taken by
spearfishermen along the Mississippi coast; these fish evidently disappear
91
from the coastal waters with the advent of cold weather.
A coastwise rate of migration of about 2 miles per day was calculated
for barracuda from the data in Table 14, for fish on both the eastern and
western coasts of Florida.
E cology
Barracuda are abundant about the reefs, shallow flats, and mangrove
habitat of Florida. They are common along the southeastern, windward
side of the Florida Keys, but occur infrequently in Florida Bay and among
the Ten Thousand Islands and northward. The barracuda occurs both
inshore and offshore, although north of Miami it is more often found
offshore, presumably because it associates with the warm Gulf Stream which
subsequently swings away from coastal waters. In littoral waters, the
barracuda shows definite preferences at various ages and sizes in its
choice of habitat.
Pollution .— Although pollution of the coastal waters adjacent to Miami
is not yet extensive, sewage disposal, dredging operations, siltation of
reefs, and formation of spoil banks have changed some of the environ­
ment, particularly in Biscayne Bay, which was formerly more favorable
to barracuda and its food. Similar operations are altering the environment
in the same way in the Florida Keys. Such alterations or destruction of
the habitat are of great importance in limiting the range and abundance of
a species, probably far more than is the inefficient harvesting by man.
Living coral reefs, the associates of which adult barracuda so greatly de­
pend upon for food, are particularly susceptible to minor physical changes
(Kuenen, 1950: 414-419), and some reefs in the Miami-Fort Lauderdale
area which were formerly flourishing have been since affected adversely
by siltation and probably by pollution.
Salinity .— Adult barracuda tend to avoid waters of low salinity. The
salinity of the coastal waters adjacent to Miami, where adult barracuda
are still somewhat common, is typically oceanic. The mean salinity of these
waters is about 35%o, with a range of about 24 to 36.6%c (Dole, 1914;
Dole and Chambers, 1918). Fresh waters from the Miami River and the
Coral Gables Canal extend into lower Biscayne Bay, Miami (Smith, et al.y
1950). Juvenile barracuda freely enter the lower reaches of these waters
in which the salinity is comparatively low, but I have never heard of
adults being taken from them1. Kilby (1955) collected a 48-mm specimen
of S. barracuda from Bayport, Florida, in water having a salinity of \6.1%0.
Neither Regan (1908) nor Gunter (1942) reported barracuda from the
fresh waters of Central or North America, but Thompson [1944?] reported
that the barracuda (species?) . . ascends up into quite brackish
water . . .” in British Honduras. Bryan (1915: 349ff) and Hiatt (1947)
stated that barracuda is a serious predator in the mullet ponds of Hawaii,
lOn July 29, 1963, a 25-pound barracuda was caught in the Student Lake, University of
Miami by Mr. Ray Taylor. N o salinity value was obtained, but these waters are reportedly
fresh and are well above the area of salt intrusion.
92
which are presumably brackish. Jordan and Evermann (1926) reported
that in Hawaii barracuda actually enter “. . . mouths of streams, some
of them extending their range into fresh water.” Boulenger (1901: 363)
recorded S. guachancho from the fresh waters of the Congo River, and
Irvine (1947: 195-197) also listed this species from the fresh waters of
Africa. Reeve (1912: 273) stated that barracuda (species?) penetrate
the upper waters of the Gambia to a distance of 160 miles from the sea.
Depth .— Young barracuda are found usually in less than 10 feet of water,
but adults may swim in somewhat deeper strata. A school of sphyraenids
was observed by Quilici (1954: 67) at a depth of 120 feet. The species
cannot be identified with certainty even from his excellent photograph
but it appears to be S. obtusata. Mr. A1 Pflueger informed me that during
the Ocean Reef Fishing Contest off Key Largo, Florida, in August, 1955,
several barracuda of about 15 pounds were taken on live bait at a depth
of 240 feet. I have seen a barracuda near the bottom at a depth of 160
feet off West End, Bahamas, which weighed perhaps 25 pounds. Large
individuals are taken by charterboatmen by still-fishing with live baits in
the deep waters off Miami. But most barracuda are taken at or near the
surface, perhaps because most angling for oceanic species is by surface
trolling. During longlining explorations, U.S. Fish and Wildlife Service
personnel have taken barracuda at depths of over 200 feet in the
mid-Pacific.
Temperature .— The barracuda seems to be sensitive to sudden temperature
changes. Galloway (1941) observed dead barracuda to be “plentiful” at
Key West, Florida, during the cold winter from 1939 to 1940, when the
water dropped to a low of 43°F in January. Bumpus (1957) gave 71.6°F
as the mean January water temperature for Key West.
The apparent aversion of barracuda to water colder than 74°F in
Florida waters has been previously suggested as a possible factor controlling
their migrations. During periods of cold weather in Florida, Mr. Richard
Meyer of Miami informed me that large barracuda are found in dense
aggregations in the warm-water outflow of a power plant on Biscayne
Bay, Miami.
Bumpus (1957) gave mean maximum water temperatures (August)
for Fowey Rocks Lighthouse, southeast of Miami, as 83.8°F, and for
Dry Tortugas, Florida, as 85.1 °F. Although the barracuda in southern
Florida waters seem to tolerate these high water temperatures, those north
of Miami would appear to avoid them (Fig. 32). In Bimini harbor, small
barracuda were observed to become sluggish when the water temperature
exceeded 86°F. The shallow bank waters east of Bimini are subject to
wide variations in temperature (Newell and Imbrie, 1955), sometimes
dropping to the middle forties (°F ). Dr. Louis A. Krumholz informed me
that in January, 1956, a low of 57°F was recorded, and in July, 1956, the
temperature rose to 98°F in these lagoon waters of Bimini. Although
93
barracuda are sometimes common in such areas, they perhaps avoid
extremes of temperature by moving in and out with the tidal waters.
In July 1956, Dr. Krumholz and I observed barracuda about 12 inches
long swimming near the Lerner Marine Laboratory dock at Bimini in
water about 10 to 15 inches deep, the temperature of which was 88.7°F
at the surface. The barracuda were in the company of needlefish ( Strongy-
lura ), mojarra ( Gerres ), and mullet ( Mugil) but did not seem to be
feeding upon these fishes. These associates were seen to swim into waters
as shallow as 3 or 4 inches deep in which the surface temperature was
93.2°F, but the barracuda did not follow them into these warm waters;
they had previously been observed to do so when the water was 80.6 to
82.4°F. On August 29, 1955, I collected a number of 2- to 4-inch
barracuda from water 2 to 3 feet deep at the Matheson Hammock wading
beach, Biscayne Bay, Florida, where the temperature was 94-95°F. On
other occasions when small barracuda were collected at the same place,
water temperatures commonly exceeded 88°F. Although the young would
thus appear to tolerate higher temperatures than the adults, adult barracuda
seldom enter the very shallow waters regardless of water temperature.
E cology of Young Barracuda.— Temperature evidently plays a major role
in determining the seasonal movements of both the young and adult
barracuda. Aside from the migration pattern which was discussed above,
and which apparently is partially a function of temperature, seasonal
changes in water temperature seem to be responsible for movements
between shallow and deep water. These movements are correlated with
growth and are reflected in the appearance of certain items in the food
chain (Tables 16, 30). The young occur in abundance along the shallow
beaches as late as mid-September. Their disappearance from the shallows
indicates that young fish overwinter in deeper water where there is a
smaller temperature range and changes occur less rapidly.
During the warmer months, young barracuda were taken most often
just below the surf, and over sandy or weedy bottom. Here the detritus
of mangrove seeds and leaves offers protection, as well as an excellent
vantage point from which to forage. Young barracuda find food and shelter
in floating Sargassum spp. (Breder, 1949). Occasionally, I have taken
young barracuda associated with such a small quantity of Sargassum or
C ym odocea that it would seem to afford no concealment.
The young often find protection among the blades of turtle grass
(Thalassia) and its associated algal flora such as Laurencia, Caulerpa, and
Polysiphonia. Waite (1923: 109) noted a remarkable similarity in pattern
in young specimens of S. obtusata from Australia and the blades of
dugong (Cym odocea?) grass in which the young live, and stated that
. when danger threatens, they immediately upend themselves, their
snouts directed downwards, their tails moving in unison with the blades
of grass to which the stripes on the fish bear a remarkable resemblance.”
94
 Rosen (1911) observed young barracuda in tidepools in the Bahamas.
Such niches are important for the young both as a source of shelter and
for the rich gobiid fauna which serves as food for young barracuda.
Loose aggregations of young barracuda, especially during periods of
feeding, have been observed by me. Such groups usually comprise from
10 to 30 individuals, but occasionally large numbers of individuals have
been reported. Longley and Hildebrand (1 9 4 1 : 6 9 -71) noted that on
one occasion in the Tortugas . . about a thousand young up to 150 mm
in length were observed in one school.” Gary Bennett, of Cocoa, Florida,
reported in his “Trout Capital Fishing News,” dated August 6, 1956, that
at the west end of Canaveral Harbor and along the jetties there were
“thousands of very small baby barracudas . . .” less than 1Vi inches long.
Large numbers of young are not common on the western coast of
Florida. Reid (1954) and Kilby (1955) reported only a few young
barracuda from the Cedar Keys-Bayport area as a result of several years’
intensive collecting.

Ecology of Adult Barracuda .— Large adults are comparatively rare along

the eastern coast of Florida north of Cape Canaveral. They seldom venture
into shallow, sandy localities found here where environmental changes are
often severe. Barracuda more than 3 feet long are “lone wolves” of the
sea, are usually solitary both in reef and offshore areas, and constantly
appear to be searching for food (Sauvage, 1891; Gudger, 1918). In
contrast to younger stages, adults usually do not aggregate; on occasion,
groups of two or three large individuals have been observed to cruise slowly
about. During the winter, these larger barracuda are more prone to
aggregate.
In Bimini, Bahamas, barracuda to 4 or 5 feet long have been observed
about the oolitic islands, along the deep channels, and, in the summer, at
the junction of the Gulf Stream and the coastal waters or in the Gulf
Stream. During July, 1955. and July, 1956, large barracuda were observed
and taken on hook and line southwest of Bimini, where they occur over
extensive turtle-grass communities, usually at high tide. With low tide,
they were seldom caught in these shallow regions, but could be taken on
the leeward side of the islands, which is steep and coralliferous, and where
they were seldom taken on high tide. In the Florida Keys, such a preference
seems to reflect their avoidance of coral stacks, and barracuda were seldom
observed swimming about the jagged branches of elkhorn coral (Acropora
palmata) and staghorn coral (A. cervicornis), particularly when heavy
swells might dash such long-bodied species as barracuda about. Thus, the
smaller reef fishes may find ample protection among the branches of these
imposing coral barriers (see Longley and Hildebrand, 1941, PI. 2, Fig. 1).
Large barracuda were often observed hovering nearly motionless just
off the bottom, or near the surface (Longley and Hildebrand, 1941, PI. 2,
Fig. 2). They also occur commonly at the juncture between the turtle-grass
95
beds and the extensive sand patches fringing the reefs. Others may occupy
shallow trenches a short distance from the reefs. These points are excellent
places from which to forage for reef-dwelling prey.
It seems possible to offer some hypotheses on the seasonal movements
of barracuda in Florida waters using several independent criteria. Under­
water observations on the distribution of various age classes in different
habitats have been correlated with (1) catches in seine hauls, (2)
observations made while angling for barracuda, and (3) information
from charterboatmen and other anglers who are familiar with the seasonal
occurrence of saltwater fishes in southern Florida.
Postlarval barracuda initially occur in the shallow grass flats from late
spring to early fall. During their first winter, when they are about 6 to 8
inches long, barracuda appear to move offshore into grass flats which are
approximately 1 to 2 fathoms deep; at this size they are seldom if ever
found in reef areas. At the end of their first winter, at a length of 8 to 10
inches, they remain in the shallows. Some enter mangrove habitats or the
deep channels between the keys. By the beginning of their second winter
at an average length of 20 inches, they move offshore to the coral reef, in
depths of from 3 to 10 fathoms. By the beginning of their third year of
life, the majority are mature, and although most remain on the reefs, some
enter the shallow mangrove habitat by late spring. They overwinter on
the reefs, and in spring many migrate northward or are found in the
Gulf Stream. The adults remain scattered until September when they
gradually return to the reefs. These fall concentrations on the reefs are
coincident with the southward migration along the Florida coast. During
the winter, in the deep channels of the Florida Keys, charterboatmen have
observed large aggregations of barracuda which they claimed would
seldom take a bait.

A ssociates and Predators. — The adults of the four common sphyraenids in

the western North Atlantic do not occur in the same habitat. Adult speci­
mens of guachancho are often taken in 12 to 20 fathoms of water by
shrimp trawls in the Gulf of Mexico, according to Hildebrand (1954,
1955). This species is common enough at Veracruz, Mexico, to enter into
the commercial fishery (Dr. Henry H. Hildebrand, personal communica­
tion). Both borealis and picudilla occur in deeper strata in the western
Atlantic than does barracuda. Where barracuda is not common, the other
three species replace it in the food chain in shallow water.
The young of barracuda and borealis were taken together by me, and
on other occasions specimens of barracuda and guachancho occurred in
the same seine haul. Small specimens of guachancho are not often reported
and may be overlooked, although their pattern is sufficiently distinct to
insure identification (Fig. 4).
The important groups of fishes and invertebrates which occur in the
96
 TABLE 16
G e n e r a l r e l a t i o n s h i p b e t w e e n t h e s i z e o f Sphyraena barracuda, i t s
HABITAT, AND THE TYPICAL, IMPORTANT ORGANISMS ASSOCIATED WITH IT
in e a c h . C r o ss in d ic a t e s t h a t b a r r a c u d a a t t h is s iz e a r e a b s e n t o r
RARE IN THIS NIC H E. O RGANISM S PRECEDED BY ( 1 ) ARE PREDATORS W HICH
COM PETE W ITH BARRACUDA FOR FOOD. ORG A N ISM S PRECEDED BY ( 2 ) ARE
EATEN BY BARRACUDA. O RGANISM S PRECEDED BY ( 3 ) C O M PETE W ITH AND
ARE EATEN BY BARRACUDA. ORG A N ISM S PRECEDED BY ( 4 ) NEITHER
COM PETE W ITH NOR ARE EATEN BY BARRACUDA.

Total Sandy or
length, weedy Mangrove Reef Offshore,
inches shores habitat habitat pelagic
1-12 Gobiidae 1 Lutjanidae
Gerridae (yg.) 2 Cyprinidon-
Atherinidae tiformes
Engraulidae 2 Clupei-
Belonidae (yg.) formes + +
Syngnathiformes 2 Gerridae
Tetraodonti- Scaridae
formes (yg.) (yg)
4 Mugilidae

13-18 Gerridae Belonidae 2 Atherinidae

Atherinidae Lutjanidae 2 Clupeiformes
Engraulidae Cyprinodon- 2 Cephalopoda +
Scaridae (juv.) ti'formes 3 Synodontidae
Synodontidae Clupei- 3 Belonidae
Pomadasyidae formes
Gerridae
Mugilidae
Carangidae
19-30 2 Mugilidae 1 Seriola spp.
2 Lutjanidae 2 Pomadasyidae
2 Clupeidae 2 Tetraodontiformes
3 Synodontidae 2 Lutianidae
+ 3 Belonidae 2 Hemiramphidae
2 Cephalopoda
3 Synodontidae
3 Belonidae
3 Serranidae
3 Carangidae
3 Scombridae
4 Elasmobranchii

> 30 Scomberomorus 1 Scomberomorus

spp. spp.
Seriola spp. 1 Coryphaenidae
Lutjanidae 1 Istiophoridae
(large individuals) 2 Cephalopoda
Serranidae 3 Euthynnus
(large individuals) 3 Stronpylura
+ + Tetraodontiformes raphidoma
Hemiramphidae 4 Acanthocybium
Cephalopoda 4 Thunnus spp.
Euthynnus 4 Katsuwonus
Caranx spp. 4 Exocoetidae
Belonidae 4 Elasmobranchii
Elasmobranchii

97
various habitats occupied by barracuda of different sizes in southern
Florida, and the relationship between the barracuda and these organisms,
is summarized in Table 16. Large barracuda inhabiting reefs occupy the
same predatory position as the amberjack (Seriola dumerili), large jacks
( Caranx spp.), and various species of mackerels ( Scomberomorus spp.).
Needlefish ( Strongylura spp.) also compete for food, but to a lesser extent
than other predators because they are limited in the size of the prey they
can ingest. The barracuda feeds upon jacks and needlefish, and therefore
shares the role of major predator with the amberjack. Other carnivores
which occur in this habitat are groupers (Epinephelus, Mycteroperca,
Cephalopholis), large snappers (Lutjanidae), little tuna (Euthynnus), and
sharks; sharks apparently do not compete with barracuda for the same
type of food. The barracuda is capable of taking and killing large prey
with its immense gape and powerful, shearing jaws can ingest much larger
fish than can snappers, jacks, scombrids, and small groupers.
In the offshore, pelagic habitat, common predatory species include
sailfish (Istiophorus), dolphins (Coryphaena spp.), and tuna-like fishes
(Thunnus spp., Katsuwonus). These offshore predators compete only to a
limited extent with the barracuda (unpublished data). Within the sandy-
shore habitat, young barracuda have the least competition, for the small
needlefish are the only common predators.
Gudger (1918) discussed the presence of a remora (Echeneis naucrates)
on barracuda from Florida waters, and I have seen echeneids on barracuda
underwater but the species could not be identified. Poey (1865) described
Echeneis sphyraenarum from the gills of barracuda from Cuba, and
Schultz (1943) described a specimen of S. snodgrassi (= 5 . barracuda)
from Canton Island in the Phoenix Islands.
Many popular tales relate the animosity which supposedly exists between
barracuda and sharks. This is suggested by Wright’s (1948) observations
with a group of British frogmen in the Bahamas. He reported that a
barracuda attacked a nurse shark (Ginglymostoma cirratum) which had
been speared through a gill slit and was bleeding profusely. Ogilby (1915:
8, 47) noted that in Australia, S. obtusata has been known to drive off
large sharks. During the course of about 40 underwater trips to the Florida
Keys and the western Bahamas, I never saw sharks and barracuda together,
although they are both common in these waters.
Evidently, few predators are sufficiently large and swift to capture
large barracuda, although I was told by a spearfisherman that he once
found a small barracuda in the stomach of the spotted jewfish (Epine­
phelus itajara), reportedly a rather sluggish species. Klunzinger (1884)
reported that tunas feed upon sphyraenids, and other authors have noted
traces of small specimens of Sphyraena in the diet of several species of
Pacific tunas. I have taken two barracuda of 20-30 mm from the stomach
of dolphin (Coryphaena hippurus) from Pier 5, Miami.
98
 B eh a v io r
Typical coloration and positions when resting are described for young
barracuda by Breder (1948a, 1949), and by Longley and Hildebrand
(1941: 69-71). Schlesinger (1911) briefly discussed the fins and general
swimming behavior of S. sphyraena. The young of barracuda closely re­
semble twigs, and on several occasions have been nearly overlooked as
the seine hauls were being examined. Young and juveniles blend well with
their backgrounds in nature, and can change their pattern accordingly.
Over sand bottom they are ghostly pale, but over sand mixed with turtle
grass they have irregular, greenish bands; the transition may be noted as
a barracuda swims from one type of habitat to another. The black
margins of the caudal fin and opercular flap are conspicuous during both
of these phases.
Barracuda are swift swimmers. Gray (1957) reported that the speed
of S. barracuda was clocked in a laboratory tank at 27 miles per hour.
A slow drift is characteristic of individuals when not actively feeding.
Breder (1926) described their locomotion as “pike-like movements” with
great “flexibility in continuous swimming.”
Large barracuda were often observed to lurk just beneath the surface
in the waters of Bimini and the Florida Keys. Hiatt (1947) stated that
“barracuda habitually appear half asleep and float in one position close
to the surface, often making slow movements from side to side. When
they are disturbed they move with extreme rapidity.”
On several occasions I have noted large individuals swimming erratically,
with a list or with a distinct, convex flexure in the back. Since these fish
appeared only momentarily, it was not known if it represented a temporary
behavior pattern or was a deformity. Wright (1948) noted such a hump­
backed individual in the Bahamas, and spearfishermen have observed them
in the Florida Keys.
Aggregating Behavior.— In barracuda from 10 to 30 inches long, two
types of aggregating behavior were observed: feeding and defensive.
For example, a typical haunt of barracuda from 10 to 15 inches long is
a quiet, shallow area, about 2 to 6 feet deep, with sand bottom interspersed
with patches of turtle grass and detritus, such as roots, branches, and
mangrove leaves. Numerous barracuda could be observed in a given area;
each moved only within its own territory. Occasionally one would dart
out to ensnare a passing fish but returned to its territory. Aggregating
behavior to feed begins after the onset of flood tide, coincident with the
shoreward movement of schools of small atherinids, clupeoids, and gerrids.
These schools of forage fishes remain compact and seldom scatter when a
single barracuda rushes at them. Gradually the barracuda leave their
respective territories, congregate, and harass the schools of small fishes,
picking off stragglers. Such behavior has been described by Gudger (1918),
Mowbray (1922), and Gordon (1946).
99
 When a school of prey is swimming over sand bottom, seven or eight
barracuda have been observed to lie parallel to one another over a weed
bed, facing the prey. This apparent abandonment of territoriality was also
noted on numerous occasions when a lure was cast into a weed bed con­
taining barracuda. If the lure was retrieved rapidly, often one barracuda
would dart out to take the lure. However, if the lure was retrieved slowly
a number of barracuda would converge from the weed bed toward the
lure but made no attempt to strike at it. When the lure was slowed or
stopped the barracuda would slow or stop; in the latter instance they usu­
ally returned to the weed beds.
The second type, designated as defensive behavior, was observed in
barracuda from 18 to 30 inches long. This reaction seems to be due to the
presence of an enemy. At Cat Cay, Bahamas, on two occasions barracuda
formed a semicircle about me as I entered the shallow water, but remained
about 10 to 15 feet distant. As I approached, they gradually retreated,
and the reverse was true. On each occasion, when I attempted to spear
one it would dart away with great speed but immediately returned to its
approximate place in the formation. Several barracuda swam at the
unretrieved spear; they merely bumped it with the snout, but made no
attempt to bite or slash.
Ward (1907: 313) reported that schools of sphyraenids (species?) off
Australia . . rival those of herring and mackerel for numbers and
denseness.” Walford (1932, 1933) and Nikol’skii (1954: 401) briefly
mentioned schooling behavior in Sphyraena. Schooling or aggregating
behavior seems to be confined to the small species of Sphyraena, or to
the juvenile forms of the large species.
Barracuda less than 2 feet long are sometimes found in well-marked
size aggregations, even within the same age class. Spawning barracuda
may congregate in schools of the same sex, based on my observations that
individuals caught in a restricted locality are often of the same sex.
On two occasions at Bimini three large individuals were observed to
cruise slowly at the surface, drifting offshore with the tide. In the second
instance, the three were swimming within inches of one another. The
largest barracuda was swimming slightly ahead of and between the others,
and was taken on hook and line; it was a 26-pound female with nearly ripe
eggs. On the other occasion none was caught. Gudger (1918) described this
accompanying behavior in three barracuda observed at Tortugas, Florida,
except that the three individuals were about the same size. He noted that
the two individuals on either side had “. . . their heads in the region of the
right and left pectoral fins . . .” of the middle barracuda. This may
represent pre-spawning activity, since the barracuda from Bimini had
nearly ripe ovaries.
A type of accompanying behavior is observed when a barracuda is
hooked; other individuals may be attracted and appear frenzied as they
100
swim beside the tethered companion, and occasionally will slice off the
tail of the hooked fish.
Peck Order .— Barracuda evidently are not very compatible, and it would
seem that where aggregations of individuals occur, some type of social
stabilization would develop. Evidently, a hierarchy exists in the juveniles,
as suggested in the following observations made at Bimini at the Lemer
Marine Laboratory.
On July 11, 1956, three barracuda which had been caught on hook and
line were placed in a concrete pool, measuring approximately 4 feet wide
by 2 feet deep by 6 feet long. The fish included a 450-mm male, a 398-mm
female, a 397-mm male, and a 306-mm individual (fork lengths) whose
sex could not be determined subsequently by dissection. The first three
barracuda were placed in the tank within a few minutes of one another.
Each went to and remained in a separate comer of the tank, breathed
rapidly, and attempted to maintain balance. Each lay near the bottom
and close to the side of the tank. Within a few seconds after the third
barracuda was placed in the tank all had recovered. The three individuals
raced at one another, but no contact was made, and after a few seconds
again retired to separate corners of the tank. The two fish which measured
397 mm and 398 mm lashed at and sideswiped one another; their snouts
glanced off at an angle just below the base of the first dorsal fin. This
“jousting” was repeated four times, and then one struck a deep, glancing
blow which opened a gash about 1Vi inches long and about V4 inch deep
on the side of the other fish. Again they hit each other with a sideswipe,
although this time there was no gash inflicted, and they turned once more
and attacked. They slowed momentarily, clamped the tips of their jaws
together, rocked their heads sideways for several seconds, and slowly
released their holds. This was repeated once more, and then they returned
to their respective comers.
At this time the largest fish left its corner and faced the adversary
which had inflicted the gash. Both “jousting” and jaw-clamping were re­
peated. The larger fish went to the corner previously occupied by the
smaller barracuda. The entire episode occurred in approximately two
minutes. A fourth barracuda (306 mm) was placed in the tank with the
others, and it entered the unoccupied corner. The three original individuals
congregated in one corner while the smallest barracuda occupied the di­
agonal corner. No attempt was made by any to fight with this newcomer,
nor was any fighting observed up to the time when the observations ceased
about 15 minutes later.
During the present investigations, several barracuda were caught which
bore scars below the spinous dorsal base which might have been obtained
in such a battle. This may also account in part for the high percentage of
regenerated scales which I found in this region of the body.
Mowbray (1922) wrote that in the Florida Keys, two adult male barra­
101
cuda which had been placed in the live well of a boat inflicted severe
gashes on one another soon after they were confined. Mr. Craig Phillips,
formerly of the Miami Seaquarium, informed me that shortly after intro­
duction into a tank 80 feet in diameter, two large barracuda fought, split
each other’s head open, and both died.
It is not known at what size this fighting commences. Barracuda less
than 1 inch long will devour or attempt to eat one another when placed in
a 40-gallon aquarium tank. Cannibalism was noted by De Kay (1842:39-
40) in young specimens of S. borealis.

F e e d in g H a b it s

In aquaria, barracuda up to 300 mm long have been observed to feed

more readily when in the company of congeners than when only one
specimen is present. Both young and adult barracuda have have been
observed to swallow prey either head or tail first, or to grasp the prey at
the midsection, fold it, and swallow it. On other occasions, the prey may be
seized at the midsection and sliced in two or more pieces, and the tail
section swallowed; this is more common in larger individuals. The litera­
ture on the functional morphology of feeding in this family is limited to a
reference by Yasuda (1960) on S. pinguis.
Barracuda from about 100 to 300 mm were fed portions of frozen
shrimp ( Penaeus ). Usually they allowed the shrimp to fall to the bottom,
where it was scrutinized for a few seconds before being held loosely in the
jaws momentarily prior to swallowing. Those from which food was with­
held were quite ravenous and took the shrimp as soon as it touched the
surface of the water.
Barracuda of all sizes feed upon moving organisms, and the stomach
contents suggest that they depend primarily upon sight to obtain their
food. Their predilection is for fish which flash or show some type of
abnormal, erratic movements.
Barracuda are generally daytime feeders, and probably do not move
about much at night. At Bimini, small cyprinodonts and gerrids introduced
at night into a tank containing several barracuda were still there several
hours later, whereas under daytime conditions they were readily consumed.
Barracuda caught early in the morning usually had empty stomachs or
recently swallowed food. In July, at Bimini, barracuda were caught on
hook and line from about 8 AM until about 5 PM. Toward dusk they
showed little interest in lures or fishes used as bait.
During 1961 and 1962, several periods were spent in the western
Bahamas and the Florida Keys using waterproof night lights over reefs
and in deeper water. On one occasion, a barracuda of about 3 feet long
approached to within 10 yards of the light (150 watts) but was apparently
dazed by the light and slowly drifted away. On numerous night-diving trips
to the Florida Keys, Walter A. Starck, II, and William P. Davis, of the
102
Institute of Marine Science, University of Miami (personal communica­
tion), observed a barracuda on only one occasion; this too appeared dazed
by their lights as they swam underwater.
Relatively little nocturnal trolling or drift fishing is carried on in southern
Florida, and thus deductions based upon the lack of specimens cannot be
considered conclusive. But Springer and Bullis (1956) recorded barracuda
taken on longline gear in the northern Gulf of Mexico between the hours
of 10 PM and 1 AM. Williams (1959) reported that all of their troll
catches of barracuda off East Africa were made during daylight hours,
although other sphyraenids were predominantly night-time feeders. La-
Monte (1952) stated that members of the family Sphyraenidae often feed
at night, and Doyle (1953) referred to barracuda (species?) caught by
night angling off East Africa. Palombi and Santarelli (1953:24) stated
that in the Mediterranean Sea, the commotion made by small fishes which
have been attracted to the surface lights used by fishermen attracts the
attention of Sphyraena spet (—S. sphyraena). Ommanney (1953) and
Williams (1956) noted that large numbers of S. jello were taken at night,
primarily during periods of full moon, off Amirante Bank, Seychelles, by
handline, but specimens were never taken by trolling.

F ood H a b it s

Except for the studies of Hiatt (1947) and Williams (1956), little has
been found in the literature regarding the food of the barracuda. Barracuda
stomachs were collected most of the year in southern Florida from the
sport fishery. A dependence on such a fishery leaves much to be desired,
since if the fish does strike a lure or bait it implies that the fish must have
been hungry or incited. Thus examination of the stomach may cause an
underestimation of the quantity of food taken under normal circumstances.
M ethods .— Stomachs of 901 barracuda were examined from various lo­
calities in the tropical western Atlantic Ocean. Nearly all adult fish exam­
ined (longer than 300 mm) were caught on hook and line with live or
artificial bait, and a few juveniles were taken in seines.
All young fish captured by me were taken either with a 10-foot “Com­
mon Sense” V^-inch mesh cotton minnow seine, or with a 30-foot nylon
seine of V^-inch mesh.
At Bimini, most stomachs of adults were examined in the field within
a few minutes of capture. A few adults which were caught on charter
boats at Bimini and all of the adults examined from Pier 5, Miami, had
been caught from one-half to eight hours prior to the time of dissection.
There was no evidence from present observations that barracuda re­
gurgitate food in their stomachs when they have been caught on hook and
line, such as occurs in certain scombrid and other fishes.
Food is here considered as any type of solid matter including fish bones,
flesh, scales, squid beaks and “pens,” algae, and vascular plants. Un-
103
 TABLE 17
N u m b e r o f i t e m s f o u n d i n e a c h o f 5 2 9 s t o m a c h s o f Sphyraena barracuda
FROM THE TROPICAL WESTERN ATLANTIC. YOUNG WERE COLLECTED FROM
1 9 2 0 t h r o u g h 1 9 5 6 ; a d u l t s w e r e c o l l e c t e d f r o m A p r il , 1 9 5 5 ,
through A u g u st, 1956.

Adults
Young mm
(3 0 0 -1 2 1 0
( 5 .5 -2 9 9 mm fork length) fork length) Total
No. items
per
stomach Frequency Per cent Frequency Per cent Frequency Per cent
1 77 6 7 .0 132 8 0 .0 209 7 4 .7
2 23 2 0 .0 27 1 6 .4 50 1 7 .8
3 6 5 .2 4 2 .4 10 3 .5
4 5 4 .5 1 0 .6 6 2 .1
5 2 1 .7 — — 2 0 .7
6 1 0 .8 — — 1 0 .4
11 — — 1 0 .6 1 0 .4
16 1 0 .8 — — 1 0 .4

Total 115 1 0 0 .0 165 1 0 0 .0 280 1 0 0 .0

recognizable material was recorded as “unidentifiable matter.” If only

chyme was present, the stomach was considered empty. Food is reported
as the number of stomachs containing a particular item. Volumetric data
were not taken.
Number of Items of Food per Stomach.— On the average, each stomach
contained only one item (Table 17), and this usually filled the stomach
area. The tendency for a stomach to have more than one item was only
somewhat more pronounced in the young barracuda; 33 per cent contained
more than one item, and in the adults 25.3 per cent did so. One 850-mm
barracuda, caught near Bimini, held chopped remnants of worm eels,
Ahlia egmontis, including 11 heads. A barracuda 56 mm long, from Bis-
cayne Bay, Miami, contained 16 postlarval atherinids which ranged from
11 to 20 mm in total length. In most instances when young barracuda
had ingested more than one item of food, the organisms were the post­
larvae of schooling fishes such as atherinids, engraulids, or clupeids. Bean
(1890), Gudger (1918), Longley and Hildebrand (1941 : 69-71), Gor­
don (1946), and Legand (1952) noted that the barracuda feeds upon
schools of small fishes, but did not give the size of the barracuda nor the
prey.
The percentage of stomachs containing food in young barracuda is given
in Table 18.
Food of Young from Florida .— The organisms found in the stomachs of
154 young barracuda from southern Florida are given in Tables 19 and
20. Fishes occurred in 95.5 per cent of the stomachs which contained food.
104
 TABLE 18
P ercentage o f s t o m a c h s c o n t a i n i n g f o o d i n 195 y o u n g s p e c i m e n s ( l e s s t h a n
300 m m ) o f Sphyraena barracuda c o l l e c t e d f r o m s o u t h e r n F l o r id a , 1945 t h r o u g h
1956. N u m b e r o f in d i v i d u a l s r e p r e s e n t e d i n p a r e n t h e s e s .

No. %
Fork exam- with
length (mm) Jan. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec. ined food

0 -2 4 100(1) — 100 (1) 0 (2) 100 (1) 100 (3) 5 9 .1 (2 2 ) — — 100 (1) — 31 6 4 .6

2 5 -4 9 — — 83.3 (6) 100 (3) 83.3 (6) 8 9.9 (9) 60 (31) — 85 .8 (7) 100 (3) — 68 7 9 .5

5 0 -7 4 — — 100 (1) 100 (2) 100 (10) 50 (2) 7 1 .4 (7) 100(3) 6 6 .7 (3) 75 (3) 100 (1) 32 87.5

7 5 -9 9 — — — — 100 (1) 100 (8) 100 (2) 100(1) 100 (2) 100 (3) 0 (1) 18 9 4 .5

1 0 0 -1 2 4 50(2) — — —■ 50 (2) 100 (3) 100 (3) — — 50 (2) — 12 8 1 .8

1 2 5 -1 4 9 — — — — — 100 (2) — — — 100 (3) — 5 100

1 5 0 -1 7 4 — 100(1) 100 (1) — — 100 (2) —■ — — 100 (3) 50 (2) 9 8 8 .9

1 7 5 -1 9 9 100(1) — 100 (2) — — — 100 (1) — — 100 (3) 50 (2) 9 8 8 .9

2 0 0 -2 2 4 — 100(1) 100 (1) — 33.3 (3) — — — — 0 (1) — 6 50

2 2 5 -2 4 9 0 (1) — 1 0

2 5 0 -2 7 4 0 (1) 1 0

2 7 5 -2 9 9 6 6 .7 (3) — 3 6 6 .7

Total 75(4) 100(2) 9 1 .7 (1 2 ) 7 1 .4 (7 ) 8 2 .6 (2 3 ) 9 3 .1 (2 9 ) 6 9 .6 (6 9 ) 100(4) 8 3 .3 (1 2 ) 8 1 .5 (2 6 ) 4 2 .9 (7 ) 195 7 9 .0

No. items 5 3 11 6 19 27 54 4 10 22 4 165 —

No. stomachs __
with food 3 2 11 5 19 27 48 4 10 22 3 154
 Food preferences are not marked, and the young barracuda eat
whatever is available in the appropriate size class. However, certain
groups of fishes are taken frequently (Table 20). Gobioids, clupeoids, and
atherinids were by far the most important items of food, and occurred
in 69.1 per cent of the stomachs. Gobioid fishes excepted, the striking
feature in the diet of the young barracuda is the predominance of small,
schooling fishes of the littoral regions. Gobioid fishes, however, occur in
tidepools and on sand flats in greatest abundance, where they find shelter
in bivalve shells and echinoid tests.
Invertebrates as a group were a minor food item of the young, and
occurred in only 3.9 per cent of the stomachs.
Food of Young from Bimini, Bahamas.— Collections of young barracuda
were made in July, 1955, and July, 1956, mostly along the eastern shore
of North Bimini, in front of the Lerner Marine Laboratory. The difference
between the two years in relative percentage containing food is not
significant (chi-square = 2.64) at the 5-per cent level (Table 21).
Fishes were found in 97 per cent of the stomachs of the young which
contained identifiable food; of these, 30 (46% ) were identifiable.
The kinds of food taken by young barracuda from Bimini in 1955 and
1956 are shown in Tables 22 and 23. Gobiids, atherinids, gerrids, cyprino-
dontids, and clupeoids were found in 26 (86.7% ) of the 30 stomachs
which contained identifiable fishes, with atherinids the most common.

TABLE 19
Sum m ary o f fo o d f o u n d in t h e st o m a c h s o f 154 y o u n g s p e c i m e n s ( l e s s
than 300 m m ) of Sphyraena barracuda c o l l e c t e d f r o m s o u t h e r n F l o r id a ,
1945 through 1956. N u m b e r s a r e s t o m a c h s i n w h i c h i t e m s w e r e f o u n d .
Dec.-
Group Mar. Apr. May June July Aug. Sept. Oct. Nov. Total
Gobioidei 1 — — 1 10 5 ____ ____ ____ 17
Atherinidae 1 1 1 4 1 6 — — 2 16
Clupeiformes 1 1 1 1 — 9 — — 1 14
Engraulidae — — — 1 — 1 — 1 3 6
Gerridae 2 — — 1 — 2 — — — 5
Cyprinodontiformes 2 1 3
Hemiramphidae — — — — — 3 — — — 3
Scaridae 1 1
Gobiesocidae 1 1
Sparidae 1 1
Coryphaenidae — — — — — — 1 — — 1
Unidentified fishes 4 6 3 8 10 28 2 9 11 81
Invertebrates — 2 — 1 2 1 — — — 6
Plant material 1 1
Unidentified matter — — 1 — 3 — 1 — 5 10
Total 12 11 6 19 27 55 4 10 22 166

106
 TABLE 20
F ood it e m s fo und in the st o m a c h s of 154 young sp e c im e n s ( less than 300 m m ) of Sphyraena barracuda
collected in so u th er n F l o r id a , 1945 through 1956. N um bers are st o m a c h s in w h ic h it e m s w ere fo und .

Pisces:
Clupeidae — — — — — 6 — — — 6
Redear sardine, Harengula humeralis — 1 — — — — — — — 1
Harengula sp. 1 — 1 — — — — —1 3
Dwarf herring, Jenkinsia lamprotaenia — — — 1 — 2 — — — 3
Engraulidae — — — 1 — 1 — — — 2
Striped anchovy, Anchoa hepsetus — — — — — — —1 3 4
Cyprinodontidae
Rainwater killfish, Lucania parva 1 — — — — — — — — 1
Poeciliidae
Mosquitofish, Gambusia affinis 1 — — — — — — — — 1
Sailfin molly, Mollienesia latipinna — 1 — — — — — — — 1
Hemiramphidae — — — — — 3 — — — 3
Gerridae 1 — — 1 — 1 — — — 3
Silver jenny, Eucinostomus gula 1 — — — — 1 — — — 2
Coryphaenidae — — — — — — 1 — — 1
Sparidae
Pinfish, Lagodon rhomboides — — — 1 — — — — — 1
Scaridae 1 — — — — — — — — 1
Callionymidae
Lancer dragonet, Callionymus bairdi — — — — — 1 — — — 1
Blenniidae — — — — — 1 — — — 1
Gobiidae 1 — —1 5 3 — — — 10
o
00

TABLE 20— (CONTINUED)

Item Dec.-Mar. Apr. May June July Aug. Sept. Oct. Nov. Total
Clown goby, Microgobius gulosus — — — — 3 — — — — 3
Highfin goby, Gobionellus oceanicus — — — — 1 — — — — 1
Bathygobius sp. — — — — 1 — — — — 1
Gobiesocidae — — — 1 — — — — — 1
Atherinidae 1 1 1 2 1 5 — — — 11
Hardhead silverside, Atherinomorus stipes — — — 2 — 1 — — 2 5
Unidentifiable fishes 4 6 3 8 1 0 28 2 9 1 1 81
Arthropoda-Crustacea
Grass shrimp, Palaemonetes vulgaris — — — 1 — — — — — 1
Copepoda — — — — 1 1 — — — 2
Mollusca-Cephalopoda — 2 — — — — — — — 2
Chaetognatha — — — — 1 — — — — 1
Plant material — — — 1 — — — — — 1
Unidentifiable matter — — 1 — 3 — 1 — 5 10
Total 12 11 6 20 26 54 4 10 22 165
 At Bimini, the food of the smallest barracuda consisted mainly of
gobiids, whereas those over 3 inches long fed primarily upon schooling
atherinids and clupeoids. Perhaps this indicates a transitional stage between
niches and/or types of food available. Where surf is absent, the quiet,
shallow-water zone inhabited by very small barracuda varies from about
1 to 12 inches deep, and is often very rich in detritus which offers
excellent camouflage for young barracuda. This niche is often rich in
gobiids, cyprinodontids, and small gerrids, but the abundance of schooling
fishes is not realized until somewhat deeper water is encountered, where
barracuda of slightly larger size are abundant, but are less gregarious than
in shallower waters.
A partly digested barracuda which measured about 90 mm had been
swallowed tail first by a 291-mm barracuda; this was the only instance of
cannibalism in the examination of 901 stomachs.
Food of Young from Additional Areas .— The stomach contents of 39
barracuda from museum collections were examined by me from various
localities in the tropical western Atlantic (Table 24). The food of six
postlarval barracuda ranging from 5.5 to 12.1 mm in standard length,
collected by the DANA vessels from the Virgin Islands, consisted of bits
of fish.

TABLE 21
P e r c e n t a g e o f s t o m a c h s c o n t a i n i n g f o o d i n 89 y o u n g s p e c i m e n s ( l e s s
t h a n 3 0 0 m m ) o f Sphyraena barracuda c o l l e c t e d f r o m B im in i, B a h a m a s ,
i n J u l y , 1955, a n d J u l y , 1956. N u m b e r o f i n d i v i d u a l s r e p r e s e n t e d in
pa r en th eses.

Fork Total Total Total

length, no. with no. per cent
mm July, 1955 July, 1956 food examined with food
0-24 __ 100 (1) 1 1 100
25-49 100 (1) 100 (2) 3 3 100
50-74 80 (5) 100 (14) 18 19 94.7
75-99 42.8(14) 67.5(19) 19 33 57.6
100-124 84.7(13) 84.6(13) 22 26 84.6
125-149 100 (2) 100 (3) 5 5 100
150-174 — — — — —
175-199 — — — — —
200-224 100 (2) — 2 2 100
225-249 — — — — —
250-274 — — — — —
275-299 — — — — —
Total 70.3(37) 84.6(52) 70 89 78.7
No. items 29 49 78
No. stomachs
with food: 26 44 70 — —
Mean no. items
per stomach : 1.1 1.1 1.1 — —

109
F i g u r e 3 3 . Percentage of stomachs of 4 4 6 specimens of Sphyraena barracuda
from Pier 5 , Miami, Florida, which contained food, taken from August, 1 9 5 5 ,
through August, 1 9 5 6 .

Food of Adults from Florida .— From August, 1955, through August, 1956,
I examined 446 stomachs of adult barracuda. Of these fish, 185 (41.6% )
contained food (Table 25).
Most fishes taken are associated with the coral-reef habitat (Tables
26 and 27). Fishes accounted for 93.3 per cent of identifiable food. Of
these 71.1 per cent were identifiable at least to genus.
Where samples were adequate for comparison, stomachs taken from
October through March contained only slightly less food than those taken
in the remaining months (Fig. 33). The smallest young and the largest
adults more often bad food in their stomachs (Fig. 34).
Four groups comprised nearly 64 per cent of the identifiable fishes
taken by adults. The Beloniformes, mainly hemiramphids, were the most

TABLE 22
S u m m a r y o f fo o d it e m s f o u n d in st o m a c h s o f 7 0 y o u n g s p e c im e n s ( l e s s
t h a n 3 0 0 m m ) o f Sphyraena barracuda c o l l e c t e d f r o m B i m i n i , B a h a m a s ,
in Ju l y , 1 9 5 5 , a n d Ju l y , 1 9 5 6 . N u m b e r s a r e s t o m a c h s in w h ic h it e m s
WERE FOUND.

Group 1955 1956 Total

Gobioidei 2 6 8
Atherinidae 2 4 6
Gerridae 4 1 5
Cyprinodontiformes — 4 4
Clupeiformes 2 1 3
Scaridae 1 1 2
Sphyraenidae — 1 1
Opisthognathidae 1 — 1
Unidentified fishes 8 26 34
Invertebrates ____ 1 1
Plant material 1 — 1
Unidentified matter 8 4 12
Total 29 49 78

110
important single group, and accounted for 24 (22% ) of the identifiable
fishes (Table 26). Fishes of the order Tetraodontiformes (Plectognathi)
ranked second in importance as a group, and occurred in 23 stomachs.
These were followed by carangoid and mugilid fishes.
It was not always possible to distinguish between fish taken as baits
and those taken naturally, although in the former instance hook marks
were usually diagnostic. Species of the families Hemiramphidae and
Mugilidae are popular baits (for sailfish), and those of the families Carangi-
dae and Scombridae are usually still-fished or drifted live over the reefs.
Of the reef-dwelling tetraodontiform fishes, members of the family
Balistidae were most important, and occurred in 13 stomachs; diodontids

TABLE 23
It em s o f fo o d f o u n d in t h e st o m a c h s o f 70 y o u n g s p e c i m e n s ( l e s s t h a n
300 m m ) of Sphyraena barracuda c o l l e c t e d f r o m B i m i n i , B a h a m a s , i n
Ju l y , 1955, a n d J u l y , 1956. N u m b e r s a r e s t o m a c h s i n w h i c h
it e m s w e r e f o u n d .

Item 1955 1956 Total

Pisces:
Elopidae
Ladyfish, Elops saurus — 1 1
Clupeidae
Harengula sp. 1 — 1
Dwarf herring, Jenkinsia lamprotaenia 1 — 1
Cyprinodontidae
Cyprinodon baconi — 2 2
Poeciliidae
Gambusia (manni?) — 2 2
Gerridae 1 1 2
Silver jenny, Eucinostomus gula 1 — 1
Cuban mojarra, Gerres havana 2 — 2
Scaridae — 1 1
Striped parrotfish, Scarus croicensis 1 — 1
Gobiidae 1 2 3
Bathygobius spp. — 3 3
Clown goby, Microgobius gulosus 1 — 1
Darter goby, Gobionellus boleosoma — 1 1
Opisthognathidae
Mottled jawfish, Opisthognathus maxillosus 1 — 1
Sphyraenidae
Great barracuda, Sphyraena barracuda — 1 1
Atherinidae 2 4 6
Unidentifiable fishes 8 26 34
Arthropoda
Isopoda — 1 1
Plant Material
Rhodophyceae 1 — 1
Unidentifiable matter 8 4 12
Total 29 49 78

I ll
were the second most common. To my knowledge, the tetraodontiform
fishes are never used as bait.
The size of the food ingested often bore no relation to the size of the
barracuda. A barracuda which measured 657 mm had eaten a lizardfish
which was 37 mm long. Remoras of less than 4 inches were taken from
barracuda which were over 3 feet long. Needlefish which were almost
as long as the barracuda were usually found folded over several times
in the stomach of the predator.
Food of Adults from Bimini, Bahamas.— Of the stomachs of 132 adults
which were examined from Bimini, 78 (59.1% ) contained food (Table
28). Barracuda in 1955 contained food more often (75.8% ) than did
those captured in 1956 (53.5% ). Barracuda taken in 1955 were generally
smaller and were taken in shallow-water mangrove habitats; those taken
in 1956 were larger and were caught in somewhat deeper water and
about oolite reefs.
Fishes accounted for 84.2 per cent of the identifiable items found in
64 stomachs of adult barracuda taken from Bimini in 1955 and 1956
(Table 29). The tetraodontiform fishes were the most important group,

TABLE 24
F ood f o u n d in st o m a c h s o f 29 y o u n g s p e c im e n s ( l e s s t h a n 300 m m ) o f
Sphyraena barracuda collected from C u b a , P u e r t o R i c o , B e r m u d a , V i r g in
Isla n d s, and B im in i and E l e u t h e r a , B a h a m a s , 1920 t h r o u g h 1956.

No. of stomachs
with item
Clupeidae
Harengula sp. 1
Belonidae
Strongylura sp. 1
Cyprinodontidae
Striped killifish, Fundulus majalis 1
Poeciliidae
Gambusia sp.
Gerridae 1
Sparidae 1
Gobiidae 2
Blenniidae 1
Atherinidae
Reef silverside, Allanetta harringtonensis 1
Hardhead silverside, Atherinomorus stipes 1
Mugilidae 1
Unidentifiable fishes 14
Arthropoda
Insecta, Blattidae 1
Plant material
Angiosperm flower 1
Total 29

112
 1— '— I— '— I— •— I— '— I— I—
0- 200- 400- 600- 800" 1000- 1200
99 299 499 699 8 99 1099 1299
FORK L E N G T H ,M M .
34. Percentage of stomachs of 641 specimens of Sphyraena barracuda
F ig u r e
from southern Florida, arranged by size group, which contained food. Fish
less than 300 mm long were collected from 1945 through 1956; fish longer
than 300 mm were taken from August, 1955, through August, 1956.

with the puffer occurring most frequently. Carangids, principally the bar
jack and bigeye scad, ranked second; parrotfishes were third.
Comparisons and Discussion.— The literature on the food habits of
sphyraenids contains mostly random observations, so that it is difficult
to draw conclusions about the relative importance of certain items as food.
Bean (1890) noted that barracuda from Yucatan fed upon schools of
Harengula and Stolephorus. Legand (1952) observed that in New Caledo­
nia, S. picuda (— S. barracuda) and S. jello fed upon schools of Atherina
forskali. He added that both species fed upon mullet ( Mugil sp.) which
frequented the mangrove habitat. Mowbray (1922) noted that barracuda
fed upon schools of mullet, when “barracuda are seen in the greatest
numbers.” Breder (1927) found a 10-mm barracuda in the stomach of
a larger barracuda, and three in the stomach of another individual taken
from Cuba. He also recorded Synodus and a myrid eel. Beebe (1928)
observed a barracuda feeding upon a fairy basslet ( Gramma sp.) in
Bermuda waters, but such instances of a barracuda feeding upon corallico-
lous fishes are rare. Ommanney (1953) noted that five of 12 stomachs of
S. commersoni ( = S . barracuda ) from the Seychelles Islands contained
specimens of Lethrinus ramak, and another contained an eel. Williams
(1956) working in the same region and upon the same species, collected
mostly fishes, including a belonid, a lutjanid, a scombrid, and a clupeid;
stomatopods ( Squilla sp.) and squid were also found.
Longley and Hildebrand (1941: 69-71) observed that on occasion
squid were eaten by barracuda in the Tortugas, Florida, region. Malpas
(1926) found small fishes, as well as a cephalopod (Sepia), in the stomach
of Sphyraena acutipinnis, and two specimens of Balistes mitis in specimens
of S. obtusata from Ceylon. Chacko (1949) also found Sepia in the
113
 TABLE 25
P ercentage o f sto m a c h s c o n t a in in g fo o d in 446 adult s p e c im e n s of Sphyraena
barracuda collected from P ie r 5, M i a m i , F l o r id a , fr o m A u g u st, 1955, through
A ug u st, 1956. N um ber of in d i v id u a l s r e p r e s e n t e d in parenth eses.

Fork
length,
mm Aug. Sept. Oct. Nov. Dec. Jan. Feb. Mar.
300-399 __ __ __ __ __ __ __ __
400-499 100 (3) 0 (1) 100. (2) 33.3 (3) — — 0 (1) —
500-599 50 (2) 20 (5) 0 (4) 40 (5) 10 (2) 50 (6) 100 (1) —
600-699 — 100 (1) 33.3 (6) 61.6(13) 12.5 (8) 42.9 (7) 100 (1) 100 (1)
700-799 40 (5) 33.3 (9) 36.4(11) 50 (6) 50 (6) 33.3 (3) 0 (1) —
800-899 100 (2) 36.4(11) 66.7 (3) 50 (4) 25 (4) 37.5 (8) 33.3 (3) 44.5 (9)
900-999 — 50 (2) 0 (3) 50 (2) 50 (4) 100 (2) 50 (2) 40 (5)
1000-1099 100 (2) 100 (2) — 50 (2) 0 (1) 50 (4) 66.7 (3) 42.9 (7)
1100-1199 — 100 (2) — 0 (1) 100 (1) — 0 (2) 33.3 (3)
1200-1299 100 — — — — — — —
Total 71.4(14) 42.4(33) 34.5(29) 50 (36) 32.5(26) 46.7(30) 42.9(14) 44 (25)
No. of items 10 15 10 23 14 14 6 13
No. stomachs
with food 10 14 10 18 8 14 6 11

Fork
length, No. % with
mm Apr. May June July Aug. examined food
300-399 — — — (1) 100 — 1 (100)
400-499 — (2) —
50 (1) 0 12.5 (16) 29 31.1
500-599 — — 33.3 (3) (3) 0 30 (10) 41 29.3
600-699 — 50 (2) 0 (1) (8) 25 31.3 (16) 64 39.1
700-799 — 75 (4) 42.8 (7) (8) 25 41.2 (34) 94 40.4
800-899 50 (4) 75 (4) 50 (8) (9) 22.2 45 (20) 89 43.8
900-999 100 (3) 33.3 (3) 72.8(11) (1) 0 40 (25) 63 49.2
1000-1099 66.7 (3) 25 (4) 50 (4) (4) 50 54.5 (11) 47 53.2
1100-1199 0 (3) 0 (2) 50 (2) — 0 (1) 17 29.4
1200-1299 100 (1) — — — — 1 (100)
Total 57.2(14) 47.6(21) 52.7(36) 38.9(35) 36.9(133) 446 41.6
No. of items 12 10 25 12 57 221 —
No. stomachs
with food 8 10 19 9 49 186 —

stomachs of 51 specimens of S. obtusata, which ranged in length from

17 to 45 mm, from Indian waters; clupeids, atherinids, mugilids, and
leptocephalus larvae were also eaten. Inger (1955) collected a gobiid
and a percoid (Ambassis) from a 400-mm specimen of S. jello from
Borneo. Ah Kow (1950) found anchovies ( Stolephorus sp.) to be a main
food item in the diet of Sphyraena jello from the Singapore Straits during
January, October, and December; other food taken included squids and the
larvae of decapod crustaceans. Roughley (1916) stated that in Australia
S. novaehollandiae . . plays havoc with . . . garfish [Hemiramphus ],
yellowtail [Trachurus], and mullet [M ugill” Gudger (1918) found 5
114
 TABLE 26
Sum m ary of food it e m s in st o m a c h s of 186 adult s p e c im e n s of Sphyraena barracuda collected fro m P ie r 5,
M ia m i, F l o r id a , fro m A u g u st , 1955, through A ug u st, 1956. N u m b e r s are st o m a c h s in w h ic h it e m s w e r e f o u n d .

Group Aug. Sept. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Total

Beloniformes — — 2 4 1 2 1 4 3 1 1 2 3 24

Tetraodontiformes — 3 4 — 2 2 — 2 — — — 3 7 23

Carangoidei — 2 1 — — — 2 1 2 1 1 — 6 16

Mugilidae — 1 1 1 3 — —■ 1 — — 1 4 — 12

Clupeiformes 1 2 4 — 1 8
Scombridae 2 2 — 3 7
Pomadasyidae 1 1 — — — — — 2 — — 1 — — 5
Anguilliformes — — 2 — 1 — — — — — — — 1 4

Scaridae — — — — — 1 — — 1 — 1 — 1 4

Other fishes — — — 1 1 3 1 — 2 1 3 — 4 16

Unidentified fishes 2 2 — 8 3 7 2 3 3 1 7 1 9 48

Other vertebrates 1 1

Invertebrates — 1 — 1 1 — — — — — — — 4 7
Plant material — 1 1 1 1 4

Unidentified matter 7 5 — 3 2 — — — 1 1 4 3 17 43

Total 10 15 11 18 14 15 6 13 13 10 26 14 57 222
 TABLE 27
F o o d i t e m s f o u n d i n s t o m a c h s o f 186 a d u l t s p e c i m e n s o f Sphyraena barracuda collected at P ie r 5, M ia m i,
F l o r id a , f r o m A u g u s t , 1955, t h r o u g h A u g u s t , 1956. N u m b e r s a r e s t o m a c h s i n w h ic h it e m s w ere fo und.

Item Aug. Sept. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Total

Clupeidae — — — — — — — — — — — — 1 1
Spanish sardine,
Sardinella anchovia — — — — — — — — 1 — 1 — — 2
Atlantic round herring,
Etrumeus sadina — — — — — — — — —- 2 — — — 2
Harengula sp. — — — — — — — — — — 3 — — 3
Synodontidae
Inshore lizardfish,
Synodus foetens — — — — — — — — — — — — 1 1
Muraenidae
Gymnothorax sp. — — — — 1 — — — — — — — — 1
Ophichthyidae — — 1 — — — — — — — — — — 1
Sharptail eel,
Myrichthys acuminatus — — 1 — — — — — — — — — 1 2
Belonidae
Houndfish,
Strongylura raphidoma — — — — — — — 1 1 — — — — 2
Strongylura sp. — — — — — — — — 1 — — — 2 3
Flat needlefish, Ablennes hians — — — 1 — — — — — — — — — 1
Hemiramphidae — — — — 1 2 — 2 — 1 — — 1 7
Ballyhoo,
Hemiramphus brasiliensis — — 2 3 — — 1 1 1 — 1 1 — 10
Exocoetidae — — — — — — — — — — — 1 — 1
Serranidae — — — — — — — — — — 1 — — 1
Epinephelus sp. — — — — — — — — 1 — — — — 1
Sand perch,
Diplectrum formosum — — — — — — — — — — 1 — — 1
 TABLE 27— (CONTINUED)

Item Aug. Sept.Oct. Nov. Dec. Jan. Feb. Mar. Apr. May June JulyAug. Total

Lutjanidae — — — — — — — — — — — — 1 1
Carangidae — — 1 — — — — 1 1 — — — 4 7
Rough scad, Trachurus lathami — 1 — — — — — — — — — — — 1
Blue runner, Caranx crysos — 1 — — — — 1 — 1 1 1 — 2 7
Pomadasyidae 1 — — — — — — 2 — — 1 — — 4
French grunt,
Haemulon flavolineatum — 1 — — — — — — — — — — — 1
Sciaenidae — — — — — 1 — — — — — — — 1
Chaetodontidae — — — — 1 1 — — — — — — — 2
Scaridae — — — — — 1 — — 1 — — — — 2
Rainbow parrotfish,
Scarus guacamaia — — — — — — — — — — 1 — 1 2
Scombridae — — — — — — — — — — — — 1
Chub mackerel, Scomber colias — — — — — — — — — — 1 — — 1
Little tuna,
Euthynnus alletteratus — — — — — — — — — 2 1 — 2 5
Frigate mackerel, Auxis thazard — — — — — — — — — — — — 1 1
Ophidiidae
Short-bearded cusk eel,
Lepophidium brevibarbe — — — — — — 1 — — — — — — 1
Nomeidae
Man-of-war fish,
Nomeus gronowi — — — — — — 1 — — — — — — 1
Mugilidae
Mullet, Mugil spp. — 1 1 1 3 — — 1 — — 1 4 — 12
117

Bothidae — — — 1 — — — — — — — — — 1
 TABLE 27— (CONTINUED)
00
Item Aug. Sept. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Total

Echeneidae
Marlinsucker,
Remora osteochir
Balistidae
Orange filefish,
Alutera schoepfi
Gray triggerfish,
Batistes capriscus
Ocean triggerfish,
Canthidermis sufflamen
Tetraodontidae
Diodontidae
Striped burrfish,
Chilomycterus schoepfi
Porcupinefish, Diodon hystrix
Unidentifiable fishes
Reptilia-Chelonia
Mollusca-Cephalopoda
Echinodermata-Ophiuroidea
Basket star,
Astrophyton muricatum 1 1
Plant material
Turtle grass, 1 1
Thalassia testudinum
Sargassum weed, Sargassum spp. 1 1 1 3
Unidentifiable matter 7 5 — 3 2 — — — 1 1 4 3 17 43
Total 10 15 11 18 14 14 6 13 12 10 25 14 57 219
pounds of fish chopped in large fragments in the stomach of a 55-inch
barracuda from Tortugas, including half of a margate (Haemulon album).
Beebe and Tee-Van (1928) noted a 55-mm parrotfish (Scarus croicensis)
in the stomach of a 200-mm barracuda from Haiti, an 80-mm specimen
of Chloroscombrus chrysurus in a 400-mm barracuda, and a 70-mm
specimen of Ocyurus chrysurus in a 697-mm barracuda.
In the western North Atlantic, Sphyraena borealis, a small species, has
been found to contain young fish and young gastropods (Summer, et al.,
1913), and shrimp (Linton, 1905). Atherinids and mysids are the
principal food of juveniles of borealis taken from the Delaware River
estuary and adjacent areas (de Sylva, et al., 1962).
Of the 901 barracuda of all sizes examined by me during the present
investigation from various localities in the tropical western Atlantic, 529
(58.7% ) contained food. Fishes were found in 82.2 per cent of these
stomachs, plant material in 2.8 per cent, invertebrates in 2.6 per cent,
and unidentifiable matter in 12.2 per cent. Gobiid and atherinid fishes
ranked first and second as food of the young in both the Bimini and the
Florida region. Gerrids ranked third in Bimini, whereas they were sixth in
importance in Florida. Cyprinodonts.were fourth in Bimini, but seventh
in Florida waters. Invertebrates, which ranked fifth in Florida, were tenth
in importance in Bimini. Plant material occurred fewer times in Florida
than in Bimini.
Beloniformes were more important in the diet of adult barracuda from
Florida than any other major group, but only ranked fourth in Bimini.
The Tetraodontiformes ranked first in Bimini, and second in Florida as
food. Carangids ranked second and third from Bimini and Florida,
respectively, but scarids were of considerably greater importance in
Bimini than in Florida. Mugilids, which were so commonly found in
Florida barracuda, occurred only once in 38 stomachs at Bimini, and
although mullet are comparatively scarce in Bimini, their abundance in
barracuda from Florida seems to reflect their wide use as bait. This
seems to be true also of the Beloniformes and the Carangidae, which
were so commonly found in the Florida barracuda. Small scombrids and
pomadasyids, which figured prominently in the diet of the Florida fish, did
not occur in those from Bimini.
The differences in the type of food taken by adults in Florida and
Bimini would seem to reflect a difference in habitats, rather than food
preferences of the barracuda. The waters of southeastern Florida support
luxuriant growths of massive and branching corals whereas Bimini is
surrounded by extensive sandy stretches which are less conducive as
substrate to the growth of coral reefs (Parr, 1933; Kuenen, 1950; Newell
and Imbrie, 1955).
A definite change occurs in the type of food eaten with a change in
size (Table 30). Atherinids, clupeoids, gerrids, and cyprinodontids occurred
in 76.2 per cent of the stomachs of the young. These same groups accounted
119
 TABLE 28
P ercentage of sto m a c h s c o n t a in in g fo o d in 132 adult s p e c im e n s of
Sphyraena barracuda collected fro m B im in i, B ah a m as, in Ju l y , 1955, and
Ju l y , 1956. N u m b e r o f in d iv id u a l s r e p r e s e n t e d in p a r e n t h e s e s .

Fork
length,
mm 1955 1956 Total
300-399 71.5(14) 53 (17) 61.3 (31)
400-499 50 (4) 50 (6) 50 (10)
500-599 100 (4) 75 (4) 87.5 (8)
600-699 100 (3) 100 (4) 100 (7)
700-799 100 (3) 30 (20) 43.5 (23)
800-899 33.3 (3) 64.7(17) 60 (20)
900-999 100 (1) 60.9(23) 62.5 (24)
1000-1099 — 50 (6) 50 (6)
1100-1199 — 0 (2) 0 (2)
1200-1299 100 (1) 100 (1)
Total 75.8(33) 53.5(99) 59.1(132)

TABLE 29
F ood it e m s f o u n d in the 78
st o m a c h s o f a d u l t s p e c im e n s of Sphyraena
barracuda collected fr o m B im in i, B a h a m as, in Ju l y , 1955, and Ju l y , 1956.
N um bers are st o m a c h s in w h ic h it e m s w ere fo und .

Item 1955 (N = 2 5 ) 1956 (N = 5 3 ) Total

Pisces:
Clupeidae
Redear sardine, Harengula humeralis —
Synodontidae
Snakefish, Trachinocephalus my ops 1
Echelidae
Key worm eel, Ahlia egmontis —
Speckled worm eel, Myrophis punctatus —
Belonidae
Rec An needlefish, Strongylura notata 2
Holocentridae
Blackbar soldierfish, Myripristis jacobus 1
Lutjanidae —
Carangidae
Bar jack, Caranx ruber —
Mackerel scad, Decapterus macarellus 1
Bigeye scad, Selar crumenophthalmus —
Gerridae
Cuban mojarra, Gerres havana —
Sparidae
Calamus sp. —
Chaetodontidae
Chaetodon sp. 1
Scaridae
Striped parrotfish, Scarus croicensis 1
Blue parrotfish, Scarus coeruleus 1
Rainbow parrotfish, Scarus guacamaia 2
120
 TABLE 29 (Continued)
Item 1955 (N = 2 5 ) 1956 (N = 5 3 ) Total
Mugilidae
White mullet, Mugil curema — 1 1
Atherinidae
Hardhead silverside, Atherinomorus stipes 2 — 2
Bothidae — 2 2
Balistidae
Scrawled filefish, Alutera scripta — 2 2
Queen triggerfish, Balistes vetula — 1 1
Ostraciidae — 1 1
Tetraodontidae
Bandtail puffer, Sphaeroides spengleri — 4 4
Unidentifiable fishes 8 18 26
Arthropoda-decapod Crustacea
Pink shrimp, Penaeus duorarum — 2 2
Mollusca-Cephalopoda
Squid “pen” — 1 1
Plant material-Angiospermae
Turtle grass, Thalassia testudinum 6 2 8
Manatee grass, Cymodocea manatorum 1 — 1
Unidentifiable matter 2 7 9
Total 29 56 85

for only 5.3 per cent of the food of the adult barracuda. In adults, the
Beloniformes, Tetradontiformes, Carangidae, Mugilidae, and miscellaneous
Perciformes are more important and accounted for 66.7 per cent of the
food. However, these groups are present in only 6.5 per cent of the
stomachs of the young. These changes in diet reflect in part a change in
habitat with growth of the barracuda. The adult barracuda shows definite
preferences for two main categories of fishes: comparatively fast-swimming
fishes of the surface or mid-depths, such as hemiramphids, carangids, and
scombrids; and the sluggish, reef-dwelling tetraodontiform fishes. The
scarcity or absence of such active, reef-dwelling forms as lutjanids,
serranids, labrids, and pomacentrids in the diet seems noteworthy.
From a survey of the literature, and based on personal observations,
it appears that, in general, sphyraenids which do not attain a large size
feed mainly on small fishes which school, and upon cephalopods, whereas
the large species of Sphyraenidae which tend to be solitary feed upon
larger fishes which are essentially non-schooling in nature.
A ttacks by B arracuda on H um ans
Perhaps the most colorful accounts of barracuda are those which involve
their alleged voracity. Of the early reports by explorers and naturalists, de
Rochefort (1667) appears to be one of the first to record the barracuda’s
ferocity, and he believed that in addition to its desire for human flesh,
its teeth contained poison. Gudger (1918) gave an excellent and amusing
summary of the popular beliefs, citing the works of de Rochefort (1667),
121
 TABLE 30
Sum m ary o f fo o d it e m s fo und in sto m a c h s o f 529 s p e c im e n s of Sphyraena bar-
arcuda f r o m t h e t r o p ic a l w e s t e r n A t l a n t ic c o l l e c t e d f r o m 1920 through 1956,
SHOWING CHANGES IN FOOD H A B IT S WITH GROWTH. N U M B E R S ARE STOMACHS IN
WHICH ITEMS WERE FOUND.

Fork length (mm)

0 - 100- 200- 300- 400- 500- 600- 700- 800- 900- 1000- 11 GO- 1200-
Item 99 199 299 399 499 599 699 799 899 999 1099 1199 1299 Total
Gobioidei 15 11 26
Cyprinodonti-
formes 3 7 10
Atherinidae 17 11 — 2 30
Gerridae 3 5 3 1 12
Clupeiformes 14 5 — 1 2 — 1 1 — 3 — — — 27
Scaridae — 2 1 2 1 2 — 1 1 1 1 — — 12
Bothidae — — — — — — 2 1 — — — — — 3
Beloniformes 4 — — — 3 3 9 4 3 4 2 1 — 33
Tetraodon-
tiformes 3 4 5 5 9 i
X 27
Anguilliformes 2 3 2 — — — 7
Carangoidei 1 — — — — 1 2 4 4 6 4 — 2 24
Scombridae 3 2 2 1 — — 8
Mugilidae 1 2 3 3 1 2 — — 12
Other fishes 5 3 — — 1 — 2 4 7 2 3 1 1 29
Unidentifiable
fishes 106 26 3 7 7 17 9 14 10 6 205
Arthropoda
Crustacea 3 1 2 6
Mollusca
Cephalopoda 2 _ _ _
1 2 1 _ 1 _ 7
Plant material 3 — — 3 1 1 1 5 2 — — — — 16
Unidentifiable
matter 7 11 1 5 4 4 4 6 11 8 6 2 69
Total 184 82 8 21 12 21 44 49 57 51 26 5 3 563

Du Tertre (1667), Sloane (1707), Dampier (1729), Labat (1742),

Catesby (1754), and Fermin (1769).
Attacks have been authenticated by the unmistakable tooth marks of
a barracuda. As Gudger and Breder (1928) pointed out, its cleaver-like
bite can be distinguished from that of a shark in that it consists of two,
nearly parallel rows of tooth marks, whereas that of a shark leaves a
parabolic wound of jagged tooth marks. The shark also tends to clamp
its prey and tear off flesh with a sawing motion. Photographs of a barra­
cuda’s bite are given in Gudger and Breder (1928) and Tanous and
llano (1960), and that of a shark in Bolin (1954).
My research and observations indicate that the barracuda depends
almost entirely upon sight and usually makes a single attack which may
be severe but is seldom fatal. In general, a shark is further stimulated by
the presence of blood drawn so that repeated attacks occur which often
result in fatalities.
Literature Review of Attacks .— The first authenticated report by a barra­
cuda upon a human appears to be that of Pike (1873: 286, 294), who
122
related personal experiences on Mauritius; he was attacked by a large
barracuda which he was able to harpoon after a second attack. Gunther
(1877), on the authority of Andrew Garrett, wrote that barracuda were
dangerous to bathers in the Indo-Pacific. Klunzinger (1884) reported that
a native diver in the Red Sea died of wounds after having been attacked
by a specimen of Sphyraena agam ( —S. barracuda ) . A native of Cocos-
Keeling Atoll whose calf muscles had been torn away by a barracuda
was observed by Wood-Jones (1902: 322ff). Holder (1908, 1924) stated
that in the Florida Keys a man was attacked repeatedly by several barra­
cuda. Citing another instance, he noted that a barracuda, which had been
placed in the well of a sloop and forgotten, attacked and maimed a man
who entered it later. Holder also mentioned an episode at Sea Horse Key,
Bahamas, where a man suffered a severe bite on the arm while swimming.
Gudger (1918) reported the case of a man having been bitten on the
foot by a barracuda as he was walking on a shallow reef beside deep
water. Mowbray (1922) stated that a young woman while approaching a
buoy when swimming in Florida waters was attacked by a barracuda and
bled to death. Waite (1923) remarked that in South America a champion
lady swimmer lost her leg and died as the result of an attack by a large
barracuda. Two young girls were attacked by barracuda while swimming
off Miami Beach, Florida, according to Forbin (1924).
A valuable detailed and illustrated report was made by Gudger and
Breder (1928) of an attack by a barracuda upon a man swimming in
shallow water on the Atlantic side of Panama. They also referred to a
young woman who was fatally attacked by a barracuda off St. Petersburg,
Florida, some years prior to 1928. If the locality is correct, this report
differs from that reported by Waite (1923) or Forbin (1924), although
it may be identical with that reported by Mowbray (1922). Irvine (1947)
gave an account of a man who lost both hands after he was bitten by a
barracuda off Ghana. In popular account, Schultz and Stern (1948) noted
that “barracuda have ripped off the calves of swimmers [and] have
amputated legs and arms with a single bite.” Storr (1957) reported that
he was attacked by a barracuda which had apparently become irritated by
the white shoes of another diver. According to Storr (personal communi­
cation), the attack pattern was not completed.
Coppleson (1959), in his study of shark attacks in Florida waters, listed
two fatalities attributed to barracuda bites, one in 1947 upon a swimmer
at St. Augustine, and a second at Key West in 1952. Coppleson reported
an attack on a swimmer off Durban, South Africa, which was believed to
be by a barracuda. A fatal attack on a bather in 1958 at Key West was
reported by Hutton (1959) as possibly due to a barracuda. A fourth
attack, non-fatal, was listed by Hutton as occurring in 1953 off Juno Beach.
Tanous and llano (1960) presented case histories of four attacks by
predatory fishes in Florida. From their descriptions of the injuries, I believe
that these were all due to barracuda bites, I have described two of these
123
 in some detail in the following section. Of the remainder, one attack
occurred in the vicinity of Miami Beach in September, 1949, and was
inflicted upon the leg of a 37-year-old woman while she was swimming in
the afternoon. The other case involved a 49-year-old man who was bitten
on the foot at approximately 6:45 PM in August, 1955, in the vicinity
of Miami Beach.
On May 7, 1953, the Miami (Florida) Herald reported that a boy was
bitten on the foot while swimming close to a school of mullet about 50 to
75 feet off Haulover Beach, North Miami. Several stitches were required
to close five or six deep cuts. It was reported that the attacker was a
barracuda (Tanous and llano, 1960).
The Miami Herald reported on May 8, 1955, that a 15-year-old boy
was attacked underwater by a 4-foot barracuda which he had speared
off Crandon Park, Miami. After dragging him a few feet through the
water, the barracuda broke loose from the spear and attacked his right
side, leaving four large tooth marks which required 23 stitches.
On September 18, 1955, according to Mr. Tom Morgan (personal
communication) of the Collier County News, Naples, Florida, a young
girl was severely bitten about the arm and fingers in Bateman Cove on
the south side of Marco Pass by a fish which may have been a barracuda.
The Miami Herald carried an article on July 16, 1956, about a
19-year-old girl who was attacked while floating in an inner tube about
15 miles north of Myrtle Beach, South Carolina. The physician who
treated her reputedly described the injury as a “multiple, tearing wound
of the lower left leg, ankle and foot, severe and extensive,” which in his
opinion was due to a barracuda rather than a shark.
The Miami Herald for July 19, 1956, printed a similar story in which
a 38-year-old woman was “ . . . slashed repeatedly and savagely by a
barracuda in the shallow surf off . . . Miami Beach.” She was bitten six
times on her left leg, officials said, and once slightly on the calf of her
rieht leg. The victim suffered from shock and loss of blood, and was
subsequently hospitalized. The attending physician believed that the bite
was that of a barracuda and not of a shark. There were conflicting reports
on the number of attackers (Tanous and llano, 1960).
The New York Herald Tribune of July 17, 1957, reported that “a shark
or barracuda attacked and killed a swimmer . . .” off Atlantic Beach,
North Carolina, on July 16, 1957. The victim, who was an exceptionally
good swimmer, shouted to a lifeguard that a shark had bitten him.
Subsequent examination of the wounds by ichthyologists and physicians did
not resolve the identity of the assailant.
Dr. John E. Randall, presently of the University of Puerto Rico,
informed me that on March 5, 1957, a man speared a 5-foot barracuda
south of Key Biscayne, Florida. The fish broke the line and turned to
attack, at which point the man raised his right arm to protect himself,
with the result that he was slashed deeply on the forearm, requiring about
124
 70 stitches. Another attack was reported by Dr. Randall as follows: in
August, 1926, a man was swimming about 50 yards off Miami Beach, in
murky water which was over his head. The man sustained three bites on
his left arm, which was subsequently amputated. The surgeon removed
barracuda teeth from the wounds.
The Tampa Tribune carried an article in May 3, 1959, reporting an
instance off Miami where a barracuda leaped from the water and bit a
3-inch gash in the neck of a fisherman who was sitting in a small boat,
with the result that six stitches were required for the wound. No additional
details were reported.
On July 15, 1960, the Miami Herald reported that a 34-year-old skin
diver was attacked twice by a barracuda about Va -mile off Pompano
Beach, Florida, in about 20 feet of water. The 15-pound barracuda made
four rushes at the diver, two of which ended in attacks, resulting in severe
lacerations of the knee and wrist which required 31 stitches. The diver
wore no brightly colored apparel or jewelry, nor had he aggravated the
barracuda in any way. Neither the barracuda nor other fish had been
speared. Visibility was described as “fair.”
From the literature and newspaper accounts I have listed 29 separate
attacks attributed to barracuda upon humans; 19 are documented, two
are probable, and the remaining eight are possibly due to barracuda. The
reports of Pike (1873), Klunzinger (1884), Wood-Jones (1902), Holder
(1908, 1924), Gudger (1918), Gudger and Breder (1928), Irvine
(1947), Coppleson (1959), Hutton (1959), and Tanous and llano
(1960) contain evidence as to the identity of the attacker. Personal reports
from Randall, and newspaper accounts of May 7, 1953, off Haulover
Beach, Miami, that of May 8, 1955, of the assault off Miami, that of July
19, 1956, on the attack off Miami Beach, and that of July 15, 1960, off
Pompano Beach are due to barracuda. The report on the attack of July
16, 1956, off South Carolina, and that off Durban, South Africa, in
1946 reported by Coppleson (1959) were probably made by barracuda.
One of the attacks (1952) reported by Coppleson and one (1958) by
Hutton were probably made by barracuda. The identity of the species off
Naples, Florida, on September 18, 1955, is questionable; the nature of
the wound and the single attack suggest that it was a barracuda. Literature
reports of Mowbray (1922), Waite (1923), Forbin (1924), and one of
the cases in Gudger and Breder (1928) are lacking in details and locality
of the attack. According to data presented by Coppleson (1959), possibly
these may refer to the same attack; it is debatable that they were due to
barracuda. The identity of the attacker in the fatality off North Carolina
on July 16, 1957, is questionable.
In the tropical western Atlantic, barracuda is the only sphvraenid which
attains sufficient size to inflict a wound and it probably made most of the
attacks listed above. In other areas, it is possible to identify the attacker
on an a priori basis, since only S. barracuda , S. jello Cuvier, S. jorsteri
125
Valenciennes, and S. obtusata Cuvier attain appreciable size. In general,
the distributions of these large species do not overlap.
Possible Causes of Attacks .— An interesting account of a group of about
40 British frogmen and their experiences with sharks and barracuda in
the Bahamas is given by Wright (1948), who stated that the barracuda is
feared by the natives because it can come into much shallower water than
can a large shark. He concluded that under certain conditions barracuda
and sharks should be feared, but that normally they were harmless. His
inferences were as follows:
“ (1) A man in the water, alone or in a group, acting quietly and
smoothly, did not release the attack pattern of either shark or barracuda.
(2) Blood without movement did not release a feeding pattern in shark
or barracuda in the Bahamas. (3) Fish blood and jerky, rapid movements
released the attack pattern in both shark and barracuda. (4) Jerky, rapid
movements alone released the attack pattern in both shark and barracuda.”
The frenzied reaction of a barracuda to a vibrating or bleeding fish on
the end of a spear is a rather well-known but unnerving experience to
spearfishermen. Wright (1948) reported a case where a barracuda rushed
in and seized a small, bleeding sting ray and jerked the ray and spear
out of the man’s hand and made off with both. Off Cat Cay, Bahamas,
I speared a Nassau grouper (Epinephelus striatus) which subsequently
wrenched the spear from my hand and became wedged under a rock
ledge. During my efforts to dislodge it, the grouper struggled violently
and emitted croaking sounds, and shortly four barracuda from 3 to 4
feet long appeared in the vicinity. When dislodged the grouper was
bleeding profusely and swimming erratically from the weight of the spear,
which was still inserted; as I grasped the spear and swam toward the
surface, one of the barracuda seized the grouper and made off with it and
the spear. In the Florida Keys, while collecting specimens I have noted
that parrotfish, groupers, and grunts, in particular, all flap rapidly and
bleed profusely when speared, and groupers and grunts also emit croaking
sounds, and often barracuda are attracted; however, only in the instance
described above was the specimen carried off. An injured fish or live
grunt with several diagonal cuts is a well-known bait for large barracuda
in the Florida Keys.
A flashing object is quickly discerned by a barracuda; hence, in waters
inhabited by barracuda it is advisable to remove shiny objects such as a
ring, bracelet, or wristwatch. On one occasion, in rather turbid water off
Key Largo, Florida, a barracuda about 2 feet long charged at and hit
my face mask which may have reflected the sun’s rays. In another
instance I was photographing some coral formations in about 10 feet of
water using a camera enclosed in a lucite case when a barracuda about
3 V2 feet long appeared. It approached suddenly to within about 4 feet of
the camera but soon backed off to a distance of about 15 feet and began
to swim erratically at first and eventually in a figure-8 pattern. As the
126
camera was pointed downward, the barracuda showed no excitement,
but each time the camera case was directed toward the fish it showed
interest. On several occasions it approached the camera rapidly but turned
sharply away and swam in a wide circle. Its behavior was similar to
that of a barracuda confronted with a speared fish.
On about 40 occasions when swimming about Bimini and Cat Cay,
Bahamas, and in the Florida Keys, I encountered barracuda on nearly
every trip. They ranged in size to perhaps 5 feet, but on no occasion was
I ever attacked. However, on any occasion it was possible to elicit an attack
pattern by spearing a small grunt or parrotfish when barracuda were close
by.
If present in the area, barracuda congregated immediately upon one’s
entrance into the water, particularly if it was accompanied by splashing,
but shortly thereafter they usually disappeared. Occasionally a stray would
be encountered and would follow at a distance of perhaps only 10 or
15 feet as I moved from one reef to the next.
Based on literature and newspaper reports, as well as personal experi­
ences, there is only one instance known in which an unprovoked, in the
strictest sense, barracuda has attacked a swimmer who was completely
submerged (July 14, 1960). In the May 8, 1955 report of an attack on
a 15-year-old boy off Crandon Park, Miami, and the similar attack on
the man off Key Biscayne in 1957 the attacks were made by an injured
barracuda. Numerous barracuda are speared in the Florida Keys annually,
but few, if any, attempt to attack the diver. Miss Kathleen Parker
(personal communication) informed me that a barracuda slid down a
spear and made a slash in a boy’s leg which required 12 stitches. Although
I have never speared a barracuda weighing over 8 pounds, divers who
have taken them up to 50 pounds informed me that they were never
menaced or attacked after the fish was speared.
There are 15 separate instances where barracuda were definitely or
probably involved in attacks where the swimmer was splashing at or near
the surface, swimming in turbid waters, swimming toward dusk when
visibility is reduced, or a combination of these factors. If the “possible”
attacks are added, then another seven attacks can be considered as resulting
from splashing or swimming. Such attacks, although they might be
considered ones of “mistaken identity,” probably should be considered as
being provoked in the sense that the victim was splashing or flashing
suddenly under conditions which elicited an attack pattern. Probably
barracuda are less able to discern their prey in turbid waters, and in such
situations the risk of attack is higher than in clear waters.
In summary, conditions which appear favorable to an attack by an
unprovoked barracuda are: (1) if the swimmer were splashing at the
surface of the water; (2) if the water were turbid; or (3) a combination
of these two. To these, Springer (1943) added flashing light, color, great
speed, and irregularity of movement.
127
 P o is o n in g o f H u m a n s b y B a r r a c u d a
Fish and Cobb (1954) wrote that there is probably more controversy
over the toxic properties of the family Sphyraenidae than in any other
group of poisonous fishes. Some specimens of barracuda are poisonous,
while others are safe to eat, according to where and when they are caught,
and the size of the fish. Since poisonings can occur from eating a freshly
caught barracuda, the toxin is evidently not related to bacterial decompo­
sition, and is probably associated with certain factors in its life history
and ecology. Thus it seems apropos to investigate some of these factors
which may be responsible for its occasionally poisonous nature.
References to poisonings by the barracuda which date back several
hundred years are found in papers by Gudger (1918, 1930). The subject
of poisonous aquatic organisms has been reviewed in some detail by the
writer in his doctoral dissertation (de Sylva, 1958) on file in the Cornell
University library, and the discussion herein will be limited essentially to
the poisonous properties of barracuda and its food.
General R eview of Poisonous Fishes.— Resumes on the historical back­
ground of poisonous fishes are found in the works of Moreau de Jonnes
(1821), Dumeril (1866), Poey (1866), Pellegrin (1899), Byam and
Archibald (1921), Phisalix-Picot (1922), Hoffmann (1927, et se q .),
Pavlovskii (1927), Matsuo (1934), Maass (1937), Hiyama (1943),
Halstead (1953), Phillips and Brady (1953), Halstead and Lively
(1954), Wheeler (1954), Fish and Cobb (1954), Loison (1955), Jones
(1956), Mills (1956), Ebroin (1956), Randall (1958), and Bartsch, et
al. (1959). Certain groups of fishes are more often reported to be poi­
sonous when eaten. These include large individuals of the phvlo^enetically
unrelated families Sphyraenidae, Carangidae, Serranidae, Lutjanidae, Mu-
raenidae, and most families of the order Tetraodontiformes.
Fish poisoning in the sense used here refers to that in humans due to
the ingestion of flesh which is perfectly fresh (Fereuson, 1823; Dunlop,
1917; Clark, 1920; Yasukawa, 1934; Jones, 1937; Gilman, 1942; Hiyama,
1943; Goe and Halstead, 1955), and in which leucomaine-like symptoms
are produced. This type is generally known as ichthyosarcotoxism (Hal­
stead and Lively, 1954), and as ciguatera in the tropical western Atlantic
(Randall, 1958).
Characteristics of Poisonous Barracuda. — The most commonly implicated
member of the family Sphyraenidae is S. barracuda. Other species which
have been cited as being poisonous are S. jello Cuvier (Fonssa<?rives and
de Mericourt, 1861), S. forsteri Cuvier (Hiyama, 1943; Smith, 1947), and
S. sphyraena L. when caught in the tropical waters of West Africa (Irvine,
1947). Sphyraenids which are considered safe to eat generally do not
attain a large size. Barracuda weighing less than 3 pounds are considered
non-toxic, and it is known that larger specimens of a given species are
more likely to be poisonous (Fonssagrives and de Mericourt, 1861;
128
Jouan, 1863b; Poey, 1866; Dumeril, 1866; Pellegrin, 1899; Rogers, 1899;
Nichols, 1912; Clarke, 1920; Byam and Archibald, 1921; Walker, 1922;
Bloedorn and Hakansson, 1926; Hoffmann, 1927; O’Neill, 1938; Gilman,
1942; Strong, 1942; Hiyama, 1943; Paetro, 1956; de Sylva, 1956; Calvo
and Alvarez, 1957). It also appears that the larger the barracuda, the
more violent will be the action of the toxin on humans (Yasukawa, 1934;
Hiyama, 1943; Smith, 1947).
In the barracuda as well as in species of some other families, the visceral
organs and blood are thought to be most poisonous. Gunther (1880) and
Mills (1956) believed that poisonous fishes are often edible if the head
and intestines are removed immediately after capture. Yasukawa (1934)
observed that the location of the poison varied with each species. The
viscera, particularly the gonads, and the skin are known to be most toxic
in tetraodontiform fishes (Remy, 1883; Hiyama, 1943; Tani, 1945), but
the flesh is often toxic, depending upon the stage of the reproductive cycle.
Fortunately, barracuda poisoning is infrequent, and deaths are rare.
However, mortalities resulting from the ingestion of fresh barracuda have
been reported by Guichenot (1843: 164), Morton (1868), Dunlop
(1917), and Calvo and Alvarez (1957). Poey (1866) and Clarke (1920)
each reported two fatalities, Morice (1958) reported three, and Norman
and Fraser (1938: 172-173) noted that an entire family perished after
having eaten barracuda. Von Fraenkel and Krick (1945), citing a native
sanitary officer in the Marianas Islands, noted that during the German
occupation there was an 8 to 10 per cent mortality rate among the natives
who had been poisoned from eating barracuda. In February, 1961, a
43-year-old native of Bimini, Bahamas, died following a meal of fresh
barracuda, and there is a report of an American sailor who died of anal
rupture following a severe case of poisoning from a barracuda caught in
Cuba.
Non-fatal cases of poisoning resulting from eatinq fresh barracuda are
reported by Moreau de Jonnes (1821), Sauvage (1865), Dumeril (1866),
Poey (1866), Gatewood (1909), Mowbray (1916), Gudeer (1918),
Castellani and Chalmers (1919), Clarke (1920), Byam and Archibald
(1921), Hoffmann (1927, 1929), Hoffmann and Embil (1928), O’Neill
(1938), Gilman (1942), Hiyama (1943), Von Fraenkel and Krick
(1945), Arcisz (1950), Hashimoto (1956), Paetro (1956), Ebroin
(1956), de Sylva (1956), Calvo and Alvarez (1957), Morice (1958),
and Ellington (1959).
Corroborated poisoning records by fresh barracuda in the tropical
western Atlantic which have appeared in the literature, supplemented by
information supplied me by victims of 13 separate incidents of barracuda
poisoning, mostly from southern Florida, which involved at least 89 per­
sons, appear in Table 31. In all parts of the Caribbean, there were 32
cases reported which involved at least 249 persons: these do not include
those by Ellington (1959) who reported at least 50 cases of barracuda
129
- TABLE 31
o A u t h e n t ic a t e d records of p o is o n in g in hum ans fro m e a t in g barracuda in the t r o p ic a l w ester n A t l a n t ic ,
COLLATED FROM THE LITERATURE AND SUPPLEMENTED BY PERSONAL INTERVIEWS.

Date of Date of Locality of No. of persons Size of

poisoning capture capture affected fish Author
Sept., 1921 ? Virgin Islands 1 About 3 lbs. a
Oct., 1921 ? Virgin Islands 2 About 3 lbs. a
May 20, 1941 May 19, 1941 Culebra, Virgin Islands 10 20 lbs. (male) b
“Middle of 1926” “Previous San Antonio, Cuba 23 ? c
afternoon”
Oct., 1927 “Previous day” Rio Almendares, Cuba 8 ? c
April, 1931 ? Camajuam, Cuba 36 ? c
“Middle of 1931” “A few hours Havana, Cuba 1 ? c
before”
May, 1944 ? Virgin Islands 2 ? d
Aug., 1945 ? Virgin Islands 3 ? d
Aug. 8 , 1945 Aug. 6 , 1945 Harbor of St. Thomas, 3 4!/2 ft. d
Virgin Islands, near sub­
marine base
July 11, 1951 ? Terre-de-Bas, 15 ? e
lies de Saintes, Guadeloupe
April, 1954 Same day Off Miami, Florida 5 “Large” h
May 12, 1954 May 11, 1954 Ft. Lauderdale, Florida 5 61/2 lbs. f
June 27, 1954 ? Hillsborough Inlet, Pompano 7 ? f
Beach, Florida
July 9, 1954 July 8 , 1954 Ft. Lauderdale, Florida 8 10 lbs. f
Aug. 24, 1954 July 4-6, 1954 Pompano Beach to 5 15 lbs. f
Boca Raton, Florida
Aug., 1954 ? Rancho Veloz, 21 ? g
Havana, Cuba
Nov., 1954 Oct., 1954 Off Miami, Florida 4 10 lbs. h
Feb. 27, 1955 Feb. 27, 1955 Tavernier, Florida 5 About 16 lbs. h
July 17, 1955 July 8 , 1955 Fowey Rocks, 4 3 ft. h
Miami, Florida
 TABLE 31 (Continued)

Date of Date of Locality of No. of persons Size of

poisoning capture capture affected fish Authority
Mar. 18, 1956 Mar. 17, 1956 Off Miami, Florida 10 4 ft. h
April 1, 1956 April 1, 1956 Biscayne Bay, 3 10 lbs. h
Miami, Florida
June 27, 1956 June 22, 1956 Off Miami, Florida 2 ? h
July 29, 1956 July 29, 1956 5 miles off Miami 12 15 lbs. h
Beach, Florida
Aug. 27, 1956 Aug. 26, 1956 Fowey Rocks, 7 22-25 lbs. h
Miami, Florida
Feb., 1957 Same day Off Miami, Florida 5 “Large” h
Aug., 1958 ? Vicinity of ? ? i
Cocoa Beach, Florida
Feb. 21, 1959 Feb. 20, 1959 5 miles north of 5 10 lbs. i
Fowey Rocks,
Miami, Florida
Dec. 10, 1960 Dec. 9, 1960 Key Largo, Florida 5 151/2 lbs. h
Feb. 8, 1961 Feb. 7, 1961 Bimini, Bahamas 11 “Large” h
Aug. 18, 1962 Aug. 16, 1962 Off Government 12 3 specimens, h
Cut, Miami, Florida 4-20 lbs.
Oct. 31, 1962 Same day 1-3 miles off 9 10-12 lbs. h
Palm Beach Inlet
aWalker, 1922
bGilman, 1942
cHoffmann and Embil, 1931
dArcisz, 1950
eEbroin, 1956; Morice, 1958
fPaetro, 1956
gCalvo and Alvarez, 1957
hPersonal interview or newspaper account
'Randall, personal communication
poisoning in the Cayman Islands over 18 months, but no further
data are available on dates. All symptoms were diagnostic of ichthyosarco-
toxism rather than bacterial poisoning. Undoubtedly there are many
unreported cases, since barracuda is usually sold under other names, and
because mild cases are often unreported or are blamed upon bacterial
poisonings.
A sample of flesh was obtained from a barracuda about 3 feet long
which had poisoned a family of four. It was caught on July 8, 1955, in
the vicinity of Fowey Rocks, south of Miami, and was immediately placed
on ice but was not cleaned until the family reached home. Shortly after
eating the fish the members of the family experienced typical symptoms
of fish poisoning; the effects were felt for about four months.
Dr. Louis F. Westbrook of the Dade County Department of Health
made a bacteriological analysis of a sample of the frozen flesh and found
that an alpha-type Streptococcus, which could not have caused the severe
symptoms reported, was the only pathogenic microorganism isolated.
Paetro (1956) reported that a sample of flesh from a barracuda which had
poisoned five persons was found to be negative for organisms commonly
associated with bacterial food poisoning when analyzed by the Florida State
Board of Health. Other workers such as Yasukawa (1934) and Arcisz
(1950) who have attempted to isolate pathogenic bacteria from the fresh
flesh of toxic fishes have met with negative results.
Identification of poisonous barracuda has not been accomplished (de
Sylva, 1956; Calvo and Alvarez, 1957), other than by feeding the flesh
to experimental animals. Popular beliefs by natives to distinguish toxic
barracuda include blackish or purplish color at the base of the teeth,
blackening of silver coins or spoons when cooked with the flesh, the
bitterness of the liver, the presence of a milky substance under the scales,
and loose or deciduous scales (Plee in Cuvier and Valenciennes, 1829;
Dumeril, 1866; Gudger, 1918; Byam and Archibald, 1921; Halstead and
Lively, 1954). Mowbray (1916) and Gudger (1918) reported that
barracuda having bluish backs (“blue-back” ) are considered poisonous.
The term “blue-blue” is applied at Bimini, Bahamas, to barracuda thought
to be poisonous, and Ellington (1959) reports that the flesh of poisonous
barracuda in the Cayman Islands has a bluish or blue-black appearance in
contrast to the normal light grey.
A widespread belief among the natives from various parts of the world
is that a lean fish, including barracuda, particularly one in which the liver
tastes bitter and peppery, is poisonous (Labat, 1724; Dampier, 1729;
Fermin, 1769; Poey, 1866; Hiroa, 1938), but that fat fish are safe to
eat (Mills, 1956). The flesh of barracuda is of excellent flavor and is
preferred to other species by natives of Cayman Islands, according to
Thompson [1945?!, even though barracuda are often poisonous there
(Ellington, 1959). Victims of barracuda poisoning have often commented
upon the unusually fine flavor (O’Neill, 1938), or that it was the best
132
fish ever tasted (Paetro, 1956). The only other comment on the uniqueness
of the fish made by any of the victims seems to be that the fish was
unusually difficult to scale or to skin. Mills (1956) wrote that poisonous
individuals . . are dry and hard on cooking and the skin creases like
rubber. If the flesh of a fish is fatty it is always safe.” In Cuba, it is
believed that fresh barracuda are poisonous which have scales which stand
out or are deciduous, and in which the meat is soft. However, none of the
above is infallible, and Von Fraenkel and Krick (1945) stated that even
in the Marianas Islands, where the incidence of fish poisoning is high,
the natives are unable to identify a fish which is poisonous.
Toxin from Barracuda Flesh.— In some of the tetraodontiform fishes, the
chemical nature of the poison has been studied by Takahashi and Inoko
(1889), Tahara (1910), and Ishihara (1918). In other groups the
chemical nature of the toxin is unknown but is thought to be of an
alkaloid type which produces effects similar to that of curare (Strong,
1942; Simmons, et al., 1944) or aconitine (Ross, 1947).
A virulent poison was isolated by Hashimoto (1956) from a speci­
men of S. picuda ( = S. barracuda) about 1 meter long, presumably
from the tropical western Pacific. The barracuda, which was preserved in
good condition after it was caught, had poisoned about 30 persons of five
families. The symptoms in those afflicted were typically those of ichthyo-
sarcotoxism. Hashimoto found that the skin of this individual produced
toxic symptoms when fed to animals, but he refrained from deriving
conclusions about the toxicity of the skin because it was taken from the
boiled fillet. Hashimoto concluded that “. . . the poison is of quite a new
type, which has never been found in aquatic animals,” and differed
from the paralytic poison found in the family Tetraodontidae (Tahara,
1910; Hashimoto and Migita, 1951), shellfish (Gibbard and Naubert,
1948; Hashimoto, et al., 1950; Hashimoto and Migita, 1950; McFarren
and Bartsch, 1960), and the histamine-like substances isolated from
scombroids (Halstead and Lively, 1954). Hashimoto (1956) found that
the toxin was not harmful when administered to mice, but had severe
effects upon cats. Poisonous barracuda may have a more marked effect
upon cats than upon humans, and ingestion of even small amounts of flesh
invariably ends with the death of the cat (Locke, 1675; Hoffmann, 1929;
Hoffmann and Embil, 1931; O’Neill, 1938; Gilman, 1942; Hiyama, 1943;
Paetro, 1956; Larson and Rivas, 1962). O’Neill (1938) reported that in
the Virgin Islands a fisherman’s cat which had ingested some of the blood
of a barracuda was paralyzed in the legs the next day and eventually died.
Hiyama (1943) noted that a cat which was fed cooked flesh from the
region of the pelvic fin of a 91-cm barracuda died about 14 hours later;
however, another cat which was fed raw flesh from the tail of the same
fish did not die.
Etiology of Fish Poisoning.— The theories which have proposed to account
for the occasional toxicity of barracuda, as well as other species, are
133
summarized by Gilman (1942), Halstead (1953), Paetro (1956), de Sylva
(1956, 1958), and Randall (1958). Knowledge of the etiological factors
responsible have progressed little beyond Martyr’s conclusion in 1530 (in
Gudger, 1930) that barracuda become poisonous because they feed on
the berries of the manchineel plant, Hippomane mancinella L. (Gudger,
1930; Calvo and Alvarez, 1957).
Of the various hypotheses which have been proposed to account for
the origin of the toxin(s) in barracuda, only two seem plausible: (1) the
poison occurs as a result of spawning (Costa Mandry, 1933; Strong, 1942;
Brown, [1945?]), or (2) the toxin is acquired through the food
(Moreau de Jonnes, 1821; Dumeril, 1866; Gunther, 1880; Rogers, 1899;
Pellegrin, 1899; Yasukawa, 1934; Hiyama, 1943; Halstead and Lively,
1954).
Relation to Spawning .— It is known that the gonads of tetraodontid fishes
become poisonous during the spawning season, after which some of the
toxin may diffuse to other organs, as well as the flesh and the skin (Remy,
1883; Yasukawa, 1934; Hiyama, 1943; Tani, 1945). Yet there is little
evidence at hand to disclose why such predatory species as certain scom-
brids, serranids, carangids, and lutjanids are more apt to be poisonous
when large. The widespread belief that barracuda weighing less than 3
pounds are non-poisonous (Poey, 1866; Hoffmann, 1927; Hiyama, 1943,
and others) seems generally to be true. This arbitrary figure of 3 pounds
might correspond to the onset of maturity, since females mature at a
weight of from 3.1 to 3.9 pounds (580 to 660 mm fork length); however,
males mature earlier, at a weight of from 1.3 to 1.9 pounds (460 to 500
mm). Thus while it is possible that a toxic substance develops in the
female at the onset of maturity, the males would also be expected to be­
come toxic at maturity. If this were true, however, certainly a few poison­
ings by sexually mature males weighing less than 2 pounds would have
occurred.
Watanabe (1946) has suggested that when fishes caught only in a cer­
tain locality are poisonous these represent mature fish which have schooled
for spawning. This explanation would not, however, explain why only one
of two fish of identical size and taken from the same place can cause
poisoning. Hiyama (1943) found no differences in toxicity between the
sexes in a large number of poisonous individuals from widely different
families. Some freshwater fishes, notably Lepisosteus and some European
cyprinids, are among those which have poisonous ovaries (Phisalix-Picot,
1922; Pavlovskii, 1927). Although Brown [1945?] noted that in the Lee­
ward Islands the ovaries of certain fishes, including the barracuda, were
sometimes regarded as dangerous to eat, Ackerman (1952) wrote that in
Florida, ripe ovaries of barracuda are considered a delicacy and are eaten
without harm. But the spawning habits of most tropical marine
species are unknown, and there is little evidence relating to the production
of toxin to spawning.
134
 The spawning season of barracuda in southern Florida is late March to
mid-October. The distribution of poisoning by freshly eaten barracuda
which caused 30 separate instances of poisoning taken from the data in
Table 31 is as follows:
MONTH JAN. FEB. MAR. APR. MAY JU N E JULY
Frequency 0 4 1 3 3 2 4
AUG. SEPT. OCT. NOV. DEC. TOTAL
7 1 3 1 1 30
Poisonings occurred in February, November, and December, which do
not include the spawning season, and hence it seems unlikely that spawning
is responsible for toxin production in barracuda. However, work by Takata
(quoted in Randall, 1958) indicating that toxin could be maintained in
living fish flesh for at least 14 months may conceivably invalidate the
above analysis.
Relation to Diet. — That barracuda acquire a toxin from eating poisonous
food is supported by a large body of circumstantial evidence. Certain
tropical marine animals appear to be poisonous nearly all of the time in
some localities. Yet there seems to be no evolutionary pattern followed
within a group in the development of a toxin, with the exception of tetra­
odontiform and anguilliform fishes. The former group includes the families
Diodontidae, Tetraodontidae (sensu lato), Balistidae, Monacanthidae, and
Ostraciidae, as well as the primitive tetraodontiform families Acanthuridae
and Chaetodontidae. However, families such as the Clupeidae, Sphyraeni-
dae, Carangidae, Scombridae, Serranidae, Lutjanidae, Muraenidae, and
Scaridae are not closely related; these groups and the tetraodontiform fishes
account for most of the poisonings.
We must consider the poisonous principle of the organisms in relation
to their habits and/or position in the food web. Those who have attempted
to explain fish poisoning in light of the latter have often oversimplified the
food web in the tropical reef environment. I have attempted to present
the generalized interrelationship among organisms found on a typical coral
reef in southern Florida (Fig. 35). The complexity of this relationship
indicates that several paths may be taken in the transmission of a toxin
from its source to the ultimate consumer.
Toxic Organisms in the Food Web of the Barracuda.— The following dis­
cussion pertains only to species found in shallow, reef areas where the
incidence of toxic organisms is high. The families Sphyraenidae, Carangi­
dae, Scombridae, Muraenidae, and large individuals of the families Ser­
ranidae and Lutjanidae include the most important fish-eating teleosts, and
are also the most frequently incriminated in outbreaks of fish poisoning,
save for the tetraodontiform fishes. A few of the many references on the
poisonous nature of predatory fishes are those of Dumeril (1866), Mow­
bray (1916), Dunlop (1917), Clarke (1920), Byam and Archibald
(1921), Gregory (1925), Hoffmann (1927), Pavlovskii (1927), Costa
135
 F igure
of southern Florida.
FISH-EATING TELEOSTS

(large individuals of the genera Sphyraena, Seriola, Caranx, Scomberomorus, Epinephelus, rca, Lutjanus)
35. Generalized relations among organisms of the coral-reef habitat

DETRITUS-FEEDERS AND BROWSERS INVERTEBRATE-FEEDERS

(Sphaeroides. Balistes, Monacanthus. (Haemulon, small serranids,
Acanthurus. Chaetodon, Alutera) carangids and lutjanids)

\ ZOOPLANKTON
(copepods, zoea)

DETRITUS BENTHIC ALGAE

(Chlorophyceae, Myxophyceae, PHYTOPLANKTON
coralline Rhodophyceae) (Myxophyceae, Dinoflagellata)
Mandry (1933), Yasukawa (1934), Mann (1938), O’Neill (1938),
Gilman (1942), Hiyama (1943), Brown [1945?], and Halstead and
Lively (1954).
Among the fishes which are essentially detritus-feeders, the tetraodonti­
form fishes occupy a wide variety of habitats but are all closely associated
with the coral-reef habitat. The highly poisonous nature of this group is
treated specifically by Anderson (1776), Pappe (1853), Richardson
(1861), Jouan (1867), Remy (1883), Takahashi and Inoko (1889),
Seale (1912), Kerr (1912), Bryan (1915), Ishihara (1918), Phisalix-
Picot (1922), Nagai and Ito (1939), Larsen (1939), Hiyama (1943),
Tani (1945), Yudkin (1944; 1945), Hashimoto (1950), Clark and Gohar
(1953), Jones (1956), and Benson (1956).
Those feeding on invertebrates such as pomadasyids, labrids, and small
individuals of the families Serranidae, Lutjanidae, and Carangidae have
not been often implicated in cases of poisoning, although when small some
pomadasyids and lutjanids are said to be poisonous in the Virgin Islands
(Walker, 1922). When large serranids, lutjanids, and carangids are prin­
cipally fish-eaters.
Coral-feeders are represented by numerous members of the family
Scaridae, and certain balistids and tetraodontids. Cases of poisoning by
scarids and balistids have been recorded by Sonnerat (1774), Cloquet
(1821), Moreau de Jonnes (1821), Valenciennes in Cuvier and Valen­
ciennes (1839), Poey (1866), Dumeril (1866), Hill (1868), Gunther
(1880), Tybring (1887), Dunlop (1917), Clarke (1920), Byam and
Archibald (1921), Walker (1922), Pavlovskii (1927), Simmons, et al.
(1944), Halstead and Bunker (1954a), and Mills (1956). The scarid
genera Scarus and Pseudoscarus, as well as several related genera, have
often been implicated in cases of death, but the geographic variation in
toxicity is marked, perhaps more so than in any other group.
The next groups in the food chain depicted in Figure 35 are benthic
invertebrates and the forage fishes. The following invertebrates reputedly
have caused poisoning in humans:
Coelenterates (Calmette, 1908)
Polychaetes (Burrows, 1945)
Prawns (MacGowan, 1887)
Crabs (Du Tertre, 1667; Sloane, 1725; Damoier, 1729; Poey, 1866;
Jouan, 1867; Roger, 1895; Banner, 1952)
Spiny lobsters (Dunlop, 1917; Clarke, 1920)
Gastropods (Poey, 1866; Banner, in Banner and Randall, 1952)
Pelecypods (Gibbard and Naubert, 1948; Riegel, et a l, 1949; Hashi­
moto, et a l, 1950; Edwards, 1956)
Holothurians and echinoids (Frey, 1951; Nigrelli and Zahl, 1952;
Catala, 1957)
The above groups are essentially browsers and filter-feeders, and serve
as food for a number of fishes. Hashimoto, et al. (1950) have isolated a
137
toxin, venerupin, from Japanese mussels, and toxins have been extracted
from Atlantic mussels (Gibbard and Naubert, 1948) and Pacific mussels
(Riegel, et al., 1949).
Food habits of planktivorous filter-feeding fishes, such as the clupeids,
in tropical waters are poorly understood. Since they strain large quantities
of zoo- and phytoplankton from the waters, they probably ingest a number
of toxic plankton organisms, particularly in the tropics where blue-green
algae and dinoflagellates are the dominant phytoplankters. In New Cale­
donia and in certain parts of the tropical western Atlantic, Meletta (—Clu~
panodon) venenosa and Meletta thrissa (—Opisthonema oglinum) are well
known for their poisonous nature (Locke, 1675; Moreau de Jonnes, 1821;
Cloquet, 1821; Ferguson, 1823; Stephenson, 1838; Montrouzier, 1857;
Fonssagrives and de Mericourt, 1861; Hill, 1868; Gunther, 1880; Remy,
1883; Rogers, 1899; Dunlop, 1917; Clarke, 1920; Simmons, et al., 1944;
Mills, 1956). Ingestion of these species often rapidly causes fatalities.
Certain species of Harengula, notably macrophthalma and humeralis, also
acquire toxic properties in some areas (Duhamel du Monceau, 1782;
Hill, 1868; Remy, 1883).
The toxicity of phytoplankton has been reviewed by Mason and Wheeler
(1942), Steyn (1945), Prescott (1948), Louw (1950), Vinberg (1954),
Ingram and Prescott (1954), Dawson, et al. (1955), and Habekost, et al.
(1955).
Dawson, et al. (1955) and Habekost, et a l (1955) have shown that
certain marine green and blue-green algae, principally Caulerpa and
Lyngbya, as well as some coralline red algae which are toxic to laboratory
mice, are important in the diet of some toxic marine fishes (e.g., Acanthu -
rus, Chaetodon, Abudefduf, Arothron).
From the above review, the presence of toxins in various marine organ­
isms is well established, and the hypothesis that a basic toxin is transmitted
through several paths of the food web of marine organisms should be in­
vestigated pharmacologically. Studies by Takahashi and Inoko (1889),
Ishihara (1918), Gibbard and Naubert (1948), Hashimoto and Migita
(1951), Vinberg (1954), and Banner, et al. (1960) indicate that the
pharmacological properties of the toxins in various trophic levels may
differ, although the generic toxins within a trophic level are generally
similar. Much work remains to prove that these toxins do not undergo
modification during transference between successive trophic levels, in spite
of the apparent pharmacological differences among these toxins.
Although the symptoms of these toxins are not identical, with the pos­
sible exception of shellfish and puffer poisoning, the symptoms fall within
the ciguatera-like syndrome.
Food Habits and Ecological Relations of Organisms in the Food Web of
the Barracuda.— A number of groups which include poisonous species are
eaten by the barracuda (Table 30). These include members of the Tetra-
138
odontiformes, Gobiiformes (Pavlovskii, 1927), Clupeiformes, Carangidae,
Scaridae, Beloniformes (Moreau de Jonnes, 1821; Cloquet, 1821), Scom-
bridae, and Anguilliformes. Barracuda over 3 pounds generally cause
poisoning, and the important dietary items of the adult are the Tetraodon-
tiformes, Beloniformes, Carangidae, and Scaridae, although there is doubt
as to the importance of the species of Beloniformes as natural food.
Tetraodontiform fishes occurred most often in the diet of the adult barra­
cuda in southern Florida and Bimini, being found in 20 per cent of the
stomachs which contained identifiable fishes. Species which have reputedly
caused poisoning in humans were found in 58.1 per cent of the stomachs
of the adults which contained identifiable food (Tables 26-30).
Geographical Distribution of Toxic Barracuda .— One of the most perplex­
ing problems in the study of poisonous barracuda is their bizarre geo­
graphical distribution within a relatively small area. Native fishermen from
such widely scattered points as New Caledonia, the Marianas Islands, and
the Bahamas are reported to insist that barracuda, as well as many other
species, caught on one side of a reef or island are safe to eat, while they
are poisonous on the other side of the reef only a short distance away
(Jouan, 1861, 1863b; Rogers, 1899; Pellegrin, 1899; Mowbray, 1916;
Walker, 1922; Yasukawa, 1934; Hiroa, 1938; Mann, 1938; Thompson,
[1945?]; Brown [1945?]; Mills, 1956). Yasukawa (1934) observed that
poisonous fishes may or may not be toxic at different Pacific islands or even
at different places on the same reef, and suggested that perhaps a particular
type of food was found only in certain places. Hiyama (1943) was evi­
dently the first to advance the theory that there was a relation between the
distribution of poisonous fishes and that of coral reefs, more specifically the
nematocysts of the coral polyps. Daget (1954) noted that a puffer, Tetrao-
don fahaka, which is poisonous elsewhere in its range, is eaten without ill
effects in the upper Niger River, where no corals exist.
Ecological relationships in the food web of poisonous fishes have been
briefly discussed by Yasukawa (1934), Mann (1938), Brown [1945?],
Halstead and Lively (1954), Mills (1956), and Randall (1958). These
workers have attempted to pursue a single train of events which would
explain the occurrence of toxins in all poisonous fishes. Randall (1958)
presented an intriguing ecological hypothesis on the origin of toxins in
the tropical marine environment which partially accounts for the geograph­
ical paradox involved. He considered that toxic blue-green algae are merely
the first organisms growing in normal ecological succession, and that poi­
sonous fishes are found where new substrate is being colonized by such
aleal growth. This hypothesis nicely explains the sometimes sudden appear­
ance of poisonous fishes following anchoring of ships or reef destruction.
More recentlv, Dawson (1959) also arrived at the hypothesis of ecological
succession. While this theory is meritorious in most instances, it is not
adequate to explain the etiology of an area which harbors toxic planktonic,
139
pelagic, as well as benthic, organisms, and hydrographic, chemical, and
meteorological factors must also be considered.
It is not suggested that toxins may be transmitted from algae in only
one way, even though the symptoms produced might be similar. Thus,
poisoning by barracuda might occur from its having eaten poisonous fishes
which had eaten poisonous invertebrates; these could have become poison­
ous from filtering out large quantities of phytoplankton or from browsing
on toxic, benthic algae. Barracuda could also become toxic from ingesting
tetraodontids which had become poisonous as a result of toxins produced
in their gonads during the reproductive cycle.
Brown [1945?] reported upon the distribution of poisonous and non-
poisonous areas in the Windward and Leeward islands of the Caribbean.
He concluded that in most cases the windward or eastern steep, exposed
areas have a reputation for poisonous species and the leeward or shallow-
bank areas are regarded as safe for the same species of fish at the same
time. Watanabe (1946) related that native Okinawans believe that red
snapper, Lutjanus vaigiensis, inhabiting shallow places where the currents
are weak are non-poisonous, while those living in deep localities within
swift currents are poisonous. Gibbard and Naubert (1948) observed that
mussels which had become poisonous from ingesting toxic, planktonic
dinofiagellates in offshore areas were consistently free from poison inshore.
The relation between toxic mussels and hydrography has been reported on
by Waldichuk (1958) and Prakash and Medcof (1962).
All points in the tropical western Atlantic for which information (Table
32) is available designating so-called poisonous and non-poisonous areas
are plotted in Fig. 36. Most refer to the edibility of the barracuda, but each
point may also pertain to other species such as carangids, lutjanids, or
serranids; tetraodontiform fishes are not included. The prevailing ocean
currents, based upon Whiteleather and Brown (1945), and the 100- and
1000-fathom curves are also shown. With few exceptions, localities where
fishes are said to be consistently poisonous are characterized by a steep
gradient along the coastal shelf, and strong, onshore currents. Conversely,
localities where fishes are said never to be poisonous are generally charac­
terized by a relatively gentle-sloping coastal shelf associated with weak
currents. Where fishes are occasionally poisonous, the areas have a steep
coastal shelf gradient but lack the strong, onshore currents, or the converse.
In the Pacific, Randall (1958) observed that poisonous fishes were often
found in areas of seasonal estuarine runoff, which he ascribed to the suc­
cession of euryhaline blue-green algae following the destruction of steno-
haline algae from runoff. In the West Indies, however, there is no indication
that poisonous fishes are ever associated with estuaries, and it appears
that here areas having estuarine runoff never harbor toxic organisms.
Relation of Toxic Organisms to Hydrographic and Meteorological Condi­
tions.—Basically, the problem of toxin production seems to be one of
14 0
 TABLE 32
L o c a l it ie s w here f is h e s are reputedly p o is o n o u s or n o n - p o is o n o u s to eat

Species When poisonous Poisonous locality Non-poisonous locality Authority

All fishes All year Berry Islands, Bahamas — Personal
communication
Barracuda February through Southern Florida Northern and eastern Personal
October and Gulf of Mexico communication
December
(sometimes)
Barracuda Always Havana, Cuba Roig, 1957
Barracuda, Sometimes Bimini, Bahamas Personal
grouper communication
•‘All species but Hormigas Banks, Bahamas Banks, Poey, 1866
groupers” Islandes las Mujeres,
Sanda de Campeche
“All fishes” “Always”? Formigas, Anegada — Rogers, 1899
Barracuda, cero, ? Bahamas (general); Grand Turk Bermuda; Grand Turk Mowbray, 1916
carangids Island: north side more often Island; south side less
often
Barracuda. East and south of St. Thomas, — O’Neill, 1938
Caranx latus, Virgin Islands; Culebrita; north
other caran­ coast of Vieques Island; east of
gids; large king Culebra Island
mackerel
Barracuda “Always” Northern coast of Cuba South coast of Cuba Hoffmann, 1931
Barracuda Never Isle of Pines, Cuba Personal
communication
 TABLE 32— (CONTINUED)

Species When poisonous Poisonous locality Non-poisonous locality Authority

All fishes Sail Rock, St. Thomas, Virgin Islands; Arcisz, 1950
Peter Island, Virgin Islands; from Salinas
Playas to Fajardo, Puerto Rico
Barracuda, cero, Always Virgin Islands: Jost Van Dyke Island; Arcisz, 1950
carangids, waters between St. Thomas and St. John;
lutjanids, west south west of Sail Rock
Lachnolaimus
Barracuda — Yucatan Bean, 1890
Red snapper and Windward grounds and bank around Re- Grenada; St. Lucia; Brown [1945?]
“other fishes” donda; Nevis; Statia; Saba; between St. Dominica, “practically
Kitts and Statia, and to the east and unknown except for
southeast of St. Barts; windward of Mont- barracuda which has a
serrat; east and southeast of the Saba doubtful reputation”;
Bank; Anguilla; seaward of Dog Island; waters close to
Flat Cay and “outside water’’; Ragged Anguilla
Island, Bahamas
Barracuda ? Montserrat; St. Kitts — Brown [1945?]
All fishes Always Islands and cays along Norman Island to — Clarke, 1920
Virgin Gorda, British Virgin Islands; to
the north and northeast of Necker Island;
waters to the southeast at Westend, Tor­
tola; Barbados; Turks Island, Virgin
Islands
Harengula St. Christopher’s (“one end of island”) St. Christopher’s Clarke, 1920
humeralis ( “another end”)
Harengula Occasionally South side of Jamaica North side of Jamaica Clarke, 1920
humeralis
 TABLE 32— (CONTINUED)

Species When poisonous Poisonous locality Non-poisonous locality Authority

Barracuda Ponce, Puerto Rico Clarke, 1920
Carangids “Some seasons” Christ Church Parish, Barbados Dunlop, 1917
Barracuda “Spawning Puerto Rico Costa Mandry, 1933
season”
Barracuda Occasionally Trinidad Dumeril, 1866
(large)
Barracuda Occasionally Near St. Eustatius, Barbados Muller and Troschel,
1847
Barracuda ? Northern end of the St. Martin-St. Bar- Carupano and Puerto Morice, 1958
thelemy islands; Terre-de-Bas, lies de la Cruz, Venezuela
Saintes, Guadeloupe
Barracuda, April Leeward shore of Guadeloupe; in chan- Bay of Basseterre Ferguson, 1823
Opisthonema, nel between St. Kitt and St. Eustatia (Opisthonema); Barba­
large carangids dos (barracuda)
Some carangids Northeast end of Barbados Ferguson, 1823
Barracuda, Smith Bay and Luongo Cay, east and Gilman, 1942
amberjack, southeast of St. Thomas, Virgin Islands,
king mackerel, respectively.
small
“mackerel”
Barracuda Luis Pena Cay, Culebra Island, Virgin Gilman, 1942
Islands
Barracuda Never Bermuda Goode, 1876
Barracuda ? St. Christopher and Nevis Castellani and
Chalmers, 1919
 TABLE 32 (CONTINUED)

Species When poisonous Poisonous locality Non-poisonous locality Authority

Barracuda Occasionally Rio de Janeiro, Brazil — Fonssagrives and
de Mericourt, 1861
All large fishes Always North coast of Cuba and Jamaica; most South coast of Cuba Hoffmann, 1927, 1929
over 3 lbs. other northern coasts in the West Indies and Jamaica; most
other southern coasts
in West Indies
Barracuda Never Cartagena; Islas del Personal
Rosario, Colombia communication
Barracuda Occasionally? “Reefs off Key West are regarded with Jordan, 1885
suspicion.”
Barracuda, porgy, Bahamas Locke, 1675
grouper, king
mackerel,
cowfish, puffer,
Opisthonema
Barracuda Never — Key West, Florida Lonnberg, 1894;
Mowbray, 1916;
Holder, 1924;
Schroeder, 1924
Barracuda Never — Tortugas, Florida Gudger, 1918
Barracuda, king ? North side of Turks Islands, Bahamas South side of Turks Mowbray, 1916
mackerel, three Islands, Bahamas
species of Car-
anxy red rock-
fish, tiger
rockfish
Barracuda Havana and Matanzas, Cuba Nichols, 1912
 TABLE 32— (CONTINUED)

Species When poisonous Poisonous locality Non-poisonous locality Authority

Barracuda May to October Santo Domingo — Pavlovskii, 1927
Sardines ? San Cristobal, Cuba, “one place” San Cristobal, Cuba, Poey, 1866
“another place”
Barracuda Occasionally Kingston, Jamaica — Rogers, 1899
“Grey snapper” ? St. Georges, Grenada; Bridgetown, — Rogers, 1899
Barbados
Scomberomorus ? Barbados — Rogers, 1899
(sp.?)
Barracuda ? — Spanish Honduras Strong, 1942
Barracuda Occasionally British Honduras — Thompson, [1944?]
Barracuda ? West end of Grand Cayman Island “Other parts” Thompson, [1945?]
Lachnolaimus ? Turks and Caicos islands — Thompson, [1945?]
Barracuda, small ? — Cha Cha Town, St. Walker, 1922
carangids Thomas, Virgin Is­
lands; Honduras
Barracuda and Occasionally Martinique — Moreau de
other species Jonnes, 1821
Barracuda Occasionally Fajardo and Humacao, Puerto Rico Aguadilla, Arecibo, Personal
Ponce1, Mayagiiez communication
Anchovies ? North shore of Jamaica South shore of Poey, 1866
Jamaica
Barracuda Always Jabor Island, Jaluit Atoll, Marianas; All other islands of Hiyama, 1943
145

Saipan, Marianas Jaluit Atoll; Japan

 TABLE 32 (CONTINUED)

Species When poisonous Poisonous locality Non-poisonous locality Authority

Barracuda Always Marshall and Marianas islands Caroline Islands Von Fraenkel
and Krick, 1945
Barracuda Occasionally Gilbert and Ellice Islands Castellani and
Chalmers, 1919
Barracuda, Occasionally Raroia Atoll, Tuamotu Archipelago Harry, 1953
some snappers,
groupers,
parrotfish
All species ? Leeward side of Canton Island, Phoenix Halstead and
Islands Bunker, 1954a
“Fishes” ? Leeward side of Hull Island, Phoenix Halstead and
Islands; northwest, leeward side of Sidney Bunker, 1954a
Island
“Fishes” Lagoon of Johnson Island Outside of lagoon Halstead and
Bunker, 1954b
Barracuda Never Hawaii Jordan and
Evermann, 1905
Nearly all species Always Hood Island, near Mangareva Hiroa, 1938
Clupanodon ? New Caledonia Islands to the north of Jouan, 1863b
venenosa New Caledonia
Barracuda New Caledonia — Jouan, 1863b
Barracuda, August to March New Hebrides — Mills, 1956
snappers
 TABLE 32— (CONTINUED)

Species When poisonous Poisonous locality Non-poisonous locality Authority

Snappers and ? New Caledonia; north coast of Australia; Sikiana, Solomon Is­ Mills, 1956
“other species” Fiji; Micronesia; Ellice Island; French lands; Tonga ( “no
Oceania; Philippines and Hawaii; espec­ poisonous fishes”)
ially at Lolowai, Aoba, New Caledonia
Mugil cephalus ? Aoba, Maewo, Espiritu Santo; Maske- “Elsewhere in the Mills, 1956
leynes; Malekula group”
“Reef cod” 9 Aoba, Maewo, New Caledonia Espiritu Santo, Mills, 1956
New Caledonia
“Large, elongate Always Tuamotus — Pellegrin, 1899
fishes”
“Fishes” ? Jaluit Atoll, Marianas Carolines: Palau, Yap, Yasukawa, 1934
Ponape, Truk, Kusaie
“Fishes” Commonly Canala, New Caledonia (on east coast) La Foa, New Caledo­ Loison, 1955
nia (on west coast)
“Fishes” ? Outside the harbor at Saipan Inside the harbor at Yasukawa, 1934
Saipan
“Fishes” Occasionally “A certain section of the reef near the r Randall, 1952
northern part [of Onotoa Atoll, Gilbert
Islands] was poisonous”
Tetraodontids ? Outside reefs of Caroline Islands and Inside reefs of Caro­ Yasukawa, 1934
Tinian line Islands and Tinian
All fishes ? — Nauru and Ocean Yasukawa, 1934
islands
“Fish and “Extremely rare” — Gilbert Islands Catala, 1957
shellfish”
 TABLE 32— (CONTINUED)

Species When poisonous Poisonous locality Non-poisonous locality Authority

Barracuda ? — ?Seychelles Islands Smith, 1956
Barracuda ? lie de Maurice, off Madagascar — Lienard, 1843
Coryphaena ? Madagascar; Madeira Western Atlantic Poey, 1866
Ocean
Sphyraenidae ? — Japan Hiyama, 1943
Barracuda ? — ?Ceylon Malpas, 1926
S. sphyraena Occasionally Free Town, Sierra Leone Mediterranean Irvine, 1947
Barracuda ? Marshall Islands — Smith, 1947
Tetraodon fahaka Never — Upper waters of Niger Daget, 1954
River
Tetraodontiform ? A1 Ghardaqa, Egypt Suez, Egypt Clark and Bohar,
fishes 1953
1A spillage of crude oil near Ponce in July, 1962, was followed by an outbreak of fish poisoning in late 1962 (personal communication).
primary productivity or, more specifically, why certain localities are more
apt to produce toxic algae than are others. Since the toxic blue-green algae
and dinoflagellates are also the dominant phytoplankters in tropical waters
and are the base of the food chain, it is only necessary to determine the
factors limiting abundance of phytoplankton and benthic algae in general,
i.e., light, temperature, nutrients, and plankton spores. The first two would
not appear to be limiting factors in clear, warm tropical seas. Dissolved
nutrients, however, are relatively scarce in the euphotic layers, since tur­
bulence and up welling processes are seldom operative in tropical waters.
In tropical areas where there is a dearth of either nutrients or phytoplank­
ters, the incidence of toxic organisms might be low, whereas poisonous
areas might represent areas of high productivity.
With respect to the possible effects of currents upon an area, it has
already been suggested that the sides of reefs or islands which are exposed
to an onshore current more often have typically poisonous organisms than
do the leeward sides. Similarly, coral reefs flourish far better on the ex­
posed side of an island than they do on the leeward side which may be a
barren waste of sand and rubble (Kuenen, 1950 : 414-479). Poisonous
fishes are seldom found in areas of extensive siltation. Thus, the thesis of
Hiyama (1943), that the distribution of poisonous marine organisms cor­
responds generally to the distribution of reef-building corals, as well as
coralline algae, which flourish along areas exposed to onshore currents or
onshore components of these currents, is reinforced.
With respect to mechanisms which might influence the supply of nu­
trients, physical factors advecting dissolved nutrients to euphotic waters,
such as turbulence and upwelling, are rarely operative in tropical seas;
neither is there any indication from the prevailing currents in relation to
the distribution of poisonous organisms (Fig. 36) that any upwelling exists.
In fact, conditions appear to be ideal in most instances for sinking of water
along the exposed faces of reefs and islands (Sverdrup, et al., 1946: 501,
Fig. 124). Such conditions would be ideal for the operation of the “plank-
ton-trap” mechanism described by Clarke [MS]. He noted that many
phytoplankters are able to adjust their level of flotation, and the swim­
ming activity of the organisms permits them to aggregate at various levels
in the water column. A water mass converging at the shoreline would sink,
yet positively phototactic organisms such as dinoflagellates would tend to
remain in the upper layers of the water resulting in a “plankton trap.” It
would not be necessary for the current to move at right angles. Interaction
with prevailing winds could supply an onshore component, similar to that
depicted by Sverdrup, et al. (1946: 500-501). Local variation in the pro­
ductivity could thus be caused by slight variations in winds, ocean currents,
or bottom topography, with a resulting variation in the toxicity of organ­
isms from one place to another.
In poisonous terrestrial plants, Muenscher (1951: 16-17) noted that
there is a similarity in toxic symptoms produced in animals as a result of
149
their feeding on certain crop plants, grasses, weeds, and algae, and those
produced by feeding the animals potassium nitrate. Plants may store con­
siderable quantities of potassium nitrate in cell sap. Muenscher stated that

3 6 . Distribution of localities in the tropical western Atlantic based

F ig u r e
on literature reports and personal communications where fishes are reputed to
be “poisonous” or “non-poisonous.” Current patterns are based on Whiteleather
and Brown (1945).
150
“the toxic quality of such vegetation is not due to the nitrate directly bui
when the nitrate is ingested it is reduced to nitrite which is toxic.” Muen-
scher also noted that “the accumulation of potassium nitrate varies greatly
in different kinds of plants. The quantity of potassium nitrate may be af­
fected by the amounts of nitrogen and potassium in the soil and the amount,
of light coming in contact with a growing plant. A field may be ‘spotty,5
some areas producing poisonous plants while others may be harmless.”
Even though terrestrial and aquatic situations may not be directly compar­
able, I suggest an analogy between the spotty distribution of poisonous
marine organisms and that of terrestrial plants, with a common underlying
principle. Perhaps such poisonous aquatic areas could be due to a differen­
tial accumulation of nutrients, with the “toxic principle” in marine organ­
isms merely being due initially to various levels of nitrates. Possibly, local
variations in the currents or nutrient supply could favor the growth of
toxic species.
Studies by Thomson, et al. (1957) and Hughes, et al. (1958) have
thrown new light on toxin production in blue-green algae, which suggest
that bacteria present in the polysaccharide sheath of the blue-green algae
may be active in toxin production. These and subsequent studies by Banner,
et al. (1960), which showed that areas of toxic seaweed in Hawaii do
not harbor poisonous fishes, indicate that a species of blue-green algae
may vary spatially and temporally. In light of Muenscher’s (1951) studies,
an analysis of the relationship between hydrographic and chemical factors
to the toxicity of bacteria associated with blue-green algae, as well as coral
and other algae symbionts, might be revealing.
The significance of the steep, exposed face of a reef or island in deter­
mining the presence of poisonous organisms is not clear. Randall (1958)
suggested that continual breaking of reef materials might offer a new sub­
strate. Perhaps the greater depth of the euphotic zone on the exposed side
permits a significantly greater productivity, which is further enhanced by
the reduced turbidity of these deeper strata. Another possibility is that
along steep faces where onshore currents converge, the resultant sinking
of water would be more pronounced in deep water, thus permitting the
formation of a more discrete “plankton trap.” Furthermore, a small but
steady suoply of nutrients and/or phytoplankters would be available to
the benthic organisms growing along the vertical, exposed surface of a
steep reef. The prevailing currents could also act by supplying certain
nutrients such as nitrates or phosphates which are favorable to the pro­
duction of phytoplankton and sessile algae, or reefs with phytoplankton.
On the windward side of islands where coral growth is generally more
luxuriant, a steady supply of plankton is brought onshore by the prevailing
currents. Reef-building corals which contain symbiotic algae, some of
which belong to the toxic dinoflagellate group, utilize the metabolites of
the algae which might be concentrated by the coral polyps. These metabo­
lites could be taken up when the coral was eaten by fishes such as scarids
151
and tetraodontiform fishes, which have already been shown to be poison­
ous and are eaten by barracuda.
Certain localities may become toxic because they were favorable to
accumulations of wind- or current-driven algae or dinoflagellates (Migita
and Kanna, 1957; Waldichuk, 1958), which are eaten by organisms on
which barracuda feed. Freshwater blue-green algae accumulate along
windward shores after a gentle but steady breeze (Mohler, 1941; Vinberg,
1954), and the toxicity of the algae increases as the algae decomposes
(Steyn, 1945).
C o n c l u s io n s

With the above sequence of events in mind, it becomes clear how some
of the paradoxes in the problem of poisonous barracuda fit into the ecolo­
gical scheme. It has been already pointed out that barracuda less than
3 pounds are non-toxic. A plausible explanation for this cryptic figure is
seen in an event which occurs at about the time the barracuda reaches a
weight of 3 pounds. This is a movement from a shallow-water, grass-bed
habitat to the deeper waters of the coral reefs, which is associated with a
change in behavior and food habits. There is a rather abrupt change in the
diet of the barracuda (Table 30) at this size from small fishes such as
atherinids and gobiids to tetraodontiform, carangid, and beloniform fishes.
It is at this time in their life when the barracuda themselves are first
considered poisonous.
This change in diet would explain why only larger barracuda are toxic.
For this same reason, it would explain why sphyraenids which do not
reach a large size are seldom if ever toxic. Thus, S. picudilla and S. gua-
chancho, which seldom exceed 2 feet in length, are commonly eaten
throughout the tropical western Atlantic without ill effects (Guichenot,
1843; Poey, 1858; Nichols, 1912). Other small species such as S. japonica,
S. obtusata, and S. novaehollandiae are eaten throughout the tropical Indo-
Pacific region, and S. sphyraena is of great importance as food in the
Mediterranean Sea where it is eaten without ill effects (Palombi and San-
tarelli, 1953). S. pinguis and S. nigripinnis are prized food fish in Japan,
where poisoning by them is unknown (Hiyama, 1943). None of the above
species attains a large size. If the onset of sexual maturity were the cause
of poisonings, surely some large individuals of the smaller species would
have been incriminated, but apparently this has not occurred. Evidently,
the lack of poisoning by these sphyraenids in part reflects their different
dietary habits, for these non-toxic species feed essentially upon cephalopods
and small atherinids and clupeoids none of which is poisonous except for
the special cases of certain tropical clupeids of the genera Ovisthonema,
Harengula, and Clupanodon. The important California food fish Sphyraena
argentea is eaten safely, and even thoush it reaches a length of 3 V2 feet
and a weight of at least 10 pounds, the lack of poisoning by it seems to
be due to the dearth of poisonous fishes along the California coast, which
152
again would appear to be related to the absence of coral-reef fauna and
flora in the temperate California waters.
The poisonous nature of only large individuals belonging to other
predatory groups, such as the amberjack, king mackerel, lutjanids, and
serranids could be explained in that only the large individuals of these
groups are capable of swallowing large tetraodontiform fishes. Poisoning
in humans by large barracuda is often more severe than when poisoning
results from eating smaller barracuda (Yasukawa, 1934; Smith, 1947);
this is true also for tetraodontids (Remy, 1883; Tani, 1945). This suggests
that the toxin is either cumulative in the fish or that larger predatory fish
are able to ingest larger, and therefore presumably more toxic, prey,
whereas in prey such as the tetraodontiform fishes the amount of toxin
produced may be a function of the size of the reproductive organs.
Another enigma is why barracuda are more often poisonous than are
other predators. It is suggested that the tetraodontiform fishes, which are so
important in the barracuda’s diet, discourage by their protective covering,
or cannot be eaten by, most other predatory species which lack both the
gape and the sharp, shearing teeth of the barracuda. The hard integument
which is developed in the adults of these slow-swimming creatures and
their large size offers little opposition to the barracuda.

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